seco1adary mipests of plants--philippines reproduction and

53
A. PRIMARY I. SUBJECT Agriculture AHTA-0000-G732 CLASSI- SECO1ADARY 1972 MiPests of plants--Philippines (TIsN 2. TITLE AND SUBTITLE Reproduction and seasonal abundance of the ricefield rat (Rattus rattus mindanensis Mearns) at Siniloar,Laguna 3. AUTHORS Marges,1B.1E. OF PAMES ARC NUME 1NUMBER BS. 4. DOCUENT DATE 1972I 52p.I ARC 7. REFERENCE ORGANIZATION NAME AND ADDRESS Interior Publishers, Availability) n 1, SUPPLEMENTARY NOTES (Sponsoring Organization (Thesis M.S.--Univ.of the Philippines) 9 ABSTRACT 11. PRICE OF DOCUMENT 10. CONTROL NUMBER PN-RAA-624 12. DESCRIPTORS 13. PROJECT NUMBER Fecundity Laguna,Philippines? Seasonal variations 14. CONTRACT NUMBER Philippines PASA RACID) I-67 Res. TYPE OF DOCUMENT Rats 1s.

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Page 1: SECO1ADARY MiPests of plants--Philippines Reproduction and

A PRIMARY

I SUBJECT Agriculture AHTA-0000-G732 CLASSI-

SECO1ADARY1972 MiPests of plants--Philippines(TIsN

2 TITLE AND SUBTITLE

Reproduction and seasonal abundance of the ricefield rat (Rattus rattus mindanensis

Mearns) at SiniloarLaguna

3 AUTHORSMarges1B1E

OF PAMES ARC NUME 1NUMBER BS 4 DOCUENT DATE1972I 52pI ARC

7 REFERENCE ORGANIZATION NAME AND ADDRESS

Interior

Publishers Availability)n1 SUPPLEMENTARY NOTES (Sponsoring Organization

(Thesis MS--Univof the Philippines)

9 ABSTRACT

11 PRICE OF DOCUMENT10 CONTROL NUMBER

PN-RAA-624

12 DESCRIPTORS 13 PROJECT NUMBER Fecundity LagunaPhilippines Seasonal variations 14 CONTRACT NUMBER

Philippines PASA RACID) I-67 Res TYPE OF DOCUMENTRats 1s

HEMOUCTIO AND SEASOML ABUNDANCE OF THE RIMML~D MT (RAk1US EATTUS AIMNNIS -REAMS)

AT sInIIAm IAMUA

MRMMABD E 4RCES

SUB4~DTO THE WADUAE FCUIIf OF THE COLIECE OF AMIUIIL UNJMlVRSITY CF TIJE PHLIPPflEs 3NI

BkRTIA FUILILI NqT OF TOE IUIIEITS

LSTE OF SCIENCE (Applied Zooloar)

pRomber -X972

The thesis attached hereto enitled Reproduction and Seasoal

Abundance of the Riceflield Rat -(1Rttusrattus mindanensis Marns) at Siniloan Lagunma prepared and submitted by Bernardo E mirges in

Partial fulfillment of the requirements for the degree of iMster of

Science in Applied Zoology is hereby accepted

Wamber Guidance Comittee Ymber Guidance Comittee

(Date signed) (Date signed)

1vmber Guidance Committee Adviser and chairman Guidance Committee

(Da-te iigued) (Date signed)

Accepted as partial fulnil wn of the requirements for the

degree of aster of Science (Applied Zoology)

FT ORILLO Dean

College of Agriculture University of the Philippines

on (Date)

iernrdo Eifrittu rges WOB -M on August 19- 1937 in Thda6ig

14 thethird sonof Proceso Mrges and Mesta Espitu He took up his- primarycourse at Caiguchi Barrio School at

IfdAngCvjte Hefihisthd his intermediate (with hontors) and

coreesat LunM9nd~i~ n - lmfrbtZJ -School nd -Indan H~ Sdhool respectively IHe attended the University of the pailppines3

College of Agriculturd in 1955 and obtained his Bachelor of Science

in Agriculture in 1959

He joined the Bureau of Plant industry in 1960 Plant Itst Control Officer and mas assiged in Zamboanga del Sur In 1968 he

as treisferred to Davao City Where he assumed supervisory f nztions along withthe Regionis regulatory activities later he Masappointed as Regioal Vegetable ProgramICbordinator for Region No 7 with station

at Zambanga ityi

In 1a9yv as avarded aColombo training grant -o th United Kingdom-ih6r hd ccmpleteafthe Past1inagement coursep He was admitted o he U1A 6afECUSAI graduate fellow later thatyeapr H

qualified as a college scholar kort ie f rst semestet-of the school

year 19704-7 Theb author-k aa to D Escuata j deli ed Thbi o tgos Dao

Sur Their daughters are Ifzel and 1 Cecilia

ERMPDO E YA)GES

ii

ACKN(3IDGEMNTS

I am grateful to nanypeople who helped me throughout the study

Profound gratitude and sincere WpprcIation U expre6ssed to ny major

Prof Pablo J Alfonso for the years of instruction guidance bdviser( i

Thanks areand ndrstanding he bas devqWd to m graduate program

also due to the Bureau of Plant Industry especially to Director Eliseo

C Carandang Messrs Jose I Males Hipolito A Cutodio and Jesus

Po Suangil for their continued encouragement and financial support

I am also grateful to NEC and US-AID for the financial assistance

during nr academic and research enrollment at UPA

Sincere thanks and appreciation are alsoftd to thle Dinector and

members of theRodent Research Center staff for theirassisiaiice and

cooperation during the collection f specimens and in the preparation

of-the manuscript For the geperWs help extend1ed by the teaching

staf of Bkvbay National Agricultural and Vocational School especially

to Messrs Bienvenido RomerosaVnd Roqulo Aloaa for allowing us the

use of their qurters and ohet facilities I am extremely grateful

I wish to thank Wn conmittee DrB Benja~iin L Cariaso and

Vicente G -M0onn and Mr Saztiago M Alviar for their suggestions

and helpful criticisms of the manuscript

In conclusion- I owe a debt of gratitude to zqvife for her

patience understandipg and encpuragement vhen morle s low

---

TABLE OF CONTENTS

Chapter Page

I INTRODUCTION------ --- ----------- -- 1

II TAXONOMY AND DISTRIBUTION- -------------- 4

III DESCRIPTION OF THE STUDY AREA- ----------- - 5

IV MATERIALS AND METHODS 9

A Collection of rats------ ------------ 9

B Autopsy -I----- ---- ---- --- ---- 10

C Statistical analyses - ---- 11i-----

V RESULTS AND DISCUSSION- 1

A Criteria of maturity - ----- ---- ----- 11

1 Testes position size and maturity ------ 14

B -Reproduction -- - - - - - ---- -- 16

1 Breeding season ------------ ----- 16

2 Prevalence of pregnancy ------------- 23

3 Incidence of pregnancy -- -------------- 24

4 Litter size ------ ----- ------- 25

5 Productivity - ------ --- ---- --- 29

6 Juvenile incidence --------- -- ---- 30 C Sex ratio- - shy - - -

D Rat population indices and seasonal fluctuations 36

VI LITEATURE CITED -- 40

iv

LIST OF TALES

Table

i Monthly mean rainfall and average temperature in Siniloan Laguna------------ 8

2 Size of the ricefield rat t various stages of mturity ------ ---- ---- ---- ----- 13

3 Moithly composition of adult male population of Hattus rattus mindanensis with respect to the positionof testes - -------- - ------- 15

4 Monthly changes in the prevalence of pregnant Rattus rattus mindanensis collected in Siniloan August 1971 through Jul 1972 - - - - 22

5 Monthly mean distribution of fetuses in the two horns of the uterus of pregnant rats examined in Siniloan August 1971 through July 1972 ------ 26

6 Analysis of variance for litter size ------- --- 28

7 Mean litter size separation for all pregnant rats examined for this study in Siniloan August 1971 through July 1972 ---- ------- -- ------ 28

8 Productivity of Rattus rattus mindanensis captured during the monthly trapping in Siniloan Auguit C 1971 through July 1972 ------- --- ----- 29

9 Monthly changes in the young-adult ratio of all rats collected for this study in Siniloapi Laguna- - 31

10 Mothly changes in the sex ratio of all rats trapped in Siniloan August 1971 through July 1972 - - - - 33

11 Number of pregnant females aptured in relation to staesof DremancY 35

12 Sex ratios in various weight classes of attus rgttus mindanensis ------------ - --- - 36

13 Population density index of ricefield rats in Siniloan based on monthly trapline catches - ---- 38

v

LIST OF FIGURES

Page

1 Portion ofthebull qtudy area showing condition of the rice paddies after harvest- ------------- 6

2 Scatter diagram of length of testis against weight of rats showing presence or absence of sperm 17

3- Graph showing extent of breeding season Judged by percent pregnant and immature ----------- 1

4 Monthly changes in the prevalence of pregnancy in -relation to total monthly rainfall and number of rainy days per month -- ---- ---- -- ---- 21

5 Graph showing relationship between breeding and monthly sex ratio - ------------------- 34

6 Monthly trapline catches of field rats in relation to rainfall and temperature -------- -- 39

ABSTACT

Mrges Bernardo Espiritu University of the Tilippines

November 1972 Reproduction and Seasonal Abundance of the Ricefield

it (lMttus rattus mindanensis Yearns) at Siniloan laguna

ljor Professor blo J Alfonso

The reproductive patterns and seasonal abundance of the common

ricefield rat were studied in an irrigated ricefield in Siniloan

IAguna from August 1971 through July 1972 The study area ias

planted to two crops of rice every year Of the two rat species

found the ricefield rat dominated the entire riceland comunity

comprising more than 99 percent of the total collection

Two age classes were recognized Juveniles and adults Adults

were further subdivided into breeding and non-breeding The vaginal

orifice became perforate at 5445 grams among fenales while the

testes descended to the scrotum at 13493 grams among males For

both sexes the skull ossified at 9079 grams The length of the testes

vas significantly correlated with the body weight Active sperms

were present in the cauda epididymis when the testes reached lengtha

of 2229 mm Once sexual maturity was attained males remained fertile

throughout the year

Reproductive pattern was sarply bimodal corresponding very

closely with the crop cycle Most litters were produced during the

reproductive and ripening stages of rice plants with little or no

breeding activity when the fields lie fallow There was no direct

relationship between breeding and seasonal or climtic chages Mean

vii

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

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41

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43

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TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 2: SECO1ADARY MiPests of plants--Philippines Reproduction and

HEMOUCTIO AND SEASOML ABUNDANCE OF THE RIMML~D MT (RAk1US EATTUS AIMNNIS -REAMS)

AT sInIIAm IAMUA

MRMMABD E 4RCES

SUB4~DTO THE WADUAE FCUIIf OF THE COLIECE OF AMIUIIL UNJMlVRSITY CF TIJE PHLIPPflEs 3NI

BkRTIA FUILILI NqT OF TOE IUIIEITS

LSTE OF SCIENCE (Applied Zooloar)

pRomber -X972

The thesis attached hereto enitled Reproduction and Seasoal

Abundance of the Riceflield Rat -(1Rttusrattus mindanensis Marns) at Siniloan Lagunma prepared and submitted by Bernardo E mirges in

Partial fulfillment of the requirements for the degree of iMster of

Science in Applied Zoology is hereby accepted

Wamber Guidance Comittee Ymber Guidance Comittee

(Date signed) (Date signed)

1vmber Guidance Committee Adviser and chairman Guidance Committee

(Da-te iigued) (Date signed)

Accepted as partial fulnil wn of the requirements for the

degree of aster of Science (Applied Zoology)

FT ORILLO Dean

College of Agriculture University of the Philippines

on (Date)

iernrdo Eifrittu rges WOB -M on August 19- 1937 in Thda6ig

14 thethird sonof Proceso Mrges and Mesta Espitu He took up his- primarycourse at Caiguchi Barrio School at

IfdAngCvjte Hefihisthd his intermediate (with hontors) and

coreesat LunM9nd~i~ n - lmfrbtZJ -School nd -Indan H~ Sdhool respectively IHe attended the University of the pailppines3

College of Agriculturd in 1955 and obtained his Bachelor of Science

in Agriculture in 1959

He joined the Bureau of Plant industry in 1960 Plant Itst Control Officer and mas assiged in Zamboanga del Sur In 1968 he

as treisferred to Davao City Where he assumed supervisory f nztions along withthe Regionis regulatory activities later he Masappointed as Regioal Vegetable ProgramICbordinator for Region No 7 with station

at Zambanga ityi

In 1a9yv as avarded aColombo training grant -o th United Kingdom-ih6r hd ccmpleteafthe Past1inagement coursep He was admitted o he U1A 6afECUSAI graduate fellow later thatyeapr H

qualified as a college scholar kort ie f rst semestet-of the school

year 19704-7 Theb author-k aa to D Escuata j deli ed Thbi o tgos Dao

Sur Their daughters are Ifzel and 1 Cecilia

ERMPDO E YA)GES

ii

ACKN(3IDGEMNTS

I am grateful to nanypeople who helped me throughout the study

Profound gratitude and sincere WpprcIation U expre6ssed to ny major

Prof Pablo J Alfonso for the years of instruction guidance bdviser( i

Thanks areand ndrstanding he bas devqWd to m graduate program

also due to the Bureau of Plant Industry especially to Director Eliseo

C Carandang Messrs Jose I Males Hipolito A Cutodio and Jesus

Po Suangil for their continued encouragement and financial support

I am also grateful to NEC and US-AID for the financial assistance

during nr academic and research enrollment at UPA

Sincere thanks and appreciation are alsoftd to thle Dinector and

members of theRodent Research Center staff for theirassisiaiice and

cooperation during the collection f specimens and in the preparation

of-the manuscript For the geperWs help extend1ed by the teaching

staf of Bkvbay National Agricultural and Vocational School especially

to Messrs Bienvenido RomerosaVnd Roqulo Aloaa for allowing us the

use of their qurters and ohet facilities I am extremely grateful

I wish to thank Wn conmittee DrB Benja~iin L Cariaso and

Vicente G -M0onn and Mr Saztiago M Alviar for their suggestions

and helpful criticisms of the manuscript

In conclusion- I owe a debt of gratitude to zqvife for her

patience understandipg and encpuragement vhen morle s low

---

TABLE OF CONTENTS

Chapter Page

I INTRODUCTION------ --- ----------- -- 1

II TAXONOMY AND DISTRIBUTION- -------------- 4

III DESCRIPTION OF THE STUDY AREA- ----------- - 5

IV MATERIALS AND METHODS 9

A Collection of rats------ ------------ 9

B Autopsy -I----- ---- ---- --- ---- 10

C Statistical analyses - ---- 11i-----

V RESULTS AND DISCUSSION- 1

A Criteria of maturity - ----- ---- ----- 11

1 Testes position size and maturity ------ 14

B -Reproduction -- - - - - - ---- -- 16

1 Breeding season ------------ ----- 16

2 Prevalence of pregnancy ------------- 23

3 Incidence of pregnancy -- -------------- 24

4 Litter size ------ ----- ------- 25

5 Productivity - ------ --- ---- --- 29

6 Juvenile incidence --------- -- ---- 30 C Sex ratio- - shy - - -

D Rat population indices and seasonal fluctuations 36

VI LITEATURE CITED -- 40

iv

LIST OF TALES

Table

i Monthly mean rainfall and average temperature in Siniloan Laguna------------ 8

2 Size of the ricefield rat t various stages of mturity ------ ---- ---- ---- ----- 13

3 Moithly composition of adult male population of Hattus rattus mindanensis with respect to the positionof testes - -------- - ------- 15

4 Monthly changes in the prevalence of pregnant Rattus rattus mindanensis collected in Siniloan August 1971 through Jul 1972 - - - - 22

5 Monthly mean distribution of fetuses in the two horns of the uterus of pregnant rats examined in Siniloan August 1971 through July 1972 ------ 26

6 Analysis of variance for litter size ------- --- 28

7 Mean litter size separation for all pregnant rats examined for this study in Siniloan August 1971 through July 1972 ---- ------- -- ------ 28

8 Productivity of Rattus rattus mindanensis captured during the monthly trapping in Siniloan Auguit C 1971 through July 1972 ------- --- ----- 29

9 Monthly changes in the young-adult ratio of all rats collected for this study in Siniloapi Laguna- - 31

10 Mothly changes in the sex ratio of all rats trapped in Siniloan August 1971 through July 1972 - - - - 33

11 Number of pregnant females aptured in relation to staesof DremancY 35

12 Sex ratios in various weight classes of attus rgttus mindanensis ------------ - --- - 36

13 Population density index of ricefield rats in Siniloan based on monthly trapline catches - ---- 38

v

LIST OF FIGURES

Page

1 Portion ofthebull qtudy area showing condition of the rice paddies after harvest- ------------- 6

2 Scatter diagram of length of testis against weight of rats showing presence or absence of sperm 17

3- Graph showing extent of breeding season Judged by percent pregnant and immature ----------- 1

4 Monthly changes in the prevalence of pregnancy in -relation to total monthly rainfall and number of rainy days per month -- ---- ---- -- ---- 21

5 Graph showing relationship between breeding and monthly sex ratio - ------------------- 34

6 Monthly trapline catches of field rats in relation to rainfall and temperature -------- -- 39

ABSTACT

Mrges Bernardo Espiritu University of the Tilippines

November 1972 Reproduction and Seasonal Abundance of the Ricefield

it (lMttus rattus mindanensis Yearns) at Siniloan laguna

ljor Professor blo J Alfonso

The reproductive patterns and seasonal abundance of the common

ricefield rat were studied in an irrigated ricefield in Siniloan

IAguna from August 1971 through July 1972 The study area ias

planted to two crops of rice every year Of the two rat species

found the ricefield rat dominated the entire riceland comunity

comprising more than 99 percent of the total collection

Two age classes were recognized Juveniles and adults Adults

were further subdivided into breeding and non-breeding The vaginal

orifice became perforate at 5445 grams among fenales while the

testes descended to the scrotum at 13493 grams among males For

both sexes the skull ossified at 9079 grams The length of the testes

vas significantly correlated with the body weight Active sperms

were present in the cauda epididymis when the testes reached lengtha

of 2229 mm Once sexual maturity was attained males remained fertile

throughout the year

Reproductive pattern was sarply bimodal corresponding very

closely with the crop cycle Most litters were produced during the

reproductive and ripening stages of rice plants with little or no

breeding activity when the fields lie fallow There was no direct

relationship between breeding and seasonal or climtic chages Mean

vii

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

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and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

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NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

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SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

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0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

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UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 3: SECO1ADARY MiPests of plants--Philippines Reproduction and

The thesis attached hereto enitled Reproduction and Seasoal

Abundance of the Riceflield Rat -(1Rttusrattus mindanensis Marns) at Siniloan Lagunma prepared and submitted by Bernardo E mirges in

Partial fulfillment of the requirements for the degree of iMster of

Science in Applied Zoology is hereby accepted

Wamber Guidance Comittee Ymber Guidance Comittee

(Date signed) (Date signed)

1vmber Guidance Committee Adviser and chairman Guidance Committee

(Da-te iigued) (Date signed)

Accepted as partial fulnil wn of the requirements for the

degree of aster of Science (Applied Zoology)

FT ORILLO Dean

College of Agriculture University of the Philippines

on (Date)

iernrdo Eifrittu rges WOB -M on August 19- 1937 in Thda6ig

14 thethird sonof Proceso Mrges and Mesta Espitu He took up his- primarycourse at Caiguchi Barrio School at

IfdAngCvjte Hefihisthd his intermediate (with hontors) and

coreesat LunM9nd~i~ n - lmfrbtZJ -School nd -Indan H~ Sdhool respectively IHe attended the University of the pailppines3

College of Agriculturd in 1955 and obtained his Bachelor of Science

in Agriculture in 1959

He joined the Bureau of Plant industry in 1960 Plant Itst Control Officer and mas assiged in Zamboanga del Sur In 1968 he

as treisferred to Davao City Where he assumed supervisory f nztions along withthe Regionis regulatory activities later he Masappointed as Regioal Vegetable ProgramICbordinator for Region No 7 with station

at Zambanga ityi

In 1a9yv as avarded aColombo training grant -o th United Kingdom-ih6r hd ccmpleteafthe Past1inagement coursep He was admitted o he U1A 6afECUSAI graduate fellow later thatyeapr H

qualified as a college scholar kort ie f rst semestet-of the school

year 19704-7 Theb author-k aa to D Escuata j deli ed Thbi o tgos Dao

Sur Their daughters are Ifzel and 1 Cecilia

ERMPDO E YA)GES

ii

ACKN(3IDGEMNTS

I am grateful to nanypeople who helped me throughout the study

Profound gratitude and sincere WpprcIation U expre6ssed to ny major

Prof Pablo J Alfonso for the years of instruction guidance bdviser( i

Thanks areand ndrstanding he bas devqWd to m graduate program

also due to the Bureau of Plant Industry especially to Director Eliseo

C Carandang Messrs Jose I Males Hipolito A Cutodio and Jesus

Po Suangil for their continued encouragement and financial support

I am also grateful to NEC and US-AID for the financial assistance

during nr academic and research enrollment at UPA

Sincere thanks and appreciation are alsoftd to thle Dinector and

members of theRodent Research Center staff for theirassisiaiice and

cooperation during the collection f specimens and in the preparation

of-the manuscript For the geperWs help extend1ed by the teaching

staf of Bkvbay National Agricultural and Vocational School especially

to Messrs Bienvenido RomerosaVnd Roqulo Aloaa for allowing us the

use of their qurters and ohet facilities I am extremely grateful

I wish to thank Wn conmittee DrB Benja~iin L Cariaso and

Vicente G -M0onn and Mr Saztiago M Alviar for their suggestions

and helpful criticisms of the manuscript

In conclusion- I owe a debt of gratitude to zqvife for her

patience understandipg and encpuragement vhen morle s low

---

TABLE OF CONTENTS

Chapter Page

I INTRODUCTION------ --- ----------- -- 1

II TAXONOMY AND DISTRIBUTION- -------------- 4

III DESCRIPTION OF THE STUDY AREA- ----------- - 5

IV MATERIALS AND METHODS 9

A Collection of rats------ ------------ 9

B Autopsy -I----- ---- ---- --- ---- 10

C Statistical analyses - ---- 11i-----

V RESULTS AND DISCUSSION- 1

A Criteria of maturity - ----- ---- ----- 11

1 Testes position size and maturity ------ 14

B -Reproduction -- - - - - - ---- -- 16

1 Breeding season ------------ ----- 16

2 Prevalence of pregnancy ------------- 23

3 Incidence of pregnancy -- -------------- 24

4 Litter size ------ ----- ------- 25

5 Productivity - ------ --- ---- --- 29

6 Juvenile incidence --------- -- ---- 30 C Sex ratio- - shy - - -

D Rat population indices and seasonal fluctuations 36

VI LITEATURE CITED -- 40

iv

LIST OF TALES

Table

i Monthly mean rainfall and average temperature in Siniloan Laguna------------ 8

2 Size of the ricefield rat t various stages of mturity ------ ---- ---- ---- ----- 13

3 Moithly composition of adult male population of Hattus rattus mindanensis with respect to the positionof testes - -------- - ------- 15

4 Monthly changes in the prevalence of pregnant Rattus rattus mindanensis collected in Siniloan August 1971 through Jul 1972 - - - - 22

5 Monthly mean distribution of fetuses in the two horns of the uterus of pregnant rats examined in Siniloan August 1971 through July 1972 ------ 26

6 Analysis of variance for litter size ------- --- 28

7 Mean litter size separation for all pregnant rats examined for this study in Siniloan August 1971 through July 1972 ---- ------- -- ------ 28

8 Productivity of Rattus rattus mindanensis captured during the monthly trapping in Siniloan Auguit C 1971 through July 1972 ------- --- ----- 29

9 Monthly changes in the young-adult ratio of all rats collected for this study in Siniloapi Laguna- - 31

10 Mothly changes in the sex ratio of all rats trapped in Siniloan August 1971 through July 1972 - - - - 33

11 Number of pregnant females aptured in relation to staesof DremancY 35

12 Sex ratios in various weight classes of attus rgttus mindanensis ------------ - --- - 36

13 Population density index of ricefield rats in Siniloan based on monthly trapline catches - ---- 38

v

LIST OF FIGURES

Page

1 Portion ofthebull qtudy area showing condition of the rice paddies after harvest- ------------- 6

2 Scatter diagram of length of testis against weight of rats showing presence or absence of sperm 17

3- Graph showing extent of breeding season Judged by percent pregnant and immature ----------- 1

4 Monthly changes in the prevalence of pregnancy in -relation to total monthly rainfall and number of rainy days per month -- ---- ---- -- ---- 21

5 Graph showing relationship between breeding and monthly sex ratio - ------------------- 34

6 Monthly trapline catches of field rats in relation to rainfall and temperature -------- -- 39

ABSTACT

Mrges Bernardo Espiritu University of the Tilippines

November 1972 Reproduction and Seasonal Abundance of the Ricefield

it (lMttus rattus mindanensis Yearns) at Siniloan laguna

ljor Professor blo J Alfonso

The reproductive patterns and seasonal abundance of the common

ricefield rat were studied in an irrigated ricefield in Siniloan

IAguna from August 1971 through July 1972 The study area ias

planted to two crops of rice every year Of the two rat species

found the ricefield rat dominated the entire riceland comunity

comprising more than 99 percent of the total collection

Two age classes were recognized Juveniles and adults Adults

were further subdivided into breeding and non-breeding The vaginal

orifice became perforate at 5445 grams among fenales while the

testes descended to the scrotum at 13493 grams among males For

both sexes the skull ossified at 9079 grams The length of the testes

vas significantly correlated with the body weight Active sperms

were present in the cauda epididymis when the testes reached lengtha

of 2229 mm Once sexual maturity was attained males remained fertile

throughout the year

Reproductive pattern was sarply bimodal corresponding very

closely with the crop cycle Most litters were produced during the

reproductive and ripening stages of rice plants with little or no

breeding activity when the fields lie fallow There was no direct

relationship between breeding and seasonal or climtic chages Mean

vii

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 4: SECO1ADARY MiPests of plants--Philippines Reproduction and

iernrdo Eifrittu rges WOB -M on August 19- 1937 in Thda6ig

14 thethird sonof Proceso Mrges and Mesta Espitu He took up his- primarycourse at Caiguchi Barrio School at

IfdAngCvjte Hefihisthd his intermediate (with hontors) and

coreesat LunM9nd~i~ n - lmfrbtZJ -School nd -Indan H~ Sdhool respectively IHe attended the University of the pailppines3

College of Agriculturd in 1955 and obtained his Bachelor of Science

in Agriculture in 1959

He joined the Bureau of Plant industry in 1960 Plant Itst Control Officer and mas assiged in Zamboanga del Sur In 1968 he

as treisferred to Davao City Where he assumed supervisory f nztions along withthe Regionis regulatory activities later he Masappointed as Regioal Vegetable ProgramICbordinator for Region No 7 with station

at Zambanga ityi

In 1a9yv as avarded aColombo training grant -o th United Kingdom-ih6r hd ccmpleteafthe Past1inagement coursep He was admitted o he U1A 6afECUSAI graduate fellow later thatyeapr H

qualified as a college scholar kort ie f rst semestet-of the school

year 19704-7 Theb author-k aa to D Escuata j deli ed Thbi o tgos Dao

Sur Their daughters are Ifzel and 1 Cecilia

ERMPDO E YA)GES

ii

ACKN(3IDGEMNTS

I am grateful to nanypeople who helped me throughout the study

Profound gratitude and sincere WpprcIation U expre6ssed to ny major

Prof Pablo J Alfonso for the years of instruction guidance bdviser( i

Thanks areand ndrstanding he bas devqWd to m graduate program

also due to the Bureau of Plant Industry especially to Director Eliseo

C Carandang Messrs Jose I Males Hipolito A Cutodio and Jesus

Po Suangil for their continued encouragement and financial support

I am also grateful to NEC and US-AID for the financial assistance

during nr academic and research enrollment at UPA

Sincere thanks and appreciation are alsoftd to thle Dinector and

members of theRodent Research Center staff for theirassisiaiice and

cooperation during the collection f specimens and in the preparation

of-the manuscript For the geperWs help extend1ed by the teaching

staf of Bkvbay National Agricultural and Vocational School especially

to Messrs Bienvenido RomerosaVnd Roqulo Aloaa for allowing us the

use of their qurters and ohet facilities I am extremely grateful

I wish to thank Wn conmittee DrB Benja~iin L Cariaso and

Vicente G -M0onn and Mr Saztiago M Alviar for their suggestions

and helpful criticisms of the manuscript

In conclusion- I owe a debt of gratitude to zqvife for her

patience understandipg and encpuragement vhen morle s low

---

TABLE OF CONTENTS

Chapter Page

I INTRODUCTION------ --- ----------- -- 1

II TAXONOMY AND DISTRIBUTION- -------------- 4

III DESCRIPTION OF THE STUDY AREA- ----------- - 5

IV MATERIALS AND METHODS 9

A Collection of rats------ ------------ 9

B Autopsy -I----- ---- ---- --- ---- 10

C Statistical analyses - ---- 11i-----

V RESULTS AND DISCUSSION- 1

A Criteria of maturity - ----- ---- ----- 11

1 Testes position size and maturity ------ 14

B -Reproduction -- - - - - - ---- -- 16

1 Breeding season ------------ ----- 16

2 Prevalence of pregnancy ------------- 23

3 Incidence of pregnancy -- -------------- 24

4 Litter size ------ ----- ------- 25

5 Productivity - ------ --- ---- --- 29

6 Juvenile incidence --------- -- ---- 30 C Sex ratio- - shy - - -

D Rat population indices and seasonal fluctuations 36

VI LITEATURE CITED -- 40

iv

LIST OF TALES

Table

i Monthly mean rainfall and average temperature in Siniloan Laguna------------ 8

2 Size of the ricefield rat t various stages of mturity ------ ---- ---- ---- ----- 13

3 Moithly composition of adult male population of Hattus rattus mindanensis with respect to the positionof testes - -------- - ------- 15

4 Monthly changes in the prevalence of pregnant Rattus rattus mindanensis collected in Siniloan August 1971 through Jul 1972 - - - - 22

5 Monthly mean distribution of fetuses in the two horns of the uterus of pregnant rats examined in Siniloan August 1971 through July 1972 ------ 26

6 Analysis of variance for litter size ------- --- 28

7 Mean litter size separation for all pregnant rats examined for this study in Siniloan August 1971 through July 1972 ---- ------- -- ------ 28

8 Productivity of Rattus rattus mindanensis captured during the monthly trapping in Siniloan Auguit C 1971 through July 1972 ------- --- ----- 29

9 Monthly changes in the young-adult ratio of all rats collected for this study in Siniloapi Laguna- - 31

10 Mothly changes in the sex ratio of all rats trapped in Siniloan August 1971 through July 1972 - - - - 33

11 Number of pregnant females aptured in relation to staesof DremancY 35

12 Sex ratios in various weight classes of attus rgttus mindanensis ------------ - --- - 36

13 Population density index of ricefield rats in Siniloan based on monthly trapline catches - ---- 38

v

LIST OF FIGURES

Page

1 Portion ofthebull qtudy area showing condition of the rice paddies after harvest- ------------- 6

2 Scatter diagram of length of testis against weight of rats showing presence or absence of sperm 17

3- Graph showing extent of breeding season Judged by percent pregnant and immature ----------- 1

4 Monthly changes in the prevalence of pregnancy in -relation to total monthly rainfall and number of rainy days per month -- ---- ---- -- ---- 21

5 Graph showing relationship between breeding and monthly sex ratio - ------------------- 34

6 Monthly trapline catches of field rats in relation to rainfall and temperature -------- -- 39

ABSTACT

Mrges Bernardo Espiritu University of the Tilippines

November 1972 Reproduction and Seasonal Abundance of the Ricefield

it (lMttus rattus mindanensis Yearns) at Siniloan laguna

ljor Professor blo J Alfonso

The reproductive patterns and seasonal abundance of the common

ricefield rat were studied in an irrigated ricefield in Siniloan

IAguna from August 1971 through July 1972 The study area ias

planted to two crops of rice every year Of the two rat species

found the ricefield rat dominated the entire riceland comunity

comprising more than 99 percent of the total collection

Two age classes were recognized Juveniles and adults Adults

were further subdivided into breeding and non-breeding The vaginal

orifice became perforate at 5445 grams among fenales while the

testes descended to the scrotum at 13493 grams among males For

both sexes the skull ossified at 9079 grams The length of the testes

vas significantly correlated with the body weight Active sperms

were present in the cauda epididymis when the testes reached lengtha

of 2229 mm Once sexual maturity was attained males remained fertile

throughout the year

Reproductive pattern was sarply bimodal corresponding very

closely with the crop cycle Most litters were produced during the

reproductive and ripening stages of rice plants with little or no

breeding activity when the fields lie fallow There was no direct

relationship between breeding and seasonal or climtic chages Mean

vii

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 5: SECO1ADARY MiPests of plants--Philippines Reproduction and

ACKN(3IDGEMNTS

I am grateful to nanypeople who helped me throughout the study

Profound gratitude and sincere WpprcIation U expre6ssed to ny major

Prof Pablo J Alfonso for the years of instruction guidance bdviser( i

Thanks areand ndrstanding he bas devqWd to m graduate program

also due to the Bureau of Plant Industry especially to Director Eliseo

C Carandang Messrs Jose I Males Hipolito A Cutodio and Jesus

Po Suangil for their continued encouragement and financial support

I am also grateful to NEC and US-AID for the financial assistance

during nr academic and research enrollment at UPA

Sincere thanks and appreciation are alsoftd to thle Dinector and

members of theRodent Research Center staff for theirassisiaiice and

cooperation during the collection f specimens and in the preparation

of-the manuscript For the geperWs help extend1ed by the teaching

staf of Bkvbay National Agricultural and Vocational School especially

to Messrs Bienvenido RomerosaVnd Roqulo Aloaa for allowing us the

use of their qurters and ohet facilities I am extremely grateful

I wish to thank Wn conmittee DrB Benja~iin L Cariaso and

Vicente G -M0onn and Mr Saztiago M Alviar for their suggestions

and helpful criticisms of the manuscript

In conclusion- I owe a debt of gratitude to zqvife for her

patience understandipg and encpuragement vhen morle s low

---

TABLE OF CONTENTS

Chapter Page

I INTRODUCTION------ --- ----------- -- 1

II TAXONOMY AND DISTRIBUTION- -------------- 4

III DESCRIPTION OF THE STUDY AREA- ----------- - 5

IV MATERIALS AND METHODS 9

A Collection of rats------ ------------ 9

B Autopsy -I----- ---- ---- --- ---- 10

C Statistical analyses - ---- 11i-----

V RESULTS AND DISCUSSION- 1

A Criteria of maturity - ----- ---- ----- 11

1 Testes position size and maturity ------ 14

B -Reproduction -- - - - - - ---- -- 16

1 Breeding season ------------ ----- 16

2 Prevalence of pregnancy ------------- 23

3 Incidence of pregnancy -- -------------- 24

4 Litter size ------ ----- ------- 25

5 Productivity - ------ --- ---- --- 29

6 Juvenile incidence --------- -- ---- 30 C Sex ratio- - shy - - -

D Rat population indices and seasonal fluctuations 36

VI LITEATURE CITED -- 40

iv

LIST OF TALES

Table

i Monthly mean rainfall and average temperature in Siniloan Laguna------------ 8

2 Size of the ricefield rat t various stages of mturity ------ ---- ---- ---- ----- 13

3 Moithly composition of adult male population of Hattus rattus mindanensis with respect to the positionof testes - -------- - ------- 15

4 Monthly changes in the prevalence of pregnant Rattus rattus mindanensis collected in Siniloan August 1971 through Jul 1972 - - - - 22

5 Monthly mean distribution of fetuses in the two horns of the uterus of pregnant rats examined in Siniloan August 1971 through July 1972 ------ 26

6 Analysis of variance for litter size ------- --- 28

7 Mean litter size separation for all pregnant rats examined for this study in Siniloan August 1971 through July 1972 ---- ------- -- ------ 28

8 Productivity of Rattus rattus mindanensis captured during the monthly trapping in Siniloan Auguit C 1971 through July 1972 ------- --- ----- 29

9 Monthly changes in the young-adult ratio of all rats collected for this study in Siniloapi Laguna- - 31

10 Mothly changes in the sex ratio of all rats trapped in Siniloan August 1971 through July 1972 - - - - 33

11 Number of pregnant females aptured in relation to staesof DremancY 35

12 Sex ratios in various weight classes of attus rgttus mindanensis ------------ - --- - 36

13 Population density index of ricefield rats in Siniloan based on monthly trapline catches - ---- 38

v

LIST OF FIGURES

Page

1 Portion ofthebull qtudy area showing condition of the rice paddies after harvest- ------------- 6

2 Scatter diagram of length of testis against weight of rats showing presence or absence of sperm 17

3- Graph showing extent of breeding season Judged by percent pregnant and immature ----------- 1

4 Monthly changes in the prevalence of pregnancy in -relation to total monthly rainfall and number of rainy days per month -- ---- ---- -- ---- 21

5 Graph showing relationship between breeding and monthly sex ratio - ------------------- 34

6 Monthly trapline catches of field rats in relation to rainfall and temperature -------- -- 39

ABSTACT

Mrges Bernardo Espiritu University of the Tilippines

November 1972 Reproduction and Seasonal Abundance of the Ricefield

it (lMttus rattus mindanensis Yearns) at Siniloan laguna

ljor Professor blo J Alfonso

The reproductive patterns and seasonal abundance of the common

ricefield rat were studied in an irrigated ricefield in Siniloan

IAguna from August 1971 through July 1972 The study area ias

planted to two crops of rice every year Of the two rat species

found the ricefield rat dominated the entire riceland comunity

comprising more than 99 percent of the total collection

Two age classes were recognized Juveniles and adults Adults

were further subdivided into breeding and non-breeding The vaginal

orifice became perforate at 5445 grams among fenales while the

testes descended to the scrotum at 13493 grams among males For

both sexes the skull ossified at 9079 grams The length of the testes

vas significantly correlated with the body weight Active sperms

were present in the cauda epididymis when the testes reached lengtha

of 2229 mm Once sexual maturity was attained males remained fertile

throughout the year

Reproductive pattern was sarply bimodal corresponding very

closely with the crop cycle Most litters were produced during the

reproductive and ripening stages of rice plants with little or no

breeding activity when the fields lie fallow There was no direct

relationship between breeding and seasonal or climtic chages Mean

vii

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 6: SECO1ADARY MiPests of plants--Philippines Reproduction and

---

TABLE OF CONTENTS

Chapter Page

I INTRODUCTION------ --- ----------- -- 1

II TAXONOMY AND DISTRIBUTION- -------------- 4

III DESCRIPTION OF THE STUDY AREA- ----------- - 5

IV MATERIALS AND METHODS 9

A Collection of rats------ ------------ 9

B Autopsy -I----- ---- ---- --- ---- 10

C Statistical analyses - ---- 11i-----

V RESULTS AND DISCUSSION- 1

A Criteria of maturity - ----- ---- ----- 11

1 Testes position size and maturity ------ 14

B -Reproduction -- - - - - - ---- -- 16

1 Breeding season ------------ ----- 16

2 Prevalence of pregnancy ------------- 23

3 Incidence of pregnancy -- -------------- 24

4 Litter size ------ ----- ------- 25

5 Productivity - ------ --- ---- --- 29

6 Juvenile incidence --------- -- ---- 30 C Sex ratio- - shy - - -

D Rat population indices and seasonal fluctuations 36

VI LITEATURE CITED -- 40

iv

LIST OF TALES

Table

i Monthly mean rainfall and average temperature in Siniloan Laguna------------ 8

2 Size of the ricefield rat t various stages of mturity ------ ---- ---- ---- ----- 13

3 Moithly composition of adult male population of Hattus rattus mindanensis with respect to the positionof testes - -------- - ------- 15

4 Monthly changes in the prevalence of pregnant Rattus rattus mindanensis collected in Siniloan August 1971 through Jul 1972 - - - - 22

5 Monthly mean distribution of fetuses in the two horns of the uterus of pregnant rats examined in Siniloan August 1971 through July 1972 ------ 26

6 Analysis of variance for litter size ------- --- 28

7 Mean litter size separation for all pregnant rats examined for this study in Siniloan August 1971 through July 1972 ---- ------- -- ------ 28

8 Productivity of Rattus rattus mindanensis captured during the monthly trapping in Siniloan Auguit C 1971 through July 1972 ------- --- ----- 29

9 Monthly changes in the young-adult ratio of all rats collected for this study in Siniloapi Laguna- - 31

10 Mothly changes in the sex ratio of all rats trapped in Siniloan August 1971 through July 1972 - - - - 33

11 Number of pregnant females aptured in relation to staesof DremancY 35

12 Sex ratios in various weight classes of attus rgttus mindanensis ------------ - --- - 36

13 Population density index of ricefield rats in Siniloan based on monthly trapline catches - ---- 38

v

LIST OF FIGURES

Page

1 Portion ofthebull qtudy area showing condition of the rice paddies after harvest- ------------- 6

2 Scatter diagram of length of testis against weight of rats showing presence or absence of sperm 17

3- Graph showing extent of breeding season Judged by percent pregnant and immature ----------- 1

4 Monthly changes in the prevalence of pregnancy in -relation to total monthly rainfall and number of rainy days per month -- ---- ---- -- ---- 21

5 Graph showing relationship between breeding and monthly sex ratio - ------------------- 34

6 Monthly trapline catches of field rats in relation to rainfall and temperature -------- -- 39

ABSTACT

Mrges Bernardo Espiritu University of the Tilippines

November 1972 Reproduction and Seasonal Abundance of the Ricefield

it (lMttus rattus mindanensis Yearns) at Siniloan laguna

ljor Professor blo J Alfonso

The reproductive patterns and seasonal abundance of the common

ricefield rat were studied in an irrigated ricefield in Siniloan

IAguna from August 1971 through July 1972 The study area ias

planted to two crops of rice every year Of the two rat species

found the ricefield rat dominated the entire riceland comunity

comprising more than 99 percent of the total collection

Two age classes were recognized Juveniles and adults Adults

were further subdivided into breeding and non-breeding The vaginal

orifice became perforate at 5445 grams among fenales while the

testes descended to the scrotum at 13493 grams among males For

both sexes the skull ossified at 9079 grams The length of the testes

vas significantly correlated with the body weight Active sperms

were present in the cauda epididymis when the testes reached lengtha

of 2229 mm Once sexual maturity was attained males remained fertile

throughout the year

Reproductive pattern was sarply bimodal corresponding very

closely with the crop cycle Most litters were produced during the

reproductive and ripening stages of rice plants with little or no

breeding activity when the fields lie fallow There was no direct

relationship between breeding and seasonal or climtic chages Mean

vii

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 7: SECO1ADARY MiPests of plants--Philippines Reproduction and

LIST OF TALES

Table

i Monthly mean rainfall and average temperature in Siniloan Laguna------------ 8

2 Size of the ricefield rat t various stages of mturity ------ ---- ---- ---- ----- 13

3 Moithly composition of adult male population of Hattus rattus mindanensis with respect to the positionof testes - -------- - ------- 15

4 Monthly changes in the prevalence of pregnant Rattus rattus mindanensis collected in Siniloan August 1971 through Jul 1972 - - - - 22

5 Monthly mean distribution of fetuses in the two horns of the uterus of pregnant rats examined in Siniloan August 1971 through July 1972 ------ 26

6 Analysis of variance for litter size ------- --- 28

7 Mean litter size separation for all pregnant rats examined for this study in Siniloan August 1971 through July 1972 ---- ------- -- ------ 28

8 Productivity of Rattus rattus mindanensis captured during the monthly trapping in Siniloan Auguit C 1971 through July 1972 ------- --- ----- 29

9 Monthly changes in the young-adult ratio of all rats collected for this study in Siniloapi Laguna- - 31

10 Mothly changes in the sex ratio of all rats trapped in Siniloan August 1971 through July 1972 - - - - 33

11 Number of pregnant females aptured in relation to staesof DremancY 35

12 Sex ratios in various weight classes of attus rgttus mindanensis ------------ - --- - 36

13 Population density index of ricefield rats in Siniloan based on monthly trapline catches - ---- 38

v

LIST OF FIGURES

Page

1 Portion ofthebull qtudy area showing condition of the rice paddies after harvest- ------------- 6

2 Scatter diagram of length of testis against weight of rats showing presence or absence of sperm 17

3- Graph showing extent of breeding season Judged by percent pregnant and immature ----------- 1

4 Monthly changes in the prevalence of pregnancy in -relation to total monthly rainfall and number of rainy days per month -- ---- ---- -- ---- 21

5 Graph showing relationship between breeding and monthly sex ratio - ------------------- 34

6 Monthly trapline catches of field rats in relation to rainfall and temperature -------- -- 39

ABSTACT

Mrges Bernardo Espiritu University of the Tilippines

November 1972 Reproduction and Seasonal Abundance of the Ricefield

it (lMttus rattus mindanensis Yearns) at Siniloan laguna

ljor Professor blo J Alfonso

The reproductive patterns and seasonal abundance of the common

ricefield rat were studied in an irrigated ricefield in Siniloan

IAguna from August 1971 through July 1972 The study area ias

planted to two crops of rice every year Of the two rat species

found the ricefield rat dominated the entire riceland comunity

comprising more than 99 percent of the total collection

Two age classes were recognized Juveniles and adults Adults

were further subdivided into breeding and non-breeding The vaginal

orifice became perforate at 5445 grams among fenales while the

testes descended to the scrotum at 13493 grams among males For

both sexes the skull ossified at 9079 grams The length of the testes

vas significantly correlated with the body weight Active sperms

were present in the cauda epididymis when the testes reached lengtha

of 2229 mm Once sexual maturity was attained males remained fertile

throughout the year

Reproductive pattern was sarply bimodal corresponding very

closely with the crop cycle Most litters were produced during the

reproductive and ripening stages of rice plants with little or no

breeding activity when the fields lie fallow There was no direct

relationship between breeding and seasonal or climtic chages Mean

vii

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 8: SECO1ADARY MiPests of plants--Philippines Reproduction and

LIST OF FIGURES

Page

1 Portion ofthebull qtudy area showing condition of the rice paddies after harvest- ------------- 6

2 Scatter diagram of length of testis against weight of rats showing presence or absence of sperm 17

3- Graph showing extent of breeding season Judged by percent pregnant and immature ----------- 1

4 Monthly changes in the prevalence of pregnancy in -relation to total monthly rainfall and number of rainy days per month -- ---- ---- -- ---- 21

5 Graph showing relationship between breeding and monthly sex ratio - ------------------- 34

6 Monthly trapline catches of field rats in relation to rainfall and temperature -------- -- 39

ABSTACT

Mrges Bernardo Espiritu University of the Tilippines

November 1972 Reproduction and Seasonal Abundance of the Ricefield

it (lMttus rattus mindanensis Yearns) at Siniloan laguna

ljor Professor blo J Alfonso

The reproductive patterns and seasonal abundance of the common

ricefield rat were studied in an irrigated ricefield in Siniloan

IAguna from August 1971 through July 1972 The study area ias

planted to two crops of rice every year Of the two rat species

found the ricefield rat dominated the entire riceland comunity

comprising more than 99 percent of the total collection

Two age classes were recognized Juveniles and adults Adults

were further subdivided into breeding and non-breeding The vaginal

orifice became perforate at 5445 grams among fenales while the

testes descended to the scrotum at 13493 grams among males For

both sexes the skull ossified at 9079 grams The length of the testes

vas significantly correlated with the body weight Active sperms

were present in the cauda epididymis when the testes reached lengtha

of 2229 mm Once sexual maturity was attained males remained fertile

throughout the year

Reproductive pattern was sarply bimodal corresponding very

closely with the crop cycle Most litters were produced during the

reproductive and ripening stages of rice plants with little or no

breeding activity when the fields lie fallow There was no direct

relationship between breeding and seasonal or climtic chages Mean

vii

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

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41

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42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

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0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 9: SECO1ADARY MiPests of plants--Philippines Reproduction and

ABSTACT

Mrges Bernardo Espiritu University of the Tilippines

November 1972 Reproduction and Seasonal Abundance of the Ricefield

it (lMttus rattus mindanensis Yearns) at Siniloan laguna

ljor Professor blo J Alfonso

The reproductive patterns and seasonal abundance of the common

ricefield rat were studied in an irrigated ricefield in Siniloan

IAguna from August 1971 through July 1972 The study area ias

planted to two crops of rice every year Of the two rat species

found the ricefield rat dominated the entire riceland comunity

comprising more than 99 percent of the total collection

Two age classes were recognized Juveniles and adults Adults

were further subdivided into breeding and non-breeding The vaginal

orifice became perforate at 5445 grams among fenales while the

testes descended to the scrotum at 13493 grams among males For

both sexes the skull ossified at 9079 grams The length of the testes

vas significantly correlated with the body weight Active sperms

were present in the cauda epididymis when the testes reached lengtha

of 2229 mm Once sexual maturity was attained males remained fertile

throughout the year

Reproductive pattern was sarply bimodal corresponding very

closely with the crop cycle Most litters were produced during the

reproductive and ripening stages of rice plants with little or no

breeding activity when the fields lie fallow There was no direct

relationship between breeding and seasonal or climtic chages Mean

vii

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 10: SECO1ADARY MiPests of plants--Philippines Reproduction and

litter size was 967 but decreased significantly during the beginning and termination of the breedinp period There vas no tendency for the number of embryos to increase with body weight The distribution of embryos between the right and left horns of the uterus vas entirely independent with no marked tendency for all embryos to develop on one side or the other The annual incidence of pregnancy was 5 67 litters the number of litters produced during the dry season breeding as pore than twice that of the wet season breeding With few exceptions the sex ratio approximated equality Of the exceptions most could be shown due to differential tra Pability of both-

sexes at certain star of the

breeding period Productivity was high about 44 vouni bInft ThMiwaA

per female per year

Seasonal reproduction resulted in high PopulAtion density by August September and October sarp decline in density was observed

in January February and Mrch

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 11: SECO1ADARY MiPests of plants--Philippines Reproduction and

MTRODUp7ON

1ts have alwys been a problem in TiJippine rcuizure here

early reports of serious rat depredations in the Bicol region andwere

in the Mountain Province (Crucillo otanes and Morales 1954) but these

lacked the details that would provide better meaning and importance to

such apparently localized and short-term irruptions Simiar severe -- 2 5 i

were Imown to exist ininfestations but notably of longer durations

large sugr estates in Mindoro and Negros islands (Utzon personal

cozunication) but loss of records during the last mar denies to pre

sent workers information on rat species and their patterns of growth

and abundance in cropland conditions The near simultaneous and widespreno

rodent outbreaks in Cotabato (Sarmiento 1953) Davao a4d adjacent i~ i bull -

provinces in Mindanao in 1952-53 provided the first Ehilippine exmprience

which considers rats as an agricultural pest of national econoic

invortance (crucillo et al 1954 and Villadolid 1954) Since then

rat outbreaks occurring at times to the proportion of a plague have

severely affected rice and corn roduction before harvest while also

affecting the potentials of sugar and coconut exports in all principal

islands and regions of the country

The Thilippine ricefield rat Rattus rattus mindanensis MRrns Is

the most prevalent rodent form affecting local agriculture (Barbehenn

Sunanil and Libay in preparation) It is chiefly responsible for most

of the danage in cereal crops where the value of crop losses to rice and

corn alone vas estimted to reach 160 million pesos annually (Sun gil

et al 1970)

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 12: SECO1ADARY MiPests of plants--Philippines Reproduction and

2

Inspiteof its widspread distribuitiow the riceplusmneld rat has

been the-subject of relatively feWtailed studieson Aeproduction and

popUdtioi dnamics To Obtain relible records oa the breeding of most

wil ls onemutrely iinlj on Dbst-mortem ezingtions of an

adequate number of the-species collected at intervals over a period of

6iWVY more years These conditions were approached by Rosell (1955)

with otr 6onsecutive years of study on the breeding activity-of attus

ak~nftftte in ricefields and vastelids of Cotabat6and by

se 1and Mcaz (1963) on three intervelig-periods in the succeeding

(1958) further reported some beeding data years Singil and Aqdlno

on AR_r mindanensds and -argdentivente collected in cropl nds and

forest clearings inlknbuiAro M1idoroOccidental SiWpl6entary data

on the ontogeny and laboiatory breeding Were aiSo obtain by Rosell

(1959-)on R4 argentivefter and Medina et al (1972) on he7i mindanensis

M1iddetaill -6n the kijrodudtivity and breddingactivity of RI 4N

taidaneni-amp6 the basis of geoaaphic Iocachirwere repotd from

International Rice ReSearch Institute Colleg6 lagu (Uler 1967)

40oi cift dii lfties df lAguna (fan~bei di 41)171) land Mufiozt

N aEcj Ad(Afonsoand Castillo 1970)0 It is apliWehVr6n n thesw

reporbs- eii 6rddfed areas ampreiniclved -the in breetig period

coinede ith te rali y season and -pekiod ofrice dulti-iation (-( wto

Ddleml ii Vi~thucl reduced breeding abivity durIig-theAdVoiFufr

bull h 5 (3 btrty) This breeding pe dicitY46eS tllt 44dJ

seasonal build-up of population andMyat1o AtIdy = ther~adits

obtained by LaVoie et al (1970) who found that tiller destruction by

rats during the vegetative growth period of rice mas light but increased

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 13: SECO1ADARY MiPests of plants--Philippines Reproduction and

3

appreciaby during thereproductiveAnd ripening Stages of development

Si~i1(971) observed-tb th presence of~oie foodand qiur~r cover offered bybooting rice at acts rapement fsuod i

wastelands into rice fielp Thus 1irrtIye populations are cozsuonl1y associat to seri uscrop dam gebefore Orest

The basic breeding cycle ofanibals may be affected either favorably

or~ ~ pnionentalpoundhctor~ Trpugh bis agicltr1~b9yLuP ~asqL

pratices nan has altered his enviroment tno fvor -the existence of many

species includng fi ld rate Etensive use ofirrigation in Iag= province duin~g te~c Beasqn hs converted astelands and lowed areas

ids Hence irrigated p3nted to

rice during the dry season This provides rats with adequate supply of

food and cover even during the dry months of the year Uhlpr (1967)

observed at IRRI where irrigstion is continous and rice is grown throughshy

out the year that R r mindnensis ehibitsa year-round breeding season

The effects of env onmntAl-fctors ariculrq olhnwte and food requireshy

ments on ed cycle or rats is emzphasized by Sunangii (1965) who

states that where food is alays plentiful and olimtemore or lespeven

throughout the year rats breed and reproduce anytime oF month of the year

Knowledge on the reproductive activity of the ricqfield rat

particul ry breeding season is needed before any realistic reduction

program pan be eyolved B understanding population changes inrelation

to season and habitat reducion measures pay be instituted at a critical

stage begore apMulatcn increase

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 14: SECO1ADARY MiPests of plants--Philippines Reproduction and

4

The esent study vas undertaken from August 1971 through July

1912 to collect informtion on the reproductive pattern of rats in an

irriaatdk ricefield and to determine population changes in relation to

seasonal and habitat changes In a broader context the study offered -4

an opportunity to egamine in detail the mechanisms of population regulation

of a single mannalian species in a typical riceland ecosystem

TAXONONAID DISTRIBUTION

Rattus rattus mindanensis is the most widely distributed Philippine 7

species and is chiefly responsible for the destruction of rice and other

crops- in In Siniloan lAguna it represented thanLuzon0 and Visayas more i - 4-

99percent of the total rat collection all others are I0ttus exulans

Although originally described by Mearns as a distinct species (Taylor

1934) the ricefield rat is now designated Ratu rat mindanensis

(Barbehenn et al 1972 and Johnson 1962) It is medium-sized rat

with adults averaging about 180 gram The tail is usuilly longer than

the head and body ac S iiiforu7yblak dark brown scalation is

variable The hind foot averages 40 mm in length but the range of 36ste Jj4 owj Jsi)dE o -

to 45 overlaps that of many species of rats Plantar pads are well

developed Ears measured from the notch averaged 21 ma (range 19 to

23) 1elage color is quite variable ranging from grayish brown toV) i3 plusmn32 - r i j I

reddish chestnut dorsally and usully blending gpudually into a paler

venter which is even more variable = from crewr white to dirty grey

often with a wash of buff of variable intensity (Barbehenn et al 1972)

The ricefield rat typically hbas two pairs of pectoral and three pairs

of inguinsl e with the anterior pair actally being abdominl

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 15: SECO1ADARY MiPests of plants--Philippines Reproduction and

5 and more distinct from the middle pair tban the ps-teri- i par To the

h

non-taxonomist the various sub-species are very similar and wheramp ranges

in mea6sreent overlap it is extremeey ditficult to s jedti3 the

In the whole isiands of Mndanao and some parts of Mind6r here

1~ttus argentiventer dominates ricefield commuitiesi becomd more

cormensal in is distribution (barbebieni it a 2 cording to

Sumsngil (1968) this specisi3 apastjcUr liking for hunmn dwellings

and generally destrcys backyard fruit crops particularly coconuts and

banana plantations as well as stored rieeand-ern- - n Coflitdt more

than 90 percent of those caughi near and inside houis belong to this species narbehem (1972)et al considered R r mindanensis to be an

ede icsi--s-pecies in the Philippines since it is presumed to occur

on vfirully every island The anizls are found from grasslwnd at sea

id~jL to ove 6 ~fd n Luzon and Mindoro although they are less

conon in hiter altitudes

DASCPI OP STDY APtS

The stuy area vs located in Siniloan laguna aboub tvo kilometers

from IAguna de Day shoreline at an approxicte elevation of slightly above sea level The topography is relatively flat with sligW dec~ine

to the east tO~rd the base of Sierra Jdre mountain- Anarea of

axp~oncltely one square kilometer within and around the peraiphery of I 7

Daybay National Agricultur and Vocatio chool was used as the

prinmry collection site (pigure 1) It is dissected by a number of

sM11 and shallow creeks and lrger tributaries that result in overflow

during the rasiny season with any nmrked rise in lake water

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 16: SECO1ADARY MiPests of plants--Philippines Reproduction and

Elg 1 Portion of the study ae showing conditions of the rice pardies after harvest

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 17: SECO1ADARY MiPests of plants--Philippines Reproduction and

The climate is characteristic of the lowland areas of Iaguna

province (Table 1) The average annual precipitation for Siniloan over

two-year period vas 19904 mn Average monthly precipitation did not

seem to vary widely from year to year the average for 1970 and 1971 va 18825 and 20982 anm respectively The bulk of the rainfall usually

occurred from July through December In 1971 much of the rainfall

occurred during the period from October to December while in 1972 the

effects of monsoon rain cused a peak rainfall in July with 7670 n of

rain This abnormQly high precipitation inundated and submerged the

entire study area a week after the last specimen collection was Dade

The monthly mean temerature for Siniloan did not vary widely The

monthly mean tenerature ranged from 2469 to 28320C with an average

of 26780 C during the whole periodf study

Although there are two cropping season annually rice production

in Siniloan is entirely dependent on the depth of ater in the paddy

fields Unlike in nany other Iaguna towns the first or regular rice

crop is transplanted in ampnuary and all the paddy fields are planted

in late februsry H rvest occurs mostly in My and June The second

crop or r gad rice is transplanted in July and August and harvested

in November and December Because most of the areas are titer-logged

during the rainy season only about half of the total hectarage is

Planted to rice Hence the amount of cereal food available to the

rats during the rainy season is coortively less than during the dry

season

Weeds are a constant problem on the YAddy fields The unplanted

wras remain largely covered with succulent gasses compsed mostly of

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 18: SECO1ADARY MiPests of plants--Philippines Reproduction and

Table 1 Mont3 mean wa- -Iand ay ge t me tie J $inilcmn

aI deg - C

bullMn h z 70 3972 ~ ~ ~ ~ je 4 Ih1infal Ai ti14n~1 1ainfall

c ) -c)7

J niw 409 2555 125 2469 47o

February 168 2553 256 2541 25

M~rch 201 2718 827 2563 749

April 203 2828 183 272E 162

My 198 2926 2298 2832 1582

June 1775 a858 2511 277E 4280

Juy 1406 2773 33146 2747 7670

A t18204 147 1658 27714

September 3592 100 2085 27-90

October 5165 2-7025 418 2688 shy

5770 682 34Q 627

D376 Z8 654 4496 2597 shy

an 8825 2727 20982 2678 21340

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 19: SECO1ADARY MiPests of plants--Philippines Reproduction and

9 bayakibok (Echinoclo crusgalli) and large stands of tikiw (Scrpu

ampdsibl) are equaljcspVuos ie banks iid nin~sIal area6 ire~

coveredwith thick stands ofta3AhbIL-(Sacchrunm spontaneum) while

pddy dikes- Iirrigtioi canalsrwere dnminatedby thick nattin of

para -grass (BraShArly ica) The presence of succulent weedsand

xtooning rice plants after harveit affords the rats anple cover nd

food throughout the yehr

YATMAIS AND MTHCDS

A Coflection of Rats

lost-Drtem exminAtion of 932 rats collected at two-week intervals

from August 1971 through Ju3Ly 1972 in irrigted rice fields fo vxd

the basis of this study Animals were generally collected by Victor

traps Suplemen tl collections were made by band capture

(blaneting) of aninals ip grassy areas Snap traps were placed along

PA dikes in two or three transects each composed of 50 or more raps

Distance between traps was about 5 meters and the space between transects

Sried IccoxI4ng to the distance between paddy dikes or about 50to 100 --trap -lines wereeg usii3yoperated for2 or3--consecutive nights

Bits conisted of fresh coconuts or sliced sweet potatoes The trapshy

ping lines vere moved into a different collection site for each trapping

period and were located systenatically to enable zanpling the entire

study area- 1he number averaga of spe ns exined per month was 776

with a range om 14 to 194 rats Generally the monthly s6Mles

consisted of more than fifty awimls

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 20: SECO1ADARY MiPests of plants--Philippines Reproduction and

LO

B Autopsy

Necropsy was conducted iii the field as soon as specimens were

collectede Allan l wered identified sexed and weighed separately

In the of gravid fem3Js-the difference in scale after removingcase

the weight of the embryos and enlarge uterus was taken as the weight

of the parent In addition standard physical measurements (total

length- _tail length ear-length-and hind foot) were recorded

The reprodfobive condition of-the males was determined by examining

the position ofthe testes and redording them as scrotal or abdoiInal

Testicular length and -width was measuredin millimeters and the conditions ofthecauda epididymis werenoted FA s with enlarged scrotal testes

with visible tubules in the cauda epididymios -wcre -considered

reproductively active

The condition of the female reproductive organs were determined by

making a median incision on the -ventral surface of tbe abdomen and

examining the uterus id situ Females were classifiedas reproductively

active if they were pregnant lactatig orhad opened vWkgiolorifice

The presence of spots or placental scars in the mall of the uterus were

recorded However no attempt was made to count the uterine scars

because this criterion was not reliable for determining litter size

(Davis and Emlen 18) A rat as classed as pregnant if the embryos

are mcroscopicaly visoIle and lactating if milk can be expressed by

the fingers from the nipples For the embryos actual crown-rump length

As WDfsured when the specimen was large enough (3 mm or more) The

number of embryos contained in the right and left horn- of the uterus

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 21: SECO1ADARY MiPests of plants--Philippines Reproduction and

were recorded Dead embryos and those being resorbed (snaller than the

mjority that developet is obviously abnornal) were noted and used to esiliate the loss of embryos duning pregnancy

Ossification of the skull Mas determined qua3itativel7 by resistance

to finner vreskikd (Dalvis and HB1 1951)

Statistical Aa6es

The annual incidence of pregmncy or the number of litters born

to a female as Appr~ciiat d fvom estimtes of prevalence of pregnancy

(Emlen sad Davis 1948 Davis and Hll 1951) Sex ratios were expressed

in percent males and were tested for statistical significance by Chi-sqar

analysis Mtuikty based on the perforation of the vaginal orifice in

females presence of scrotdi testes in uesand osugfication of the

skull in both sexes were determined by correlation and regression analysis

3h determinLg reproductive -trends the data were analyzed by

=oths$ anad also by seasons Mnthly Mean litter size vas tested for

statistical significance by the use of Anaiysis of Variance and mean

differences vasdetermined by Duncan id ltiple Mnge Test

ESUIEW AD DISCUSSION

A Criteria of laturit

Determination of hrnonPogical age inferal rats isdifficult

because growth and developent rates r y vary In ear individualnd

in different PO1)l3ltioflsa were it tioable to ObserV9PnIividus over

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

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41

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43

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TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 22: SECO1ADARY MiPests of plants--Philippines Reproduction and

12 a period of time the differentages at which they attained a given level of developent could be determined However since the aninals

in the present sudy were examined at but one stage in their development

onthe presenge or absence ofa given character could be observed

Characters ofteA used for differentiating adults from Juveniles are based on the capacity of the animals for breeding Though annals are adult

by chronological criteria they can play no part in the reproductive

phases of the population dynamics until motile sperm are contained in nale reproductive tracts or until ovulation has occurred in ferles

(Jackson 1952)

Use of weight at which 50 percent of the aninls possess a given

characteristic is preferrable to 100 percent level since the latter

TV be undul affect-ed b a few slow3y naturing individuals (Jackson

1952) these 50 perdent points were calculated from the regression of

percentages on the body weight

Table 2 shows the sizes of the ricefield rat at various stages of maturity The weight at which 50 percent of the females have the vagna

foAte wks 54145 g~rams althoug femnles mysaome become perforate at wikihtas ow ai 10 grams However these newly perforate femles sho 06d t ofrpductive tracts Perforation is not indicative

sexual lturct in temas=f due to the presence in the population of

iew Per~i~ate fensles which lack ovarian developeent tIn the mles rapid increase in weight is not normally acccqsnied

bymaturation of the gonads or accessory glands The testes become

scrotal posiion a6t 13493 gras while sperms were present in the

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 23: SECO1ADARY MiPests of plants--Philippines Reproduction and

~13

V~ble 2J Sie of I ricefield rai at Varid stages of mturity

Weiwht in grams Characters No of rats 50 ercent poizt

TagIiriai iritibe per orate 450 5441

mvi1- osoiricition 90-797

st scrotalpoitn 477 134 3

Spe in -daud epidikvmis 55 15447

lAcental scars 450 18555

oauda epiidymis at 15445 grams Theyeight t which 50 percent

become fertile gives ainasure of the averageweight of onse of fershy

tility

Ossification of the skull occurred afterVt rats attined the

weight of 9079 grams

The presence of scars in the uterus is a good InUoplusmntion that

implantation has occurred Thus if the aninals are grouped by size

and classed as parous (indvidmils wi scars or visible embryos) or

non-rarous (individuals Vithout scars or visible embryos) a measure

of the size at first pre canyc beobtained adEmlen 1948)can (Davyi

The lowest weight at which first preuay ourred was 105 -grams which

can be taken asa measure ofpeurty 7pregnantof the age Of the lo+

females examined 8654 pe ighed re grams

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 24: SECO1ADARY MiPests of plants--Philippines Reproduction and

14

1 Testis Dositionl size and fertility

ibsition and size of testes have been used as the most important

criteria f breeding in mles Testes in the ricefield rats may occupy

either abdionil or pscrotal Both testes generaly occupyositions

the same position Descent my occur at an early age depending upon

the season since scrotal testes were found among males weighing well

below 50 grams during the months of April and October The proportion

of rnles with scrotal testes increases with weight thus showing some

relationships between position of testes and seuml maturity Recent

studies by the Rodent Research Center indicate that rats with active

spernatozoa invariably had scrotal testes although 45 percent of the

adult mles with no apparent sperm activity (epididyml tubules not

visible) had scrotal testes (Swink et al 1970) Descent of testes

isnot a very good index of fertility in the ricefield rats

Table 3 shows that there isa high proportion of males with scrotal

testes throughout the year Considerable reduction inthe propoplusmniOn

of males with scrotal testes in June and July mas due to the presence

of more Juveniles in the-population Since this species is capable

of producing active speriatozoa throughout the year in both frrigated

and rainfed areas (Swink et al 1970) then the occurrence of seasonal

changes in the Pregnancy rates cannot be attributed to seasonal changes

inthe capacity of nales to fertilize

Length of testis was used as an index of testis size Miller

(1946) observed that length and weight of testes among pocket gophers

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 25: SECO1ADARY MiPests of plants--Philippines Reproduction and

15

T~ble 3 Mnthr c~~sition of adult nle po tuJMon ojI rattas mindanensis with respect to position ot testes

Tota les stis 3ositiOn ercent Month exam er Abd inal Scrotal scrotal

Agst 1971 8 1 7 8750

September 40 12 28 700

October 30 8 22 7333

November 69 15 54 7826

December 114 28 86 7544

Januu 1972 22 8 14 6363

brw 10 3 7 7000

wxch 9 2 7 7777

April 43 13 30 6976

1y 38 14 24 6315

June 56 40 16 2857

4uly 43 27 16 5925

Total and average 8 171 311 6452

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 26: SECO1ADARY MiPests of plants--Philippines Reproduction and

are closely correlated (r a 087) so that length isan adequate estimate

of size Laurie (1946) found among mice that if the tubules in the

cauda epididymis are visible to the naked they invariably contained

large number ok sperms and that the indlJdil would be fecund In the

ricefield rats the presenceof sperm is usually associated with a testes

length of between 20 and 32 mm (Figure 2) It was also found that a

testes weight of 22 mm or over was almost invariably associated with

fecundity that is with large number of sperm in the cauda epididymis

Below the length of 20 mm the cauda epididymis was nearly always

without sperms that is to say the males are non-fecund

B Reproduction

1 Ereeding season

The seasonal reproductive pattern of 1ttus rattus mindanensis

in Sinilcan Iguna are sunmrized in Table 4 and Figure 3 Although

the proportion of adult sales capable of producing active sperms

remained high throughout the year (Swink et al 1970) reproductive

activity of adult females show seasonal trends Analysis of the

thlyprevalence of visible pregnancy andincidence of juveniles in

the population indicates that this species features a sharp bimodal

breeding curve peaks occurring from March to April and October to

NoNvember This is in general agreement with Uhler (19) who also

observed bimodality in the breeding pattern characteristic of this

species at the International Rice Research Institute College Laguna

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 27: SECO1ADARY MiPests of plants--Philippines Reproduction and

SPERM 0000 o NO ERM e e

0 0 0 o0 00

00S0 00

_6 0 00 U) 0 0 0

deg bull70 LI I t

Ishy

35 55 75 95 115 135155 175 195 215 235 255 275295 315 BODu Y WEi GH T (gin)

Fig 2 Scatter diagram of Iongth of losuis against woight ifrats showi~g presence or abaen~e of s~rw

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 28: SECO1ADARY MiPests of plants--Philippines Reproduction and

0 PREGNANT

901 A JUVENILE

soshy

70

oshy

0

0 o L 1

) 2

A0 6 F

40 ~bg at l eadr e

Fig 3 Graph showing extent of breeding season judged percenp3 7 by g~tandifimatiire-

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 29: SECO1ADARY MiPests of plants--Philippines Reproduction and

19

In other parts of the Philippines a unimodal curve has been found

In Cotabato the ricefield rat breeds actively during the rainy season

(June to December) followed by no breeding activity from January to

early part of Jmune (Sumangil e al 1963 Rosell 1959 and Sumngil

and Rosell 1959) Sanchez et al (1971) also observed that inNueva

Ecija this species breeds actively during the rainy season months but

ceases compltely during Mrch April My and June this period

corresponding to the dy season

Jackson (1952) stated that many environmental factors play a role

in the regulation of sall muml breeding The basis however for

the appirent relationship between these factors and increased breeding

is not well understood Bwruon (1936) observed that breeding season

appears to combine with the warm season in temperature and sub-temperature

clites But it appears that in monsoon areas the relation between

climate and breeding is not so simple It seems that breeding is largely

a matter of nutrition that my well depend on harvest and indirectly

on cliatic events

Harrison (1952) in Wayamprecorded a bi-monthly rhythm and attempted

to relate itto the lunar cycle No suchcorrelation has been attempted

with the present data Swmngil and Rosell (1959) generaly regrded

climate as the min factor affecting the rhythm of reproduction of the

ricefield rat They observed signifiant correlation between reproduction

and rainfall In laya HarriSon (1952) has shown that rats appeared

to have no breeding season although breeding activity my show fluctuations

related to raInfall

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 30: SECO1ADARY MiPests of plants--Philippines Reproduction and

20

In this study correlation of monthly prevalence of pregnancy with

rainfall showed a slight negative relationship (r a 035) When the

monthly prevalence of pregnancy was correlated with the number of rainy

fays each month the relationship renained slightly negative (r u 0446)

indi(cting that seasonal changes has very little effect on the reproductive

activity of the rats This tends to show that in effectively irrigated

areas where rice can be grown at all months of the years rats can breed

continuously regardless of seasonal fluctuations As shown in Figure 4

the rainy seamon breeding seems to be significantly correlated with

rainfal (r a o447) while dry season breeding appears to be negatively

correlated with rafall (r a -030) This mY explain the significant

relationship between reproduction and rainfall reported by Sunsngil

et al (19j3) ad S ugil and Rosell (1955) Apparently their data

were gathered from rainfed fields where seasonal cycle and crop cycle

correspond more closely than in irrigated areas In rainfed areas the

farmers entirely depends from rain to start land preparation and planting

Table 4 indicates that active brPeding in rats is directly related

with the presence of rice in the field In Siniloan dry season breeding

appears to be of greater intensity and of longer duration than during

the rainy season This my be attributed to -he fact that wider area

was planted to rice during the dry season Hence more food is available

to the rats as a source of nutrition to maintain their Lxeeding activity

Breeding my be reduced or terminated when cover and food supply are

depleted or when convetition for resources such as food nest sites and

harborage is high (Sanchez et al 1971) Sunangil and osell (1959)

observed that thu rate of reproduction of lampborstopy-rowed rats diminiches

with the reduction of protein supply

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 31: SECO1ADARY MiPests of plants--Philippines Reproduction and

1001

40

20

o A F M A M J

97F [972

600shy

500

S400

F306 200 _3sea

o 1 M 4 40100

AtS O N J F M A J J 197 1972

S20 bull bull i 4

4 4O

4

0 IS

6 bull 0 - 4 bull4 r= i

AS O N F MAMJ J 1971 192

Fig fMonthl o ge ithe preval~n~e of pregnancy in relation to total monthly rainfh11and n~mber of

rain~v days ter month

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 32: SECO1ADARY MiPests of plants--Philippines Reproduction and

22

Table b pmeaence of pregnnat M us ratty-Mnthly obnges in t-he mindanensis inSinilan August 1971 through July 1

Month No of adult

Number Percent pregnt pregnAnt

Embryos per

Growth stagesof rice

females fegale I

August 3971 14 0 0 0 transplanting

1052 650 tilleringSeptember 38 4

21 6 3157 967 booting headingOctober

79 38 4810 968 mature harestingNovenber

Decenmer 80 7 875 886 post-harvest

0 0 seedbed land prepJanuary 1972 23 0

131 3 2727 70 transplantingFebrury

5 4 8000 1075 ti11eringIiMrch

April 32 20 625o 1o45 booting heading

My 30 11 3667 1118 mature harvesting

12 2553 916 post-harvestJune 7

sepdbed 3AIn 3repJuly 37 (f2 541 750

Total-aidaverwges417 107 2802 967

aJ Prevaltie of pregnancy bised on total number of feniles with perforated vaginal orifice

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 33: SECO1ADARY MiPests of plants--Philippines Reproduction and

23

(1955) thiamine is essential forAccording to Nelson and Evans

in the rat Nelson and Evans (1955)the normal reproductive performwance

cited Even- ishob --(1922)-who itepbrtded the imediate cessation of

on vitamin B-deficient diets Ueno (1934)estrous cycles in rats placed

cited by Nelson and Evans (1955) iidependentlyand Cowrd et al (19 2) as

mentioned the occurrence iof reproductive distrUbance in rats deficient

in vitamin B or B1 These results strongly suggest that seasonal

fluctuaions inthe breeding activity is dependent on the quantity of

anIS3l and vegetAble food available to the rats

2 Prevalence of pregnancy

Prevalence of pregnancy (per cent pregnant) was based on the number

of adult teenles per month Annual prevalence mas calculated by

Table 4 shows thataveraging the estimates from the monthly samples

oriented around two peaksthe variability of the breeding intensity vas

of activity One occurred in November when about 48 percent of the

seually mture females were pregnant and a second and higher one in

Narch and April when about 65 percent of the mture females are pregnant

arch my not be reliableHigh perceamptige of pregdant females during

It is-since the number of individuals involved ws not large

possible that non-breeding animals coming from surrounding wastelands

rp hsnyaccountwre ificluded inthe samples during th~e ~aaa

This is infor the low )rproductive rate during the rainy season

agreement with the observation of Sumangil (personal cncm) at Midsayap

Cotabato where he found that one-third of the annual reproduction

occurred during the s planting

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 34: SECO1ADARY MiPests of plants--Philippines Reproduction and

Although the data from the present study are Inadequate to determine

actulf-the factors rtesponsible for initiating active breeding it is

evident that the 0wevalence of pregnancy increased markedly during the

repr6ductive and ripening phases of therice growth A prevalence

of 526 and 875 percent during the months of July and December indicates

a shrp decline and termination of the breeding seasons

3 Incidence of Meoncy

The incidence of pregnancy or the number of litters born to a

female ddtwing a known 1eriod of time was calculated from estimates Of

the prevalence of pregnancy (Emlen and Davis) 1948 Davis and Hall

1951) by using the formula I Pt where I is the incidence during a d

time P isthe prevalence of pregnancy t isthe time of interval of

observation -i days and d is the number ofdays during the gestations

period of Tatsthat embryos are visible in the uterus which is about

18 days

The annal lujaidence of pregnancy of Siniloan field rats of 568

liters wnsbased on 1417 adult fenales This isnot sigpifieantly

different frmnIthose reported by Sanchez et al (1971) of 59 liters

for this Opeciel in three southern municipalities of guM It 1si

slightly higher (I -5) though nay not be statiticall- siifitcant

from the observations of Sumangil and Rosell (1959) for this nhspi

in Occidental Mindoro and for other Speciesnfor most oher-ares of the

world (Jackson 1952)

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 35: SECO1ADARY MiPests of plants--Philippines Reproduction and

25 Ro compre the incidence between two crop cycles the data are

sumarized for seasnal differences as sugested by Davis (1953) The

pqocentage pregnapt for all months (or seasons) were summed and then

divided by the znumbprofmonthp The method essentlaiy assumes that qile numb)= of t (100 was caught in- each month --Te incidence

of pregnancy was 405 litterls during the dry season and 164 litters for

the rainy season It appears that ricefield rats in Siniloan are able

to produce more than twice-the w er f litters cbring the regular

cropplanting than during the palaad or second crop planting This

results in a rapid increasp in the population towards tb tormination

of the dry season breeding

4 Litter size

Estimates of litter size was derived from counts of presumed normal

fetuses Ased on fetus counts the mean number of embryos per female

is 967 with litters ranging fromu 5 to 17 (Table 5) Counts of very

young fetuses my be of less value in determining litter size as some

loss ofyoung my occur after this age (Wirtz 1970) Loss of ova before

fertilization or resorption of embryos are among the fctors that may

reduce littersize in ammls Resorbing embryos were found in 11 of

107 pregnnt females examined One fetus ws affected in more than half

of thp oases while 2 were affected in 3 cases Intrauterine mortalty

Of 212percent is signifant1 lowerthan that reported by Smwngil

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 36: SECO1ADARY MiPests of plants--Philippines Reproduction and

26

Table 5 Monthly mean distribution of fetuses In the two horns of the uterus of pregoant rats examined in Sinilcan laguna August 1971 through July 1972

No of Total no enno of NMths pregnant

fenles of

eMgros Right horn

Left horn

embryos

Augut 1971 0 0 0 0 0

September 4 26 12 i4 650

October 6 58 28 30 967

November 38 368 179 L89 968

December 7 62 31 31 886

January 1972 0 0 0 0 0

February 3 21 1l 10 70

IMrch 4 43 21 22 1075

April 20 209 110 99 3o45

My 1 123 67 56 1118

June 12 110 56 51 916

July 2 15 10 5 750

Total 107 1035 525 510 967

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 37: SECO1ADARY MiPests of plants--Philippines Reproduction and

aui )Rpslh 9)o 6pret This pay be attributed to tie

difficulty encountered in determi frmnnornal dresorbing fetuses

during the early stages of pregnancy

ble 5 shows the distribution of fetuses in the two horns of the

uterus Based on 107 pregnant femlesdisseotedfrom which 1035

embryos were counted 525 or 5072 percent were lodged on the right horn

of the uterus while 510 or 4928 percent were lodged on the left horn

of the uterus Except for one case where all of the 7 embryos developed

on the right side of the uterus there seems to be no tendency for the

embryos to develop on one side of the uterus or the other It is evideni

that embryo development among the ricefield rat is entirely random and

independent This is In contrast with the report of Harrison in Malaya

(195L) where one side of the uterus was favored for embryo developmnt

in Rattus argentiventer

Table 6 shows the analysis of variance for the number of embryos

of all pregnant females The mean separation are shown in Table 7 The

average litter size recorded during the months of 1ebruary Ju3y

September abd December are significantly smoller (pu 005) tba those

obtained for the other months This indicates that litter size produced

between crop cycles decreases when food is scarce nnd competition for

other resources is high There seems to be no correlation between the

weight of the paent and the size of litter (r a 02)

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 38: SECO1ADARY MiPests of plants--Philippines Reproduction and

28

Table 6 Analysis of varlaae for litter size

Source of Degrees of Sun of mean

vaziatQI r-di i sams ic Los -value

th 9 12055 1319 249

97 IMr510I 537I

Total 106 61a56

ISignificant at 5 6 i

Table 7 Mean littir size iiaxation ofall _egait rats Ii aanined for this study iniSiniloan Z =a

Sept Feb July Dec June Oct Way Apr -Met Ift

L

6z50 70- 75 886 216

992-w

268it

ioI42 1075 n 18

SNoawn mcqui b 14

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 39: SECO1ADARY MiPests of plants--Philippines Reproduction and

29

ratio~ - pro o of adult fem1esi

popltjtion Ason in Table - the embryo productivity vas 4396p shy

femle per year This values obtained bymultipJying the average

n mber of viable embryos per female and the incidence of pregnancy Bia

rate (birth per rat per year)ia calculated by multiplying the anal

Wbrm0 pozumy---he percentage of-ultIWales -itbe -populetion

and the sex ratio (Davis and Golle4r 1963) Tab4 8 shows that the

rate of birth among ricefield rats mas 931 young per rat This did nol

e 81InnI IFI Idanensin t captured du i n

rp mgmn nq mde l A 88

omonths vthout-b1fo O- it li- Vrig20

a shyAverage-tio of viable enbryosfe leaJ 774

Embryoaplusmne -ieyear 4396

Age ratio (adult) 7379

Bex ratio (nale) 5183 Adult females (of total) bull0

Birth rate (birthratyear) 931

a Base qpon in ut toconto and correeatedby an estLteof 20 uterine morl tfpibl~shedfr cl6sely -Jr tedspecies

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 40: SECO1ADARY MiPests of plants--Philippines Reproduction and

30

differ significantly from the results obtained by Sanchez et al (1971)

for bothrainfed andirigated conditions In the real population

young and even nomeadult femles do not breed hence the rate is even

lowqr These results show that uder thd conditions obtaining in the

rice fields the rats have a high potential rateof increase

6 Juvenile incidence

Supplementary and confirmtory evidence on breeding may be furnished

from other sources The age composition particularly the incidence of

Juveniles may provide valuable clues A continuous stream of juveniles

contributes to the mnthly sanples but their relative frequencies that

the gestation period of rats ranges from 21 to 22 days Considering the

lactation period of 21 days (Medina et al 1972) the highest percentage

of Juveniles appeared in the collection about two months after the peak

of the breeding period The smllest rat captured weighs only 24 grams

whichmy be just about the size at weaning

There are two peaks in the incidence of rats less than 80 grams

in body weight (Table 9) These peaks confirms the existence of two

min breeding seasons of rats in Sinilcan Both occurred about two

months ater the peaks of pregnancy has been attained A Mrked increase

in the number of wal rats uas noted in January following the peak in

November An increase in the proportion of juveniles in June and July

substantiates the hypothesis of considerable breeding during the sumer

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 41: SECO1ADARY MiPests of plants--Philippines Reproduction and

Table 9 Monthly changes inthe young - adult ratio of al raiiCollected for this study at Siniloan IASUnA

Months Toamp 1 Number Number -Pecent

Rats ofy omxl ofadlt 8 ~ n August 1971 25 21 16oo

September 78 12 66 1538

October 51 1O 41 1960

Novedber 148 15 133 1013 December 194i 31 163 1598

January 1972 62 237 4032

February 21 6 18 2857

Iftch 14 2 12 14I28 Az j7 i76 62 1842

2Y77 54 3807 JuIe 108 63 5000 uy 84 o a4762

Total 932 240 692 2621

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 42: SECO1ADARY MiPests of plants--Philippines Reproduction and

32

months of Mrch and April The low incidence of Juveniles in Nitch and

shyeroenb~e~rco susiio -tatt there is little0r-no breeding

activity during the months of January and August

C sex atio

Small mMM trappers have often foimd a slight preponderance of

nales in their returns Among the workers with the present speciew

(Tigner 1$72 Sanchez et al 1971 Swink et al 1970 Alfonso and

Castillo 1970 Uhler 1967 Simngl and Aquino 1958) and with

Rattus argetiventer(Sunangil et al 1963 Rosell 1959 Harrisonshy

1951) Timer and Alfonso were ony those whose samples were large

enough to demonstrate that sex ratio of trapped rats vas statistically

significant from a ratio of 50 percent mles in favoT of the females

A total of 760 rats were trapped from August 1971 through July

1972 consisting of 400 rles and 360 ferles (Table 10) The over-all

sex ratio of 5183 percent mas not statistically different from the

ratio of 05 mles However the monthly sex ratios Indicated a

variable pattern There appears to be a high correlation (r a0723)

TheProportion of zia1esin -thessugae increased AppreoablyOur4Ing the

height of the breeding season and tended to decrease during the noAshy

breading months (ZLgure 5) This may be attributed to the wider home

ranges of the mules than the females Studies of Venezuelan roof rats

by Gomez (1960) indicated that sales were recaptured up to 80 meters

from the point of release while most females were recaptured from 15

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 43: SECO1ADARY MiPests of plants--Philippines Reproduction and

Sln~loa c n i e sex14 Of all rats traplagura August 1971 tju6ah JMIfV I172

i

Mont s Total- Number of Number of SexaJ Chishy

rats --Males -fenes ratio s~usre August 1971 25 8 17 320 324 SepteMber 74 40 38 5128 005 October 51 30 21 5882 158 Novenber 84 50 34 5952 067 December 82 W 34 584 o86 Janwwy 1972 62 22 40 3548 522 Fmbiry 23 12 44782 0002 WrCh 14 9 5 6429 314 Aril 78 45 33 5769 185

y 71 38 33 5352 035 June 108 56 52 5185 015 u 8 41 51-9 019

Total 760 400 360 5183 21o I i f I

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 44: SECO1ADARY MiPests of plants--Philippines Reproduction and

---

100

9 PREGNANT 90

O SX RATIO

z Go-110 I - -

So50--0 40

a30shy

10shy

70

A S 0 N D J F M A M J J

1971 1972

Figure 5 Graph showing relAionship between breeding and monthly 3ex ratio

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 45: SECO1ADARY MiPests of plants--Philippines Reproduction and

35 to 20 meters from the original trap but not much farther Thus widely ranging males would come in contact with more traps and would be caught more often in relation to their absolute number than females IMrkes (1924a as cited by Laurie (196) however found among laboratory-bred

mice that the proportin of males was lowest in the height of the breeding season This sexual difference in iovement might also be induced by the care that females must give to their young in nests and burrows Femle rats n advanced stage of pregnancy become less and less mobile and their tendency to leave the nest is reduced

correspondingly Consequently their chances to meet with a trap will be reduced correspondingly (Table 11)

The Secondary sex ratio can be further altered by differential growth and mortality (jackson 1952) Analysis of the data by size group (Table 12) indicated that the males predominate in the youngest cass and females in the oldest class The deviation from the 50

Table 1i Number of pregnhfnt females captured in relation to stagesof pregnancy

embryos (nzn) Assumed-- (mame (days) Total Total Average parfeales erYos femle Less than 7 m 6-n 58 593 1022

718 mm 12-16 31 29 942 More than 19 mm 17-22 18 150 833

Total and averages 107 1035 967

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 46: SECO1ADARY MiPests of plants--Philippines Reproduction and

36 Table 12 Sex ratio in various weight classes of rattus rattus

mindanensis

Weight Total Number of Number of Per cent glassg) - rats males -feemles- males

105 183 111 72 6o65

105-184 270 116 154 4296

185-262 388 214 174 5515

265 82 39 43 4756

Total 923 480 443 5240

percent ratio in the youngest class is suggested by Ecke (1955) to

be due to the faster growth rate among males Predominance of females

in the oldest size groups suggests greater male mortality with age

D Ft Pepulation Indices and Seasonal Fluctuations

Table 13 shows the monthly population indices of rats inthe study areamp There were only two species of rats caught in the rice field

for the whole trapping period All rats caught except for four

ttus exulans were FiRttus rattus mindanensis comprising more than 99

percent of the total collection

Using the trapline catches as an index of rat population the

Population shcyed monthly fluctuations (Fig 6) The rat population

density seemed to increase with the advern of the rainy season

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 47: SECO1ADARY MiPests of plants--Philippines Reproduction and

37

Table 13 Population density index of ricefield rats in Siniloan -based on m=th3y trapline- catches

onth Number of Total rats Trappin incidence trapnights caught

ea bull

August 1971 110 25 2272

September 473 7e 1649

October 295 51 1728

November 590 84 1423

December 960 82 8541

Janaryp 1972 638 62 972

Febra 258 23 89

Mrch 376 14 372

April 717 78 1088

546 71 1300

June 600 108 1800

July 290 83 2862

Total 5853 748 1295

Average percent trapping incidence per month

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 48: SECO1ADARY MiPests of plants--Philippines Reproduction and

Legend 0 Trap index

70 A Rainfall 1700

7 0 Temperature

I 60 t 60Ql

50-1500

B 0 I

4 _

300404

Xi~ -1 -C_ c - c shy10 10

20o

A S 0 N D J F M A M d J 21971 I97

of fedirplinrats ishyeFigure 6 M onthly ctche s

relation to rainfall and femparcdwe

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 49: SECO1ADARY MiPests of plants--Philippines Reproduction and

39

reaching the peak in iily and August During the month of August

when the field lie fallow ratu can subsist on ratooning rice succulent

grasses and insects The lowest trapping incidence occurred in

January February and N rch after the Mlagd crop has been harvested

This period corresponds to the dry months in Siniloan Since rice is

a preferred habitat its absence during these months apparently increased

conpetition for food cover and other resources resulting inthe

dupletion of the rat population through increased movement or migration

into the surrounding w -telands A vigorous wat control campaign was

also maged during the dry months and this may have aided in the abrupt

reduction inthe size of the rat population In =lRI however Uhler

(1967) observed that population peaks occurred from June to August

dFebruary

dynamics of one rat species inthe different areas points to the role

of environmental influence on rat populations

and November t This marked dissimilarity inthe population

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 50: SECO1ADARY MiPests of plants--Philippines Reproduction and

4o

LI TUE CITED

AIFONSO PJ and JC CASTILLO 1970 A contribution tQ the ecology of the ricefield rat (Mttus rattus mindanensis 1earns) in Mufoz Nueva Ecija 10 pp (Typewritei

and JL LIBAY 197R Rodents of theBARPEHENN KR JP SUFANGIL Philippine Croplands 33 pp (Typewritten)

BUXTON PA 1936 Breeding rates of domestic rats trapped in I~gos Nigeria and certain other countries Jour Anim Ecol 5 53-66

CHAIANE VH 1941 A trap-removal census of smli mamalsbull J Wildl mgat 5 42-67

CRUCILLO CV FQ (FYES and JL MORALES 1954 What We Ire Doing to control field rats in Cotabato Pbilippine Islands A syrup on Rat Control Div Biol Sci Nat Res Council Phil pp 25shy45

DAVIS DAVID E 1953b The characteristics of rat populations Quart Rev Biol 28 273-401

and JT EMIN 1948 The placental scar as a

measure of fertility in rats J Wildl Mgmt 12(3) 162-166 _

and FB GOLLEY 1963 Principles in Ivmmalogy Reinhold Book Crop NY 335 P

- and 0Hall 1948 Seasonal reproductive condition

of nale brown rats in Baltimore Maryland Physiol Zool 21 272-282

_ 1951 Seasonal reproductive condition of feale brown rats in Baltimore 1ryland Physiol Zool 2926

EMLEN JT and DE DAVIS4 1948 Determination of reproductive rates in Tat population by aamiaation of carcasaes Pbysio Zoo 21(1) 59-65

ECKE DH 1955 Analysis of populations of roof xats in southwest

Georgia US Public Health Monogr 27 1-RO

EMLEN JT AW STOKES and DE DAVIS 1949 Mqthpds forestimating populations of brown rats in urban habitats Ecology 30 430-442

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 51: SECO1ADARY MiPests of plants--Philippines Reproduction and

41

GOMZ JUAN C 1960 Correlation of Population of roof ratsinVenezuela with seasonal cldnges in bitat Amer Mid- i 63(1) 177-193

HAU-ORIONJL 1951 Reproduction in Mts of th6 hubgenus 1EttusBroc Zool Soc London l21(3) 673-694

1952 Data on the reproduction of some mlAyanlmmls Proo Zoo Soc LU don 125(2) 4-146c

HAYNE DW 1949 Two methods for estimting population from trappingrecords J Yhmm 30 399-411

JACKSONJ WILLIAM B 1952 Poulation of wild mouse (Iroarseus leucous) subjected to the applications of DDT and parathionEcological Monographs 22 259-281 -

1952 Reproduction In Ihcific Islxnd tEcology T-I Storer (ed) Bernice P Bishop Ma-Bull 255 p 90-92

JOINSN DH 1962 Rodents and other Micronesian mnul1s col atctedIn Pacific Island M~t Ecology TI Storer (ed) Bernice P Bishop lMm Bull 225 p 21-38

iAU13 BMD 1946 Reproductlion of house ose ( ismusculus)living indifferent enviionmdnt Trans Royal Soc London

vol 133 (B) 248-281

IAVOE G FN S K and JP -UsINGIL 1970 Destructionof rice tillers by rats in relation to stages of rice develomentinLuion Fhil Agric 54(3-4)-175-8j

LESLIE PH UM VENABIES and LSV VENABIES 1952 The fertilityand population structure of the brown rat (M~ttus norvegicus) in9rn ricks and some other habitat Proc Zool So London 122(1) 187-238

RIANO WIAMA P 19(o A contribution tb the blology and bonto1of a rodent pest affecting coconut (Cocus nucifera Linn) Unpublished Undergraduate Thesis Univ Phil 36 pp

MlDINA F IS CL-WDRIACA BD]E IA CRUZ V poundAGUIGIAN r ADBEIAMNS ESLIT2 and To =~GORIO 197~2 A study on theada~ttion andreding of Xield rats (Rattus-rattu sp) in the laboratory MDCAECp- MUn Quezon City l7 -meograp d)

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 52: SECO1ADARY MiPests of plants--Philippines Reproduction and

42

MLIER MILTON A 1946 Reproductive rates and cycels in the pocket gopher Jour Mmmal 27() 335-358

NELSON NM and HM EVANS 1955 Relation of Thiamine to reproshyduction in rat Jour Nuteition 55 151-163

NICHOLSON AJ and DW WAMER 1953 The rodents of 4ew Caledonia Jour Mlnm4l 34 168-179

ROSELL EON C 1959 StUdies on reproduction of the ricefielJ rat rattus argentiventer (Robinson amp Kloss) in Central Cotabato Unpublished MYsters Thesis Univ mhil 44 pp

SANCHEZ FF JP SUMINGIL FN SWINK G ATWELL A DE IA FAZ M FALL J LIBAY D SANCHEZ D TOLE INO R WEST and N KVERNO 1971 Annial Report Rodent Research Center College Lagun 80ppmimeo Unpublished

SARDMINT ARRV1953 Unpublished Annual REport to the Director of PlAntIndustry Bureau of Plant Industry Cotabato Cotabato pp

SOUTHWICK CH 1966 Reproduction mortality and growth of murid rodent populations Froc Indian Rodent Symp Calcutta Tndia pp 152-176

STICKEL LF 1948 The trap line as a measure of small mammal populations Jour Wildl Mgmt 12 -153-161

SUANuIL IJEsUS k yon Hanaooox on Tne ecopogy oT riceiield rats and their control by chemical means Pest and Dis Cont Div Bur Plant Ind Mnila 27 p (mimeographed)

0 1971 Burrows and breeding of ricefield rats in Cotabato Philippines Unpublished Masters Thesis Col State Univ Denver Colorado 38 p

SLMNGIL JP and A AQUINO 1958 Studies on rat reproduction in Namburao Occidental Mindoro Pest and Dis Cont Div Bureau of Platt Ind Yanila 14 p (-rpewitten)

and NC ROSELL 1959 Notes on the studies of Philippine species of field rats Sulamizam Bull 1(10) 41-51 (Reprint)

__ s LU SEQUI and AM MACANAS 196 Mchanism and effects of seasonal or cyclic reproduction on the outbreaks of ricefield rats in Cotabato Phil Jour Agric 28(3-4) 131-140

_xPJ ALFONSO FN SWINK and JL MORAIES 1970 More effective control of rats The Phil Recommends for Rice NFAC Quezon City 43 p (Brocahure)

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp

Page 53: SECO1ADARY MiPests of plants--Philippines Reproduction and

43

JP ~S~~SAG4 IAVOE14A~6DE Lk BkZ RR WESrDToIENTIN31J ~ n iIAYKUtRNO S~1HZ 1970

AWMZi Report Rbdent Reseearch Center College Laguna 56 pmilmeo Unpublished

TAYLOR EDARD H 1934 Philippine Land Mmlis Bui Sci MongrNO 30 Bureau of frmnting Yni3A 51 f9pp

TIGER JAMS R 197 Seasonal food habits ofR-ttusxa-ttus iindasshynensis (The Philippine Ricefleld Rt) in Central Luzon PhDThesis col State Univ Denver Colorado 66 p (Unpublished)

UHERm LD 1967 The reproduction of Rattus rattus mindanensis at I IRRI C611ege Laguna Philippines Phil AGric 51(7) 576-580

1967 The seasonal abundance of the Rattus rattus mindanensi at IRRI College Laguna Thilippines Phil Agric 51M7 5131586

VILIADOLID DV 1954 Some observations on the eoology 4nd biologyof rats in Cotabato A SyMp on Rat Cotrol Div Biol Sci Nat Res Council Phil pp 3-12

WIRTZ WIL M0 1970 Ecology of the JnesmmtRkt uexun on Kure AtollRaaii PhD Thesis Coll Unive Ann Arbor Michigan USA 132 pp