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CANADIAN THESES --

he quahty of lhts rntcrollche is heav~iy depmhl ufxm the La qualrtede cette rnhrohcbe +d gbndement de la qualit8 .qualtty of the orlgrslai ?hss!s submltW fcy r??aa5imng Every de la t h k xw&tse au rnmofttm'age Nws avms p u t fart pour e m har t e n made tic, ensure the ha quatit& &p&leure dd reprductlon bon p~sslbie

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it pagesare rn!sshg. c o n k ! :he univ degree. st&&-qut a w~fBr&+e grade

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Prev~ousiy copy ng9tc!d mafer!ajs fjourna a,~c!es & biisked tests, ete ) are qot fmeu, -

c, Reproductmn in fwior m pan-of inis fi-.n IS g o v w ~ ~y fw Canadian CooyngM Act 3 S C 1970 c G 3 C

Les dwurnmts g u ~ font ~ $ 6 ~ 4 I'obpt cfun dro~t cf.auteur (articles ae revue.,eamen's pub!r&s, etc ) ne sont pas m~crofilmbs

A Ja tot can3drenite sur re drat 6 - d e u r . SRC 1970, c 0. -

. %

THIS DGSERJATQN t~ T H ~ S E A E ~ E HAS BEE'N MICROFLMED . . ~ ; ~ C R O F I L . M ~ E TEE QUE EXACTLY AS RECUVEO &€US L*Av&NS WqUE ,- $ .

J

THESIS S L l 3 , M m I S P.4RTIAL FULFILLMENT b~

, ?HE REQLIRESEATS 'FOR THE DEGREE OF

f DOCTOR OF PHILOSOPHY

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L'autorisation a PtP accordBe j

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3 l a B i b 1 i o t h g q u e n a t i o n a l e "\

6 u C a n a d a de microfilmer 1

c e t t e thPse et de prEter oa de venare des exernplalres du '

f l l m .

L f a u t e u r f t i t c i a i r e au ciroit -

5'auteur) se r e s e r v e les , . a ~ t r e s drolts de pub1 icatron; n l l a t h g s e ni de longs e x t r a i t s a e c e l l e - c l ne zazven t "ere ~irnpr lmes ou a ~ t r e m e n t r e p r ~ c t i t s sans son a3tQrlsat l o n Ecrlte.

>

- 7 - '\

- -- 1 '-; j 3

x - -- r

Dr. #. t. t tnston, -4sswiate Professor, S e n i c r Supervisur

.- , - - - - - Dr. R . f , * t h e m ~ , k s o c i a t e Professor

, - - Dr. 3 . k b r d e n , FYofessor, h b l f c Exminer

Dr. 0. rtCul&eon,$~<r~ Progrim, R.C. Hinls t sry of .br tcdtttre and food, Pubi ic ,.Exminer

to U I ~ S 0 1 the S i m Fraser Universlfy ~ l b r & , and to make partial or 1 .. ringla copies only for such users or In response to a request from the

' l ibrary of any o t h e r unlvursity, or o t h e r educatlonaI I n s i i t u t l o n , on - . i t s own bshsif or tor me of i t s ussrs, i further egree that permission

lor multiple copying of this work for scholarly purposes m y be granted . . ccJ

by me or the Dean of Graduate Stud ier. l j i s understmd that copying

or p u b i t c a t i o n ot this work for financlad gsln sha l l lol be allowed

2 < -* d*.2L+gy ax2 zar.zgezer.-, -,f xil2 t e e and dgzwst ica ted k.or,e;;_ bee - - 1;

= s $ l i - a t c r s f3- 7 7 2 ~ f+<i? p~lljratlbn A

Author:

J (signature) T -

' (date) ' . - . .

r = - . ,

The objective of this smdy was to investigm 'the b i o l ~ ~ management md .- . -

- -

-

efftca~y of wiId bees and domkricared honey bees for nee fntir p o b t i o d ObServations

were made and experiments conduced' in the Okanagm Valley. B. C from 1983 to 1985. ' C

I - Atttiougb abundant in a a W habitats, unmanaged wild bees are not reliable or

4' predicable pollinators foi orchards in tde Ohnagan Valley. ~owev?. the management of ,

"6 . dominant wild bees species such as i3cnnbu.s rerricda ocdentalis and' B. b&nw nearticur,

which diT& a strong mrdnsy toward foraging orchard crops, does have potential for - ? .

C

fruit crop pollination. Alhough Omia Iignoria popinqua was not a dominant species b

orchard habitats. it has p0,tenr.d for uee fnair pollination' in the Okanagan Valley since it is - inQgenous to this area and management systems far its use in fruit 'tree p o h t i o n have

already been established

The results of a stndy comparing the relarive pollination efficacy .of honey bees and 0.

lrgwr~a suggest that, although single 0. ligneria visits produce fruie of a amparable size to

those resulting frem one to three honey. bee visits, overall pollination ef icacy u lower due $2: L - - -

' + , to the greater number of seedless carpels per fruit and therefore an increased tendency

I

toward as)immemcal apples. In adctirim, the data s u ~ k s t that 0. l t g w i o is less e f i s e n t af

. pollination sin% it spends dp i fmd! more time &aching for and foraging on blossoms than

do honey bees.

Predicting honey k colony pollination potenngl in cherries, pears and apples utilizing e

- , such famn as colony charaaeristia forager mmce counts and weather conditions is

possible. but predictions-will have to be made on an individual cmp basis nther than by

Colony characterisries and profitability were examined for three systems of honey bee i- ---

- - - - - --

managemenL Mawemens of coIonv weight, d e d worker brd area and surplus honey a

colony uigour, yielded the 'oes~ income, and mn provide a new source in me through -afa, the

safe of packages and, nudei. /

'C . " d-

DEDICATION

for his appreciation of people and To John Corner, the environment

4 . .

F& Drs. W. D. h e . R D. M-Mullen and H. F. ~ a h v n - k&iculm &da ~ k a r c h

H- -, t'. M y . R W. Brook T. Griswd& aqd R R: Snelling for heir ' the ideatificaticm af . " t

- e

iviId bees; Dr. P. F. Torchio TOT .pqvidtnp 'Osmio lignann for the polliriauon - o

< * Y

efFiac): study; Dr. R D. Roudedge for statisual 4istance: W. Camerop, V. James. C. Day. * = P

T: Solmer. D. Clandge. A Ckridge. G. Gteq ~ ' ~ o b e n s o n . and the Coldmeam Ranch' far r * -

9 permission to use their archards and kind $&in8 this study; and S. Renyard, B. Brsusseau.

. - & . , R. R o m u . K. Woodward L Fngurnq I€. Naumanh S. MilcheU. L Chong. M. >&le , - a

3 ' 7 -T - and G. fudd Specrril t h k s are due to m y Senior Supeniwr. Dr. M, L

- *

Winston, for his understanhe and guidance. .. 7 I

" /' Finally, my sinckmt thank you - - to - Roben - - Dupree. - for his patienc?: m u r a ~ e m e n t a n d

, / undersmding

t

- -

This research was supponed b) pants f'om the Okanagan Valle) Pdiinauon Asmiation. .

rhc British Columbia Science Council (G.R.EAT. Award tr, C.D.S. and a research grant #,

M.LW.) 'and operating grant AJ774 f r m the ,htural 'kiences and hgheerjng R m c h

Council of Canada.

- 5 ..................... .......... .................................. . + c is DEDICATION ................. ,:---..--...-.------ .-.....-, .;: v I

, - ~. ,- &

*~; ." * . :% .)

v: .*.- ?.< ... ACKN0WLETX;EMENTS .- --.--... f......I..............U.I...-....Y..I..~.....-.-~.- . . . - 3

"p

................................. .......... ..1. Lisr of Tables .- ................................. -.---...- + ..,...-..........-......-..-.-....... , ix

Q A Wild -bee pollinator diversity and abundance in orchafd' md uncuitivaml bbitars e-. * . , i n the 0- Vattty, British Cdumbia ,-__..................-...-.*.~..--.-.-.--.-...-.........-..-.. 4 f - --

.................... . M A r n A L S AND m o m ,--...- ---.---.....FC1 ..........-.C1.C1.C1,..--*.C1....... 9

II. Abundance of Wild Bee Species ,,.. .............................-.-.-.----...... 23

, ,

I. Species Distribution Pawm --,--,., ....................tri.tri.tri..--..-.....+...................... 33 4

-

II. Annual Distribution hne rns -+-.,-....----.-.-,.-..;..........,.A ...................... 36 #--

IIL Orchard Pollination e l ~ .-,-.---..-........-............-.-.............-.-...- ..-. - ...-...,-........ 38 d l

B. A comparison of the pollination e f i a d e s of honey bees. A p r m e w m L and Osmia d~gniana propin& Cr- on Red Delicious apples ,,.......,,,.... 44

EL lmplicarim for Poliination Management Systeins ..-...... ,,., .,-,.,,.,....-.. 68

f PI-SCUSSION .................-....-.a..---.-...-.----...--.-.-.....-...............................-..............-......................... 102 #

- -

CONCLUSIONS L.L.--- - - . . - , ......-......,...-.......-................................-.............-.......-.-............. 105

APPENDIX I ............. I..., .... -.-L.--. -..-...... - ...--.. -.... . . . . ............................................................... 108

- 4 4 Table 4. MA capture nur (bm/h) of wilildbec families i o m c h d r and

uncultivated, habitas 111 the OIranagan Valley during 1984 and 1985 ,.-...,.., 27 ~. 3 Tabk 5..Me~b1'3$~d dara m d e d at the elo ow& BC, Mvniapal Atrport .'

Weathef Stanm - Amm~beric .Em ' r S e r u i c e , w & - ' A

periods of April and M a y l,%lf%!, and June 1984 -.---_--.-........,... 31

7 Table 7. A comparison of honey ke (Apb m e U i , h I;)' aad Omuo fig- . . propinqua Cresson +Amamam for ofchard poIIinatioo ..,.,...,.............. %

LIST OF FIGURES C i

- - -- --- - ~ -

--

1 - ' Fig L ; ?) The dimibution of resid dkatittn between -ti= species by their -between-y'*ear . . .

comawive years, 1984-1985 ( ........ ............... .*........ . Sorrnson 1948) --.-..LC..: .i...L.. .............. = .................................. 24

, .

2 Fig. 2 A . . /

of &ween-year %imiM;y' irr bee dsitation in fruit crops v e f s u s X f 3 1 i h found in three u, foix of the designated habitats. PJW ccxnmunis (pear) was not jncluded since no bee pdlinators were collected on . this fh i t aop in either 1984 or 1985 '(Sorenson'~ PresenceAbsence

- - .... ........................................................... .......... silnblity Ige& S03e~t~01f p43) w......... - .., *

1 3. capflne rates- of non-Banbur and, htnbus species in me Okanagm-

V d 5 y d& 1984 and 1985. Bloom period refeq-m the pmbined . ...... p o ~ o n period for cherries, pears and a b l e .f .................................................... . 1 .

. 4 'Fig 4 . R - & e o g v i a b i e s e c d p p u fruit and fWtWe@sfot-R& -=--

Delicious apples resulting fiom one of the following tTeatments in screened - - . endbsures: one 0. 2is;lruia (OL) visif one to three honey bee (HB) visits. Control A (blossam gusters thinned, and no pollination allowed) or Control B (blossom duqers thinned and unlimited pollination ailowed) ............................... 61 1 a 1

5 . ' ~ i g . 5. Relationshrp of 'number of viable seeds per fruit and fruit weight for Red Dezicious apples r d t i n g from one 'of the following, matmmk outdoors: one or two hwqy bee (HB) v&i&, Control A (blossom* dusters thinned and - . no pollination allowed) or Conml B (blossom clusters thinned. and unlimited

.................. pollination allowed) ............................... ................................................... 64 - - -- - -- - ppp -

6 Fig 6.' Regression analysa o f h a g e r r entering per colony and two colony chmcreristics, areas of adults and d e d brood, recorded while colonies

..-,I were located in ' cherries, pears or appkrXiiring ttie pollination period in .......... ........................ ......................... 2984 .,,-......-.,.,.,,..--.-.a L--..a , .........,..,...I. 79 .

-,- . - 7 Fig. 7. .&elation analyses of foragers entering p& colony and three wqather

conditions,' globaI sdar radiation (G.SR), relative humidity (RH.) arid . *

temperame, remrded whi l i colonies were iituatcd is cbgmes, pea.$ or -, . 4 ............. ....................... apples during the poiiinatidn period in 1984 =+ +. ~................ 80

< . <

* "

A 8 Fig. 8. Repessiw analyses of foragers exiting per colony a& pdkn bearm -

e11teTing pr=r colony. v e m s three colony characteristics, areas of adults, sealed .

brood and unsealed brood Data were recorded in 1985 while colonies were loated in an apple orchard. @can count per minute per colony was - *

Q -- ,...... ........*. . de-rived from a mtd af 18 we minute counts ........................ ..,.,...... 81 Ps 2

4- 8 > - r,

4

* Fig 5. Regression analyses of foragers exiting per colon). and pollen bearers

entering per &my v e f ~ t t ~ three coiony c~aracteristia, a . of acWts, ,&ale& \ - a - -

brood and g s d e d br& Data were recorded in 1985 me colonies were "

localeh in a chew orcbrcZ hfcar. wunt per minute per'colony - - - was derived . from &ta~ counts for 18 me- &nut@ @terZds ...................................................... 82

Fig. 10, Regression analyses of foragers exiting per colony md pollen bearers entering per colony versus three colony characteristics, areas of adults, sealed brood and unsealed b r a Data were recorded in 1985 while colonies were - located ,in a pear o~chard Mew &unt per minute per colony was deriv

............. . from to& counts for 18 one minute intervals L ...............................

11 Fig. 11. Correlation analyses of foragers exiting per colony, pollen bearers e n t e ~ g per colony and three weather amditions, global solar radiation (G.S.R.), relative humidity -(RH.), a@ temperantre. Data were recorded d m k g the

*pollination perid in 1985 white 10 adonies were Iwated in a cher;). -

or&& Mean counts per minute were- recorded for six repetitions per 1 -

colony redring in 60 data points fur each correlation ...-..,...................-................ 84

12 Fig. 12, ConelaQon analyses of foragers exrting per colony, pollen bearers entering per colony and three weather conditions, globaf solar radiation (G3.R).

* retative h+- W) and fsmpmme. Data were m r d e d during t6e pollination period in 1985 while 10 colonies were located in pears. Mean munrs per minute were recorded for six repetitions per colony resulting in 60' datar points per correlation ...,,.-., .......................... : ..--...-.. . . . . . . . . . . . . . . . . 85

Fig. 13_ConeIation analyks of foragers exiw per colony. pollen bearers entering per colony and three weather condition% giobal solar radiation (GS.R). relative humidity (RH) and temperature. Data were recorded during the- pollination-period irr ,1985 while 10 colonies were locared in an apple \orchard Mean counts per minute were recorded for six repetitions per

.............. colony resulting in &I data points per bonelation ...........,........<-...................... 86 e

> . . - - - -

,+'- - +

4ze

a i ~ ' s a g e s is &c

* occtu there are rwo major -5 rtquiremm~:

A

CMcGregor 1976). - - - - -

- - - - - - - -- - -- -- - - -

- < -c

Frrdr v;trieries which p5sxtwx~ m Q

.- aap with their awn arc vrmtd

seEkpmpau%k or seff-fmik. Vitrieties hi do m set an eamamc crop with &cir own

- - - - - - poUm viability: 3) regulan~ o f ' f i o ~ ~ g ; 4) ovnlap of p c h k e r flowering period relative to

- -- - -- - -- -- -- -- - - -

-

the &--vane%? Xb- % F a Fffiaenc'xhtiii- to fruit set (Horticula Ed-tion -

- - - -- ~%TI.

C .

commorr, and the popttfatiwt of wild bees d e a d , renting of honey bee cdonies for -- - - - - - - - - - - -- - - - - --- - - --

L'

@$tion began Cmendp, one million hone). bee denies are'rented annually in the

Urnled Sdks for tbe pdlinalion of agrictdmil oops (McGregor 1976). In 1983. 6465 hives e.

I - f 4

- -

-- -

?

indigenous populaliofis has hen limited In additian. information on the comparative - - - - -- - - - - --- - -- - - - -

- - -- -

pollination efficacib orp* bees and wild bee pollinators rehtive m fruit qll.lity .is d- . -

ne@&i%Tc -@E -bee&qasy -ve, refking honey bee 6nmapemenr ~~~s tems IO

'9

fwr major studies undertaken were to: a

1. the abundance d d i v m i q of wild bee pollinamrr in orchards and -- - u n ~ v a t e d habitats in the Okanagan Valley, and to identify wild bee pollinalqrs w i h

% 2. ,d-e the pollinatioo eflicadsr of an indigenous wild bee pollinator. 0. Irgnarrr .

\. i ' popinqu4 and h e y bets on Red Delicious apples; \.4 -- -

orchards using fwer mfrmx counts; and,

4. ~ v & g t t e A the biology and emnondcs of hL&e managemem 6 r pollination and, ,

deLtEr&ine the feasabili-r). of package brr andlor nudeus prodmion in conjunction with

ABUNDANCE

THE OE4.YAGAN VALLEY, COLUMBIA

- In w h - w e a r f s ) . pears (fynrs cmmunis). and sweet cherries

(Bums mMn] (with the exceptwn of rhe self fertile varieg Stella) to be pollinated, tiiere >

must be present in the orchard 1) pollenizer vAties which pmduce pollen capable of -

fertilihg and produdng fruit when &e pollen is mrried to the stigmas,of thk main tree .. fruit variety; and 2) an adequate supply of h e a pollinators in the form'of either managed

honey bees ar wiid !%es to wsure that &-p&imiod ocfurs. '. -- - - - - - - - - -- -/ - - - - - - -- - - - - -- -- - - --

R e pollinators are the prhaq agenrs for the transfer of pollen from one

seff-incompatible fruit cultivar to another. Wind andlor other animals are negligible factors in , & be pollieasen &&,---& pcars-fMcGreeor 2m:=ffrrwerffrrwer---

stramres -

found in qese fruits are dea& adapted for bee pollination, and the requirement for bees

has been shown by. many experiments in which negligible fruit set ocnus on trees screened

to exdude them, whereas it was satisfactory on adjacea plants to wbich bees had access

(CIaypool et al. 1931; Free 1W, I%;, Swphen 1958).

insect pollinator of agricultural crops, including: . . e -

1. Propgation - ecommidydfeasible management systems have been developed which are 6

conduave -to manipulations of colony pcpdations; a

- / /

,/I- , l- -&ty - honey bees are maintained in mlonies which can be uansponed when and

where they ar/,needed during the pollination period Also, honey bee workers cm -

reorient to new hive locations; -

' a g i v e time period for pollen andlor nectar. A h . honey bees are not oblipate visiton - - - - - --

te any @e plant mes, as arc m e &golectic wild bees;, -

4. Availability and Managemem by A ~ & - different crops have varying pdlinarion b

- . the irn$manx of aattiraliy ocaming populations of wild bees for the pollination of fruit and

f crher crops has also bees well documented In this paper, wild bees are defined as all

f

members of the superfamily Apoidea other than the genus Apis: Satisfactory to excellent fruit

- - set smmimes can be ztmmpkheb-in the absence of honey bees-(Fox-Wlson I929rmkenP -- ---

- *

1958). Brimin et al. (1933) determined that in pans of Nova Scotia -where there w& a '

virtual absence of both bumwe bees pnd honey k, the naturai populations 'of Hidictus and

The population levels of wild bee species have decreased substantially since the

aforementioned studiesbere completed,. primarily- due to changes. in agricultural practices which %

have resultad in extensive monwopped acreages, reduced plant diversity, increased use of

pestibder. mmpetihn by honey bees and dcswtion of nesting habitat (Free 1980;. Johansen

Don and Marten 1966; M- and Moulter 1977; Johansen and Shawa 1974; Morgan and

Perdval 1967: Cruden 1$% Heinrich 1975, 1976). Reductd populations and annual fluctuations - -

in abundance patterns have made mmmaged wild bee species unreliable and unpredictable

pollinators for'agriculoual crop$, neccesirating &e rental of honey bee colonib for crop t

pobatim: However. a nm,ber of wild bee species have proven to be more efficient 4

polhators of certain crops than honey bees. For example, the alkali bee, Nanio mlanderi

1972: Menke i952. 1954; Stephen 1x5; Svphen and Evans 1%0; ~ o b d s 1%4, 1967; + L

Klostermeyer 1964) and in many areas of the -US. and Canada they are considered the - -- - - -

efhiem-golltnarors of f E i 3 ~ ~ & nut aaps. Osmia l igwia propinqua qesson is a North

American spies in th; fioal stages of -devel&pment as a pollihalor of almond (Tordiio 1979. - -

1981% 1981b. 1982a) and apple crops (Torchio 1W6. 1982b. 1984, 1986). Osmio canifcofis

fRadoszBowski) has been developed as a commercial pollinator of fruit trees in 'Japan (Maeta

1978; Maeta and Kitamura 1%5a' 1%5b, 1974. 1981) and has been imported to the eastern ' .>

fiom Spain and is bemg tested as a potential pollinator of orchard crops such .as apples

(Torchio and Asensio 1985). *

crops. For example, Bmbw tenemis is being developed as a pollinator of kiwifruit in New \

Zealand (N. Pomeroyl, 'personat communication). Bumble bees are important pollinators of red.

clover in northern Europe and efforts are made, to grow &us crop for seed i n areas where _

bumble be? are abwdant (Free 1980). B& spp. were imported for red clover pollination - in New Zealazld (MtGregor 1976).

-- - - -

If we are to benefit from the pollinating activities of wild bee spedes it is essential

that we begb to augment and mainrain their. populations. Before management programs for .

wild bees can be' developed it is neceaar)- to &ey bee fauna in a pantcular area, not

only tr, determine the relarive abundance and species, but also to

determine candidacy of panicular species as naniral and d e e a &Aollimtors of target crops. 3 I

The objectives of this re&&& were ro:

%

2. Examine whether mcient n u m h of wild bees rage in the orchard' habitats to k

lktemine the most abundant (dominant) wild bee species which forage in o&d

habitats and their potential for domestication Ad management for mm&rcial m e fnrit

* i I t + =

a : . $"' .-

to early June 1984 i d mid-Apa to the end of May 1985. Founeen collection sites - <

(Appendix 1) were used representing four habitk types: 1) Orchards: Far from Natural

,(OFN) - Sites were located in an orchard surrwnded on all sides 'by other orchards.;' 2)

Orchards: Near NaDaal (ONN) - sites were locat@ in an o r c h d . which was bounded on 6

one or two sides by natlrral, UllCUltivated land; 3) U*tivated: Near- Orchards (UNO) - - - - - -- - - - - - - - - - - - ,- -- -

~1% were un&tivated land bounded on one -q two sides by orchards: and 4) Uncultivated: - -

IC

Far fiom Orchards (UFO) - Sites were d t i v a t e d land completely surrounded by naml

habitat and > 0.5 km from the nearest orchard The sites were loated from ~ummbrland to - - - -- -- - -- -- - - - - - - -- - -- - - -

. - -- - - - . -- - -

Oyama and the same w~~ectioo sites were used during both years of 'tbe study:

1

The Okanagan Valley has a xeric, cold temperate climate with an annual rainfall of . 7 ,

38.0 to 51.0 cm All sites in t& uncultivated habitat dJplations are slaaificd as 'ponderosa

pirebunchgrass" (Krajina 1969; h y s h a w 1970). The crops represented in .thi orchard habitats

orchard which was designated UFO. Cdlection at =chard sites was coincidental with fruit

crop and understory bloom su'& as Tmctracum @ciincrle (dandelion). Uncultivated sites were

rampled for the entire &week period Bee poltinatms were aollected for 1 h intervals from

0900 to 1MO h at each site to allow $thin and between-habilai @sons. Only wild '

C - bees foraging on blossoms were aught; hone)- bees were not cotlami

,. I

D m S F I Y : To determine d ive r s i~ of wild bee pollinators on an orchard crop or aatural

and cross-referenced with ahe bee c~ptwcd while foraging on them The bees were killed ,

Griswofd. Bee Biology and Systematics Laboram, Utab State University. E&m, Utah I

(Megachilidae); Dr. W. E LaBerge. State N a n d History Survey Divisioq Champ~gn, IUinop

.(Andrenidae, Megachiiidae=MeWes species); Dr. G. C. Eickwort, De$t of Entomology,

University of. California, Berkdey, (Anthophoridae = cerati/ speaes); DL R W. , - - r - --- - - - - - -

Brooks. D e p ~ of EntomologyT Uni Lamence, Kansas {Anthophridae).

Veda specimens have been r&ned by all of the abpve taxonomists and at Simon Fraser

University. In 1985, aII wild pdlinators were identified to species at Simon Fraser University. - - - - -- - - - -- - - - - -

-- -- ----- - - - - - - - - - --

In 1984 and 1985, '& bumble bee spddmens (Banbur spp.) and plants were identified to

specks at Stmoa Fraser University. Shamowwiener Diversi'iy Indices were calculated from

these &ta fMargalef 1958). Bemeyear sirmlarity in for;tging patterns was determined using

S u m ' s Presence-Absenw Similarity Index (smensm 1948). Analyses of variance, were wed

to analyze within-year family and habitat cemparisoIls '

ASWANCE: Coifection methods have tkm desaibed in the previous section Collectors -

waIked transem through eacfi site collecting wild bees foraging on* plants in close proximity

to their path. There are a numkr of problems with this collection technique which could -

result in biases in the cztpnue rate data: 1) jlIctim collectors may have differed in the

vigor with which they surveyed dectim sites; 2 ) &Terence in ropography between sires >

could have made it m e difkult TO f d l a in m e sites compared to others; and 3)

would l-ikelq. have ccmmd at all @ection sites &d therefore. 1 believe that the resultant

capturq rate data is representative of wild bee abundance in the Okanagan Valley. Abundance

( h n t . upture rates were z , . - ,

dererrmn ' ed f a each habitat A wosampk t-test was used to compare toral capture rates of '?I g - -* + -

each habitat between years '

All- H d bees (Hymenopm: Apoidea)*collected were'in the families Apidae, Haliqidae,

Andrenidae, Megachilidae, Anthophoridae, and daC@letidae (Table 1). The f d q 0kidaej" '2- E V

. - ~eIIiti&e and Fideliidae were nof represented ik ,these mllmionr i

$ R -

*& than the' orchard h%tats (OFN a6d ONN) in both ye& -(iabie 1). although @ ; *

1985 the diffmmrs arm nor as dramkc as in 1984. In 1984. a particularly low dikq - - -

--p-pp----p- - - m r ~ - f w t l r e - f w m T 3 - - ~ ~ d a e , A . ~ ~ , M e g a c h i l i & e and'

Anthophoridae showed higher d i v e @ in Marluvated habitats compared to orchard

botfi yeam In cantma, the Apidr# in 1984 h&i lower diversity in ONN and UNO '

than in OFN and UFO. In 1985. @a divqit)' for the Apidae was low in the OFN

habitats and high in the other thee habir~n (Table 1). Only two individual Cdlctidae (one ZI

species) were collected, both in the UNO habitar in 1985. - '

- -- - - b

The higher sped= diversity on naanal vegetation in both yeas was due primarily .to 4

the gream number of 5 # 3 n - - l 3 m h species odlated in the wmFtivated habitats in both

y e a n Seventy-nine maBcmbvr sped& were d e u d in the lmcultivated haxtats and only . -

the urmltivared habitas Dwpvcd to 27 at the orchard sites (Table 3). A total of 100 -. 3

wild bee W a c id;nt%ed dvnag 1984 and 1985.

+- Species 0 U a ' 6)

3 u

4

No. Individuals

* Diversity

Indelt

Nor Spe c ies-

c a No. Individuals 3 Frr

E 0 * - U)

'&

4 -a 5 ' ~ i v e r s i t ~ ' u al k -----lnde$

E: ms U

No. Species - -

aJ d c 3 7 k .

U a ' No. Individuals-

r( a Diuersity - k .- 3 Index

m 3 a No. species h z (d As

2Q-g *

Diversity .. Index m e

No. Species a cu --

:NO. I n d i v i d u a l s 3 L 2

m i > u + L No. S p e c i e s u 0

1 - o D i v e r s i t y rrJ z ' Index

m E z e No. S p e c i e s

E+ Species

- W - - - - - -

9 D i v e r s i t y a b u 3 Index

3 2 L ,KO. G I n d i v i d u a l s 5 z

s 0 4 - ah. Ind iv idua l s

I n d i v i d u a l s -

t

Divers i ty . .n Index -+

0

d - - * G

i %,Different 5

ri

h - Species 1 1 d No. Indiv iduals

a: E " Divers i ty Q U a z.1 Index u a i

k 5 No .Species i u b f 4 i 2 ~ . Ind iv idua l s 5 C4 f r

- r " " Diversity j a u a L u m Index i

5 h f i

ZEo. I n d i v i d u a l s z z

i c

4 D i v e r s i t y j 4 T 1 L .. 3

i

L 3 Y a Diversity

.. a Index

D i v e r s e IC 4

Index 4 it Tu.DiffeTeut -

0 F Species

So. I n d i v i d u a l s

.. 2 -a U

Divers i ty aJ u 3

Index

a $No. I n d i v i d u a l s !z 2 Cr.

- 4 2 b Q =So. I n d i v i d u a l s C s i Z

- d ci Diyers i t y z Z

1 ndex

r - ; 30. I n d i v i d u a l s - 4 L

D i v e r s i t y 'JY I n d e x

No.Diff erent Specf es

a No. ~ n d i v i d u a i :

cR ' -0 h m

. D i v e r s i t y .

u G f ndex a ' No-Species + 0' L l k 4 - ;L

2 :NO. I n d i v i d u a l s 3 b.4

.. m -o 'Et D i v e r s i t y aJ L u a I n d e x 9 5 -Xo.Species u 0

. -

4 D i v e r s i t v

u a NO. I n d i v i d u a l s 0 Z

2 u D i v e r s i t y a Z

E

2 $NO. i n d i v i d u a l s c &a

Divers i ty V: d

Index 5 u No,Different 0 €4 Species

Ho. I n d i v i d u a l :

.. s ,, Diversity 01 L Index

- 9 VI o 'ci D i v e r s i t y Q) u LU Index

. D i v e r s i t y - 1 b nd ex

D i v e r s i t y u (3 Z Ihdex

2 No. I n d i v i d u a l s -3

4 u No. Species 0

, , e

No. Individuals

- m . Diversity m

+- s Index

G - k r: a No. Individuals 3 c4

Index

No. Species

3 &l

2 2 No. Individuals 3 z

Diversity Index

* * 3 u No. Species

'a G L Z

2 '$ No. Indiv jduals 0 2 .

4 ' a Diversity L 3 U

Index

a. z " * E 3o .Spec i . e~

- 0 L I L .

5 ' -

Diversi ty Index -

L

yt - f - - - - -- C -

a U 0

No. Species B

No. Individuals

n a Diversi ty b . a Index

u L %I rU 5 u M c a No. Individuals

=s Fri

r3 3 > u N o . Species 4 b 11 0 7 t 5 h 3 = c E; 240. Individuals 3 2

Diversity Index

.* - d a u No. Species a a

V Q Z

cb - s k

r -- DCY er sirp

Index V) - -- - ~-*Effemr? u 0 r S p l c i e s H

No. f n d i v i d u a l s

CI1

-2 at

" Diversity '0 U a h \ u 0 Index

3 2 N o . ~ n d i v i d u a l s I Fr

> .u No. Species

4 Diversity a

-- Index .. 3

m Lt u No. S p e c i e s LI 2 ar

2 :NO. Ind iv idua l s 0 Z

3 u Diversity m z 1 ndex

m E . a No. Spec i e s L. L Q C +-. U u u a No. Ind iv idua l s CI I v e4

L

p h t spedes, 7 more than in 1984, were used as forage. An increase in the number of

4

no wild bees were collected on pears Native fiw. such as BdsmmAiza SOgitroto (spring

sunflower) and T. o_Sricide, bad higher t d diversiv of bees foraging on them thm any

the orchard crops in both years (Tables 2 and 3). T. offCcinale was 'particularly inmesting . . since it is an understory vegetatim in orchards and was found ar all habitat designations

dlrring - bofh y-, except ONN in 1984. The remainder of the native flora listed in Table 2 / ri

preseacrabwnce dmilariq. index (Sorenscm 1948) revealed Ulat man sped= of. bees (741%)

portion of plant species (76%) were visited 'by a highly variable forager population, also

determine the dmitarity of Of bee fforaging patterns on orchard sups and two of the *

dominant native plants for 1%4 and 1985 (Fig 2). P p s c m m w i r was not included in this

analysis since no wild k pollinawn were d e c t e d on this nop in eiihervcar. Wild

were more variable in &tir w e of orchard rwurces (lor. similari~ values) tban of

T. o@cide and 3. shgifram (hi@

F

n. Abundance of & species -

(dominant> sped= at the orciurd habitats during 1%4

Fig 1. 'a) The dimiiuticm of resident bak species by their simiiarity in flower itfikation . bttwttn tmsxmive years, 1984-1985: b) tite distnhttion of flower sped= by the&

between-year sin&&& in bee spedts vi9'ting them dmh& ~onsecutive years, 1984-1985 (Sorcnson's Presence-Abse~lce S-ty -ex, Sorel3s011 lw).

Ik FLOWERS VISITED

b) BETWEEN - YEAR SiMILARtTY IN BEE VtStTATlON

pnunaars. were most common at 7be UFO designation. h 1985. B. t . occidentalis and B. b. - - - - - - - - - -

--- -- -

nearctinu i& e spe&& ~-ectedbr~&ne speciorchard habitats, while H. - 4 - zag^ aod v ~~ were again dominant in the uncultivated habitats. A l i t of wild b - bee poBinauns and their abmdance in the habitats in which they were colected in 1984 b d

- - . w-.

- 1985 is given in Appendix fl.

One useful factor in demmining the potential value of the dominant wild bee species &

- as pollh&rs on tree huit crops is their preference within #ofchard habitats for understory

d ' - -

--

--

vegna&m or crop-neer(Appendix IIr Of XI the - a d bees. Z K a - - 6 - t h Z - oi&rddhabitai, -

75% of 8. r. occidentdis, 43%-of B. b. nearcticus, amd 54% of H. cineraria were captured

while foraging "on apples, sweet cherries and crabappl& Althougb none of the other bee

collccrcd qvhile foraging on these crops. The Iialictidae' displayed a m o n i preference for - I

'2 \ Lnderstory -plants with only 14% of all individuals capmed . while foraging on the *get fruit

crops. The Anthophoridae dm prefered undersmy plants with the exception of 44 H. - cinerana collected on orchard crops in 1984. No Colletidae were mllected in orchard habitats,

/' uncultivated habjtals during botb 19&1 and 1985 (Appendix U). Of all the individuals

colIene& 48% of H. cwt$usu, of Dialictur sp. 2, 70% of Dialictur sp. 5 , 76% of ""si =

, Dtolrctur &. 6. 63% of D. him&, and 6% b D. pnrinmur were caprured while

fordging -on 7. o @ d e . In 1984 and 1985, 87% and 78% r d v e l y o f all wild bees .-

mai capture rates in rmultivaed vdorchard habitan (Table 4). lo addition, the Y O

Ta

ble

4

, W

un c

ap

ture

ra

tes

(bce

s/h

) il

d h

er!

fm

\ 1 i

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ulti

vate

d O

kana

gan

Va

lle

y h

uri

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19

84

., 1

98

4 M

aan

cap

ture

ra

tes

(Ile

ca/b

)

lu

I 1

98

5 M

ean

ca

ptu

re r

ntc

s 1

8kes

lh)

' ,

'4

I *

, I

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s :

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urd

' 2

.52

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llo

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51

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w

f 01

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: F

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1___5____

6

designation had a tignipcantly 'higher capture rate than the UFO d-tion.

fmlres. 1% -

can be accounted for by the faa that a large number (n=44) of H. cineraria -

were ai one of fotn sites in this habitat designation. The Apidae had the next

highest capmre rate in Om, although it was not significantly different from the .Halictidae,

Aneenidae and GSletidae. In ONE;, the Apidae had significantly higher capture mtes than

rhe othn familia (Table 4). The Anthophoridae were scarce in this habht designation

Hafictidac h d a si@fimn~& higher capture rate than any of the otber families.

- In 1984, capture raws for the Apidae fluemated the least over the 4 habitats compared

*

in 1985. total m p w e rates uf rhe two tmculrivated hawts were an;lin significantly *

higher than both of the orchard desigx~tiom (Table 4). Within the orchard designation, the

totd capnne raq for O?$N, was signifivtIp bigher than OFN.

- In OFN and O W in 1985, the ~ p i h a e had dpnificantly higher capture .rates & a ~ any -

compared to 1984 ,and was nor bramaucdly different from the capture rare in ONN. In A

UNO, the famiiy Halicddae had rhr highat rdpm rate although it was no1 /&nifi8ntly

was spifwtt) h~&r h~ f& h e oxha d w . The higher -capmrt ram of Wctidae in F .the ur~Atiwted habitas in 19% m e similar although not as dramatic as those in 1984. As

Colletidae were higher in 1985 than 1984. - - - - -- ---- -

- - -

-- -

6

Abundance patterns for Banbur species (bumble bees) were lower but more constag - *- - - - -- - - -- --

-

than for non-Banbtrs species (all wild bee spedes excluding bumble bees) duriug the

cobation per&% 1984 and 1985 {Fig 3). Tbe~e was a gradual increase in the abundance of I B m h s spec55 and n o n - B m h species towards June in b t h years In 1984 and 1985, there

was a redimion in the abmdmce of non-Bmbus Wes during the combined blooming

period for apples, chenies, and pars (15 April to 21 May). Since the collection period in

peak * non- bus Fpeda abundance in mibApri1 1985 would have been eaauntercil in

abundyce -apparent in_ _the fm -we& a f June l9&L wuld Jhave-axumd ia -wPlrtkPr-- +

conditions in Ap:ii and May 19% were warmer and drier than in the prev~om year, while

. -,- conditions in June 1984 w m d e r and piener than normal' (Table 5). The substantial

reductions in non-E~xbus speds abtmdmx chine the middle of the bImm perid In I984

and 1985 and during Ehe ftm week in' June 1984, were not associated with umeam&d

data It appears that Bombus were nor negatively influexed by the tJacltmtnl weather

near the end of April 1983, since capture ram were comparable in- bofi years

Ndley -

cherries.

/ ..--.- NON - Bombus sp.

I985 DATA

BLOOM PERIOD

Table 5. Meteorlogical data recorded at the Kelowna, B.C., Hunicipal Airport Weather Station - Atmospheric Environment Service, during the collection periods of April'and May 1984-1985 and June 1984.

Total Precipitation -

mm

April 1985 A

April Average' 14.9

i%y 1985 2 L 9 5 - 4 13.7 29.2

M a y Average 20.3 4.2 12.2 2 8 + 0

June 1984

June Average r

Year

h a b i t a t between y e a r s .

T o t a l Cap tu re Rates, x number of Bees/h 2 S . E . - - - - -- -- - - -

-

i

Urchards: Far Orchards: Uncu l t iva t ed : from - L% t u r d Xear Xa tura l Xear Orchards

Uncu l t i va t ed : Far from Orchards

T - t e s t p r o b a b i l i t y -

between y e a r s >0.05

The dominaat wild bee species in both orchard habitats were the same in 1W and

1985. B& r . mcidentrrlis, 3. b. nearcfim~s and Hapropad4 dwr& B. t. mcidentalis and

B. 6. nemaim are both gotentid fruit crop pollinaton, while the value of H. cineraria has

blossoms. This species has been sumssfully managed as an apple pollinator flmchio 1976,

The dominant a d bee s p i e s c o l I e d in the uncultivated habitats were l-klictidae.

and all species exhibited a preference for fmgbg on T. ofiUnale, Of the dominant wild

the orchard habitats The swcqribilir). of these smli wild bees t~ pesticides (Johartsen 1972;

orchard vqetation, T. of i c ide , reduces their value as pollinam of orchard craps in the

Pears are genemy not &&red to be as amactive u, honey and wild ,hew as other

A higher diversit). and abundance of wild bee pollinators were found in uncultivated - - - - - - - - - - -- --- - ----

I-

habitars - s the ct%thad h a = - ~ e ~ ~ ~ u v ~ ~ I O W w i l d pqwhtions on apples,

- I ~.-*-Uoll~ in both abundance and diversity

sops in the Okanagan Valley. These pattern$ of pollioator d i m i on W t i v a ~ e d and

a c k d kbitars suggest that pesticide impact, corngetition with managed honey bees, and nest

l%e low diversi~ of wild bees in or-d habitats when compared with micultivated . habitats iD the Okanagao V&q was parri*cubrly miking in 1984, when 9 rpoa of wild

The higher dive&& of wild bee @inam in uncultivated hahitars was primarily due to the i

79 nm-BoPnbrrs species cdleaeb, as 'campared to only 12 non-Bombus spedcs in Lht orchard

habitats. fn 1985. 64 species of wild bees mt coltecred in uncultivated habitats an4 35 .

species in orchard habitats. Fib ~3n- lkm.h~ species were cdlecred the d t i v a t c t f #

3 habitats, c~myared ro 27 ~~ sped& in the orchard habitats. Thcse results are similar

to those obuined in 2 study in the Fraser Valley of B. C. on berry crops (MacKenzie and

with only 9 noo-~&ur sped= (only 2% of bm mlltncd) oo cultivated bluebeny. raspberry ?

t

and cranberry. ,

Human effecu on and mar Mdulonai sos& are alro rcrpomitle fur reducing the

/. ---. -

\ + kes ro paUbda (lohanun 1972; Rowright and Thaler 1978) may aawnt in pan for .the

/' . .

of wild bees in orchard babi~ts, because mcs& orchardists rent hmq bee colonies during the

biooming for gdhatioa These pollination units are recmmnended at the rate- of 3 to

honey bees were not cafntlared in this &. it was evident that more honey bees than wild

_bees -yere plrwnk &I Lhc_orrharb h a w h m g the b l o o m - q - - w k - - - -

the hypothesis that competihon wi& honey b e s reduces wild pdltnatar populaticms (Eickwort

ag~&wiil practices rtsutting in i u c m d fi&d site and x.mxm-&= operatiom have

absorbed S U I T O U I ] ~ ~ ~ ~ natrrtaf habitat and d e a d t8e amom of laad avaliable for wild bee

Valley have probably redwed nestkg sitw and therefore wild bee poptjlatiom

a

frm during !idI b l m in behies and pink bud 'we in apples kiq _have acamwd for O . < --.

4 - popkition f imt iom in the ONlt' and UNO habitats.. Available forage for honey and

h r *

wiJd kes in orchards .w&d have been slLbaanliallg reduceb-%inqe a b e e permi& of the

have k m s e d , possibly fox& bea to forage on the bardicrxnnive floh in adjacent

uwltivnted hatitars Tbb change & habitat muld have aocwnrcd for the significantly lower --

_t,-

abmdam oTpwiId kCI h OKN =-l984==eded to -1385 and L e si&%&%higher

% w=-empm%xerirr- - .

Michma i979)*-&b =*s- berwan- --- .e

5

-- - pamirdand maybe h i g b l y f v c tfl wild & -f-m 0 t - * , -- . -

.I

* ~ u r t h c k e , the d2 pmximirf i? large patctia of carfy & furage. as rcpmmid by

. - . -. E . - * ; I .

. , ~ - . - - - - - - -

f ~-

-6' C- : - - - --

* ,. - - 0 . . ' b r ; 4 - -

C - v , - - . _ _ 1

- - - - ~- , ~- . %j -

i ---: -

. - J R . . 5 . . ... . ;

apples. mmim m- aa~mpp~es, would * . % . b e . . :UNO ~- -

. . * - - - ~- + - ~ -1- - , ~ - - ---. ----A - ~--->-- L--

h . < -: -. .' . _ b p i - - ----7 nm~ -=-- we7tha~-efpergepMY io the :;Thael COIIC]IISim We

, .. 8 - * - :" : 9 ' - ,_ . I , 1 r -..-:. - . i . I - +

ch Wicaehi&a Smtf.-and. *. I .

. r : r

f , t . . . . .of wild . -, . - 1 . l : ' - ., .:-: \ I 8 * : .' .* * ,

b in and $&. *me.:* -i4743: '&$. M ~ p s l & ! i ~ s p m i v d ~ ) &pite 'the - ~ . . '1 p - . b

3 . 1 ' ' !

.,. .~ 2 .! ' @ prrsnvt of'ww w'& , - '(rde s - :).',B ~, . &;d&ti&s& 3 dif l idt

' 1 \ ' " . . r

, . ~ ~ : i * - .: . ' . .

% ' .

to avrmin which year rcjjrt$&rs the - w h ' ; - o r % .. "&erageW &ti& ip. -ty, .but . . . ~

. : I . 1 * ! . . 1 . t

1984 aar Ekel$j wne miboolm;l' . . than @~'(agw bf ,mm,d w&&& cbnditiions ; -- -~ , _- ~ . _ LA-- -_ - i . r i .- .. . c~xfunftrd - - . A d . . .- . i: -. . , . . " . - -

' * . ' = . . i . , ? . *

I

1- . . valley. lnclement weather - . seems to affect activity

of Babut spedes less rhac non-Banbrrs sped6 sin= the fomrer are' better able to regulate

. their M y taperawes and are therefore capable of survjving and foraping in conditions

8' 0 * I (fig- w b l ~ re@=tingh a ef c m ~ ~ &y have brm delayed by t$e rh&l - - -- -- - -- -- ------ 9 -- -

- - --*- 4 - 7 - 7 7 - . ./

c1 reatber mdditiom enm@pd -i+r the end of April i d a d d -d-m .the pq&--inh%-- .- -7- +

s 1 ' . 6 J

, . * s . 6 B - -specis abundage snmmed 14 tb 21 .April -1985. . . . .i a'

h * i - . .Bi & . s a

- -- LA- . T I - - - - --- - - - --- . L--

e i-; 2 -d pollination IA& , + . s - ,' . . f 1 -

A m 9 y.3 I

3

@itfaid et aZ. (1933), rmleaed"1356-18@ d d &A '& -a&'; arc&& during be -,-

' ~ a m pniod They dearmined that the avelage &ber .*of. &d bKs taken @ all 0 .-

smaller population of wild bees - fmagine .in habitats in the 0L;anagan Valley. During

1984. capme rates in OFN 'were i376 bees&, aMt 2.36 .bees/hr in the ONN (Table 4). "In - -

= 1985. capme rates in O M &&e 2.52 bees/hr. and 5.82 beeslhr in' ONN (Table 4). , . I ) ' r

. According to I3rittain et al. (1933), the 'number of wild bees present in+ my stud9 would not . j

- - . . W - h e r e is a mcieaiy ab&t poplation o f wild bees 3 a r r a t u r a j ~h~ra; amamding. . - , . .

,*. . . . s .

* . s

-. * . . , . . * , * ~ - . . - - - & .

- -- - - ~~ . . .:% ,

- . . . - - _ " - - - . ._ - - . -

i . , . 0. - , .. .. 3

. .

0 ' . * - - . . + ' . ' ?

4 0

- . I - '. > 38 1 . r '. . _ ' ,

. *

. - * . ,. - - - w. ' .

&e m e n d e d for adequare mdf set to mcur ( ~ f G r e ~ o r 1976; Philp 1930,,.b47; ~teph& ' ,f

-

-1961). (3ipidering this fec~mmendstion and Brimin et d's (133) suggestion (that 13.56-18.06 \+

b / h is equivalent to one hooey bee colony per 0.41 ha of orchard, it is that there t - i -..-

G

are insuffn=imt numbers of wildi bees in the Okamigan Valley orchards to .pollinate apples sr-

and cherri& (Table q. Furthermore, the number of hives r&mmended for pollination- of

- -- - - -- -

pzs is- double rhat-of apptes a& &ma {Comer Ti d.- - -

that even the

number of wild bees availabk in uncultivated habitats of the 0- VaHey would .be '

A: . . .insuffiaent for pear p o ~ m

Studies in the Fraser Valley of B. C , have detemined that wild bees are not present

in sufficient numbers to pdllnsl~ commercial blueberry, cranberry andd raspberry aop~

-. (Winston and Graf 1982). The average capture rate of wild k e s in blueberries, cranberries

,- - C

and raspbemes were 208-1276 bees/b 53310.89 bees5 and 1.2ftZIO bees/h repxtively.

All were well_'below the reoommeaded levels of pollinators n& to adequately potlinate - %

. t -&-- a f967- 9 M d q m 1976 -

MacKqne and W W n (1984) found capture raws of wilh beer in 1987 to be significantly

higher than in 1981 (Winsurn and Graf 1983). population levels were s@l well- below t i m e . recommended for adwuate pollination of the bezrq. crops.

strong tendency toward foraging on orchard crops does have potentid for fruit crop - - - - - - - ---- - - -

pollmation. -'rtarrbTatmcdk -&entlzfis a n & ~ ~ 3 @ @ F i ~ ~ e ~ ~ ~ t E i ~ t w o o f t h e dominant

- -

~ w p a € a t i o ~ ' fiom ye& w ' y a regardless of weather conditions Mehods of rearing and .

management have been developed for several Bombus species and may be adaplable to

pllinatitw maat S)~S~~EIPS (Rowright and fay 1966: Morgan and Percivd 1967). In

addition, B& species possess several chaaeristics which are panicularl> suitable for uee I

2. Ehmble bees are Im mcqxible to paticides than the smatler wild bees (Johansen

, (

3. Although precise nudies on the foraging beha~or of Bombus sp8aes on h i t trees are

- ~ R T n w n ~ = M P n t c t l O C t \ - ~ ~ fly f-i

to nee more ream) than honey bees and aie thus potentially better cross-pollinaton. 1

In addition, Bombus species are -large and hairy, and make contact with the stigma on

most visits whether colIeamg poiien or- nertar.

/ ~ltho& Om& ltgnariD popinqua was no1 a dominant. spedes in orchard habitats it

1W6. 1982b. 1984, 1%). Additiooal informatian on the nesting behavior. &sting habitats and

annual population 'levels of 0. I. popinqua in the Okanagan Valley would be useful for

su~cessful adaption of the minagmmt &stem utilized in the westem U.S. 4

The remaining dominznr wild bee species, Halictus conficsus, Diolicius Ilaevissirmcs. D.

fruit polhators in this area because of inffequent foraging on orchard crops and reduced

. . - - -- - - ~

c 2 -------- ~- - - ~ ~ - -- -- us - - - I r

- ~ - - - - - - - ~-

~

----- - ~ ~

- - - - - - - - - - -

In an effon to Enaintain -indigenous populations of Bombus temcda occidentaZis, B. 1

bt- nemctieus and Omtia iignmicr propinqua in the Okanagan Valley for napnesting and -1

- used by these species for foraging neahg anb hibernation shMd Ix enantraged A n u m k . .

of measures could be taker: to provide addifionaf forage for managed dee pollinawrs,

, including honey bees and these wild bee <-The+ -far -d cregs k of

-shoq duration, therefore providing forage for managed bee pollinators for a restricted period J

of the s&m. fn addition, the reduced divers@ of nrltivated and native plant species in

areas of intensive orchard production remias the plants available as nectar and pollen sources 8 .

For €he remainder of the seaan. A h n a t i v e nectar and pollen sources could be made

available to managed bee p o b m r s by using d t i v a t e d land near orchards to maintain a

wntiTluous sequence of nectar and pollen-producing plants. Uncultivated habitas, in and

I

cycles of various fruit crops aPutd r e d € in retaining pplations of both managed wild

because b i t is considered t be a competitive rattrn than cosnplement;tn. vegeiation, attracting a

large number of foraging bees hom the fruit bloom (Corner n d. 1964). The 1 9 4 capture

rate of wild bees on' dandelion in orchard habitats was 10.57 b l h , whicb was higher than

the number of bees foraging w W target fruit crops (apples= 3.74 bets&; cherries= 3.64

i

alamative forage. thus avoiding rhe los of a subaaniLal number of bees. i n - r i i of this . mnclusioq ? ~ t & r e rates in orchard babims in 1985 indicated that dandelion (3.45 beeslh)

crdp kt) maintaining wild and hoqeg bees in -cis. The wild bec which I have

not exhiit a strong tenchxy mwapds f ~ _ o a ~ w-

* available, but will udize chis aifd plant as a l t e rnah forage.

a North American species now in the final sages of development as a pollinator of almond

(Torcfiio 1979, 1%4 1983b) and apple Gorchio 1976, 1982b 1984, 19%) crops, Since 0.

, . and W b l y a cheny pdllinato; io this i@m b feasible. A ampaxkm of honey bee and

intensive than honey bee maqanrm synam. Since 0. fipmia is an obligatory lmivdrine

speck~, -tion of ne3i.q popafat i~~~~ t imagbd thp summer is weccessar)., as is --

fee- and w t i m for ovmmering popuhtiax lo comparison LO honey bee colonies. 0.

u s g '" a d a m - % E z & l b 5 C b. i . 3 m 3 3 s ';--..-i

U r i a O b d

rl e b b

Honey kes cdlming pollen walk on the anthers and usually are effective in - - - - - - - -- --- --

- - ~ ~ n r t ~ - t h e + ~ 5 ~ - a r e ? l e c t a r w f f e r t D ~ - ~ ~ t f ; e i r p l o ~ s .

4 b=u=-m-f,y-.

ihey arc a major proMem in Rtd Deliciour rppks and effstivdy redua miit otr n e r e .x

are no side-kers in 0.' lig?uzri~ and both sexes amtact the sexual column of bl- to

The ten&rarure at which flight and foraging are initateti is vCry imponap~ espdally

- - -- in Lu; growiag r e s x i z - u - ~ ~ ~ - ~ u ~ a e y WW may experiem-re~ve~y opt-- - wmther dwing spring p o b t i o a Honey bee flight is initiated from ZOOC to 1 3 ' ~ . while 0.

pollen within hives facilitates cron-pdlination and ,alleviates the p~ent ia l ly denimeml effects

, - - -' to adequately pollioatc one hsrarr of apples '(247 treks/ha). Comparable @, ; - e

- - --

- It is evident that honey bees and 0. lignaricl possess -

tree nuuitioq nee spadng and pollenker placement have a 8 k in fruit set and '3

q m H , ~ m p h r y - B a k e r et dl. 1975). However, the efficiency with which pollen is uansfered !

- and depbnited on the stigmatic surface of a receptive blossom by bee pollinators issgene&ly - 1

comic$& to be of p h a q in fruit development (Free 190). unponaerR,\ ,- - - - -- - - - - - - ) - - - - - --- - - ---

- - Apple blossoms oomin of five stigmas which unite to form a single style lead& to -

- by McGregor 1976). Harmones pmked in the seed affect fndr growth, fruit set and general

hormonal balance of the nee (LuchKilI 1949; Westwood 1978). It is not necessary in apples

\ for all ovules to develop, aad fnn't may be produced with less than / a>hnplement of L.

'see& In many varieties, including Red Waws, low seed mun@rs are assodated with the

development of msshqm fntits, because ttte flesh cells of the fhit are preferentially

=ted in the region of the ovaries aw- fertilized ovules (S&ander 1955). Therefore, -

d

the shape of an apple is, to a large extent, dependent on the numkr of seeded versus

seedless carpels per fnrit ( ~ r ~ 1970), with -pen fruit having one or dore seedless

carpels In addition, the site and weight of fnrit q e influenced by seed number and' th_e - correhrion is genemy positive {Schander 1955; reviewed by Lane 1981).

multiple bee visits may be so thar 1) .an adequate amotmt uf W e , ampatible .

diffcrmt ke species. but wili ill in i n m i n i n g or improvint the m&ct value-of tree .,

'Ihe objectives of this andy were to: +

1; Compare the pollhati011 eflicacy of hoaey bar and 0. lignmh cm Red Deliuous apple

f i t set, weight and shape; and,

2. Evaluate the effect of multiple visas tq h e y beer to Red Delidous blomms with

-- ... - * ~ a g g k d to prevent pllkitioa from 4w. duriqg the entire experiment; and, 2) Crmrrol B: '

4+-

1

i blossom dustm were thinned as pieviop$y described but were howed unlimited pollination*,

' . throughout the experiment Due to probhms encountered d t h 0. lign- release and escape

,

from the scr&med m l y the one visit treatment by this bee was tested, reducing L- , : - &

the total numb& of txatme'nts 'in each screened enclosure to six ,

Prior to the opening of the kmg blossom$ bags, made of mosquito netting were placed - . .

over entire b b s and secured at both ends to prevent bee' pollinators from contasting

recepti,ve b l a s o m On 3 May 1%5, one 0. l i g d nesthg shelter (as described by Torchio, _.- --

1979) was placed in each of the screened &~sues . Shelters were ktangular, plywmd boxes - -

- -

(each 0.9. m wide, 0.61 m high, and 05 m deep) with one 0.9 m side open and facing

south. Nesting sheltes were supported 1.3 m above the ground surface by four wooden

sakes driven into tttevsoil and seed 'to the box.'

B .' - - . hdcs (8 mm diameter) W e d thou@ eath unit wire prepzed and placed iQto a~ opes

f i a of the 'snilk cuun Rqm mws (19.7 an 1- with a 7 mm inride diadur) were I .

insned into rhe milk cams thmgh the dnlied holes. N& nat boxes were &ced in each . .+ -

I of the nest shelters (225 n s hdeskhelter).

fio& Utahz on 118 March 1985. The turpid adults had been collected frw trap-nests A

the previous fa placed in individual clear, #000 gelatin capsules, and stored at a arnstant - 8C. This t e m p w e was mairrlained during transportion to Simon 'Fraser Universiry and -

folio& arrival. Rior to heir rekax, -d adults were placed at room -&p5rattsre .(21•‹c)- -

mhil they had emerged from rhe cocoon bur were still in the capsuit. These .capsules were

tk& rsooled to 4* c so that alj adults would _be at the mge-for- rekase--CkJ M a - - - - -- - - - ir

1!385* d u l e were left a kmm temperaaue for six hours and then removed fmm the capsules - - --

- -- far release into the- screened znd(1su1es The number of females and miles released in each -

shtpment of adults "was 364 and 34.2 percem for females aca - d e s respdvely. In &cb

- 3 9

screened endosure 6.6 nest hdes per female were made avdab e, which is slightly more

'than tbe number Tmchio (19391 reccmmenQ f4.6 nest Boles/imde). 0. lignaria were allowed

a 24 hr' reorientation period prior 10 the cnmneocement of pollinator visitatioqcounts to 1

. , ot bees -Mating and fcmping of C- fig& were observed following their release in screened

mctosuk Ho n a hiriaurn onnred during the entire polhnatioo .cod and the number of

individuals in the screened endrnura dmeased over the course of the experiment possibly /

due to escape under the screened panels, t C -

A theimhypquaph was p i a d in a Stevensons screen and positioned in' the orchard

Delicious apple .were alsb pkced in the emlosures. The bouquets provided a w e n source

for e r ~ ~ ~ p o W t i o r : since the honey bees and 0. lignaria were restricted to the enclosures a

and would wt have coatria with Mher p a W pollenizer varieties in the orchard. The

babapple - vhrieties uwd were Car). Hojer R15. Baaaa Rosthem R15. and Red Splendor # ' /

Rl5. Ikxiquets were replaced every 24 hr. Honey bees and 0. lignaria were observed f

P A sirnik experiment wrts d t b c t e d ourdoors in an orchard planred with standard

-- .- =tStodr Red DeIiciouxam&--euarieties, & - ~ u t l a n d , ~ - ~ - - ~ - .

approximately 25, years dd Four t rea~~~ents were tested on three ~ e d Delicious apple uees

including ow or two honey bee visirs and the same two control treatments used in the

screened enclosures, 0. lignorirC were nut dxmved in this experimen~ Limb bagging of

blossom dusters was completed on 12 May 1985. On 13 May Ia85, thme pollination colonies.

as dmibed in Pan C wcn ptaad in &c cwdurd. X~ese colonin were allowed a 24 hr

~reorientation period prior tp the -r of honey b e visitation counts. ... -

-

/ . outdoon. The -procedure inwkveb' 1) movd of the bgs to expose several bl-m clusters.

2) watching the king biossoms cmsan* mtif a predetermined number of either honey bees I

\ or 0. i i g m h bad visited ri ie receprise blosam 'and a m t a d the stigmatic swface., 3 )

fruit (McGregor 1976). ~ol ld i tor visit counting was completed in screened enclosures oh 15 - - - - -- - -

- -- h&yand on 17 May outdam. ~ ~ b e ~ ~ % f e e n - ~ ~ ~ ~ ~ ~ e s w e r e ~ - - t l & - ~ ~ ~ ~ ~ ~ .

Pollinator visit times (WT = flight time to receptive blossom + time k t in contact

with h e stigmatic d a c e ) were"Tecorded in the 'screened enc lmes and o u t d m for honey

bees and in the screened exbsures for 0. fignark P T were recorded between 1000 and -

.) 1500 hr while blossom visirs .were being recorded

-

On 28 and 29 Sepmnba 1985, tagged apples were picked from the trees used in the -

- -- - - - -- - - -- - - - - L-- - - - - --- -

xr& @ w e ~ and o m d q sntdief. This was approximately wen days prior to the *

T

average harvest date of Red Delidous apples in the Okanagan Valley. The tagged~apples

were vansported to Simm f;raser Vniversit)-. where they -ere weighed and halved horizontally - -

- - - -- - ---- - - -

- - -- - -- - - - - - -- - - -

-

f to e m ,the carpels. The number of seeds per carpel and the Wficat ion of the seeds

were de&ed Three utd &sGficatioos were established indudmg. 1) Viable Seeds -

endosperm gorged, seed mitt f@_v devejoped and length norm& 2) Non-Viable A Seeds - 3 .. errdosperm cdapsed, seed toat M y developed, and length normal; and 3) Aborted Seeds - unfenilired and length abnmmaUy reduced (0.1 - 0.3 rnm). Carpels mnrainiog only aborted

stimulate flesh cell growth. (Westwood 198 ) .

For the purposes'af .this paper, pollination efficacy will be defined as thc effect of

polbation on percent mdt sei. fndt weigh^ and hit shape. ~'mmparison of the relative

/ was complezed for screened e n d c q m and ou- In addition, pollinator visit times and the a

poUEnarion index (PI) were uwd u, delermined pollmadm effiecy of treatments The PI ( =

-

Data fm the screened endanne and outddor experimem were subjected to analysis of

and mean fruit weight d-----

&ysis siraoe M V A d h e d dhar mean f i r weight and mean numkr of W e seeds

per fruit were significantly differat from the other three new @ < 0.05) . Fnrit wei * - f and number of via& seeds pn h i t were wt Eignifcantly different for the rnmining ma.

so data m e pooled for analysis.

#

in mean fruit weight and mean n u m b of viable seeds per fruit @ < 0.05. Table 8).

However, significant d i f f ~ ~ (p < 0.05) in these c h m c s w& found between I

- ---

- and &.- &&&@ --@a&-+ -anb- - - -

--

A

Pnrrnt fruit set in smemd endonna was substantially higher than what is expected

under ~~ pllinarioo amditiom (Table 8% Apple' trees may shed 9f percent or more -of

their flowers and young fruits =harues (Wenwmd 1978). bur an economic yield Itom

an acre of m a m e amles results when approximately 5 p e m t frnit set occun (Hortinrltural

HB visit resulted in higher pezmi W*t ut (53%) than wo or three HB visits (43% and

- 41%. respectively). Two and three HB visit5 resulted in fruit sea cmpxable b Control B

The pollination hdcr (PI) for Control A Az1.43) was the lowest of any of the t

treannmri in screened en&&&. ~ h i l e Con& 3 had the highest (PI=285) (Table 81. One .

Table 8 . The effect of me t o three honey bee (HB) and one Osmia %i&a (OL) v i s i t on Red D e l i c i o u s apple f r u i t weight , f r u i t set, man seed set per f r u i t and mean seed s e t p e r bfossorn (PIa) i n screened enc losures .

-

- - - - - - - - --- - A - -*-PA- -- - x No. Viable -

No. of Seeds per Flowers in % F r u i t x F r u i t Weight F r u i t Blossoms

TreBtmnt Treatment Set ( g ) + s . E . ~ 5S.E. 0'1)

2 visits-HB 113 4 3 98.78+ 6.45 a 5.39k0.41 ab 2.32

3 visits-KB 4 1 108.035 6.91 a 5 . 3 1 a . 4 8 ab 2. 18 > 32 Contro l A 93.4311 6.10 a 4.47r0.46 ab 1.43 f

Control B 6 0 4 5 155.29+ 8.36 b 6.5620.48 b 2.95

k-' a P I = P o l l i n a t i o n Index der ived A x B

@ b Means w i t h i n a column fa l lowe&by the same l e t t e r a r e not s i g n i f i c a n t l y

, . d i f f e r e n t (Pc0.05 l e v e l , S tudent-Sewn-Keuls t e s t ) .

-In the screened eadarnues me OL visit resulted -,

more (p < 0.05) - - - - - - - -- - - - - - -- - - - - - - - -

- --

- Seedtess-carspefs per-frulf C Z ~ M ) I - ~ ~ ~ ~ Z E ~ I F ~ ~ J I - T ~ - ~ - ~ no significant differences (p - rn me UL mt. one to SF

visit The mean number Of see&& carpels per fruit was not significantly ctiff't @ <

0.05) for one ut three HB visits and C o n d A (Table 9).

- x

In the screened encloaue cxperimen~, dgn%cant positive currelations @ < 0.05) were

- - - - - - fa& rYffen &e- sumber-ef-viabkseak per Frrrit wercanrehmhvitk fnrirweigtn Ti one

OL visit (r=.693), one to three HB visits (r=.669; r=.623; r=.443. respectively) and Control

< 0;05) between the three nees used ia the study. Therefore, data were pooled for analhs.

Tfiere were no significant differences beween one and two HB visits or Cum01 A and B' in w

& fruit weight and mean. mrmber of viabie seeds per fruit @ < 0.05. Table 10). .= *

Penrnt fntit set was lower for Ccmmf A than for any of the other aeatments (7%) --

(Table 10). Two HI3 f i t s produced about the same fruit set (24%) ai m e HB visit (20%) r

2 ~ - The pollination index for Concfol A was lower than any of the other eatm me&

outdoors (PI=0.44) (Table 10). The pollination index for Control B was lower than either of - -

the HB visitation neatments (P1=0.79). Two KB visits resulted in a slightly higher polhation - index ( P I = 1 3 ) than one HB visit @l=&%).

t

Table 9. The effects of one to three honey bre (HB) and one Omia l&na& fOL) visit. on the mean nurrber oft seedless carpels

- -- per Red Delicious apple in screened enclosures <nd outdoors.

toea tion

x No. of Seedless

Screened enclosures f visit - ' OL 2:38 + 0.59 b

1 visit - HB 1.42 + 0.23 ab

/ , 3 v i s i t s - IB 1-49 2 0.26 ab

. Control A 1.00 + 0 . 3 2 ab

Control B . 6 7 +_ 0.19 a

2dt doors Z v i s i t - HB 1.56 2 0.69 a

2 v i s i t s - HB 1.22 2 0 . 4 3 a

Q?.ieans within a column followed by the same l e t t e r are not s ignif icantly different (P < 0.05 l e v e l , Student-Fewman-Keuls t e s t ) .

Fig 4. Reiationship of number of viaMe seeds per Wt and fruit weight for Red Delidous apples resuiting from one of the fdlawing mmmnts in screened encksum: one Osmia fignari4 (OL) visit one to three hclney bee (HB) visils, Conwl A (blc$som clustm thinaed and m pdlination allowed) or Cmmd B @lossom dlaas thinned and unlindted pollination allowed).

. 2 VISITS - HB .. ..

' 2. .. . . 2 ....- . . 2.. 2 . . . . . . .

r = .623 S.

CONTROL A

r = ,271 N.S.

0 I 1 1

30 60 90 120 150 180

3 VISITS - HB

I I I I I 1 0 40 80 120 160 200 H

CONTROL B 2 r2

FRUIT WEIGHT (g)

6 -

v i s i t s on Red Delicious apple on set pe r f r u i t and mean seed

The e f f e e t of oqe o r two hone; bee f r u i t weight, f r u i t set, mean seed

, get per po l l ina ted blossom (pia) outdoors.

Z N a - V i a b l e -

,No. of Seeds per LY - Flowers i n Z F r u i t Z Frui t , Weight F r u i t Blossoms Treatment . Treatment Set fg) ~ s . E . ~ 2S.E. (PI)

(A) (B) - - -~ ~ - - - ~ ~p

1 v i s i t 46 ' 20 100.64k 8.34 ab 4.78k1.16 a - 0.96

2 visits

Control A - 60 ' 7 116.07+10.88 ab 6.2520.85 a 0.44

a ~ ~ = ~ o l l i n a t i o n Index derived A x B

k a n s within a - f n ~ . ~ ~ - t h - , , , . d i f f e r e n t (Pc0.05 l e v e l , S-tudent-~ewman-~euls t e s t ) .

- - - -

, HB visit @=1.56).

In the outdoor experimenr, no significant conelations were found when the number of *

t J

viable seeds per Wr were correlated with h i t weight for any of the treatments @ < 0.05)

Honey bees in meed' encl-es spent &gnificantly less @ < 0.05) time searching for

- and collecting pollen from a receptive blossom than did 0. l i g w i a (PVT, x= 13.9 + 0.92 1

rs: P W , R = 47.6 + 566 sx, nrpecrively). Honey bees outdo& spent si&wtly leps (p -

- ,--I- - - - -- - - -- - - - -- -- -- -- --- - -- -

< 0.05') rime searching for and collecting poll? fmm a receptive 2hsorn than either honey------ --

a i ,i--

bees or 0. lignaria in screened enclosures (PW, z= 10.4 + 0.45 sec)

fn$t and fruit r i 5. of number of viable seeds per weight for Red Delicious

1 r = ,367

0 2 N.S.

I 1 I 1 70 85 100 115 130 145

0.

2 VISITS

.

r = ,437 N.S. .

I I I I , , lo 90 105 120 135 150 165

.

12

CONTROL B

8

2 5.5 1, , ,. , ; 1 - 3 4 z

r = ,394 . . N.S. r = ,224

0 N.S. 4.0 ..

80 93 106 119 132 145 40 60 80 100 120 140

; ,lo FRUIT WEIGHT (g)

Iig- >WL Alrhough the mean number of seedless carpels and viable seeds per Fruit for =&: -=-.

. . . .- . ~- - -- - L a I---~ -----A - -L---T. . '

-. . . orie . . 0. lig& visit . . k a e .. not significantly - .$ifffre-nt -~ -~~ from. me. two o r fhrge hongy~ bee visits,, .. . & U ' - rhne were subnantiallg &ikf empty carpels fewer viable seeds than in the other

4 - 0 e *

otrcatmertr+ (Table 8 and 9). Misshapened Fruit \ve p e or more Weq -Is (Free-970).

- ~

, - I ~

Fon~6ee -wrsrs --- a t ~ - -~ - - ~- --

I I

n 1 . & . A one *empty carpel. which also indicates a

.a f ' fe -l$b~evef, the r d t s for 0.- lignario suggest tipt there- is a' i

* . @ . * P f O p O ~ ~ t . e l y greatef ps@'bili.tf of this waking compared tb honey bees. As seeds deveop

t gibberellins (~estwood' 1978; . 1969) wbch are related to. fruit s~ a n d growth.

. t - . . , ' .. . . .

' +produce more severely misshapened fnrir . . . - . * . b

- .

*

c~d&ure, 0. &nuria spent 'a ' s i p d i i t f y l o n g & - h e =dm@' for and Coltgcting . *

: t. . , * -. ' . --. -' .. , <* If fiqn receptive M v ihhn did hone;- bees. beersd~srsul t , in combination- with a .s@stqgtially ,

- -

- There were no signifikanf differences betweed a ~ p of the fTuil quality charaaeristics - B

- - - - - - - - -- - -- --

-- measured for the &&*virji - r r ~ a x i i ~ t s L - ~ ~ m & * ~ ~ ~ ~ ~ - -

at .an i n p ~ a l of &e or two days on aver&e doubled the 'wed set. per pollinated Mossom

(PI) as -pared to polhating once. The results for one and two honey bee visits in the

-

screened endosure and outdoor experiments do not support the fmding of Visser and --

- -

-

Verbaegh (19801, possibly beguv the intervalt between the first and subsequent poUinati?n

visitr were less than four hours\ at t"emperannes ranging from 9OC to 19S•‹Ci Visser and -L

w h a the &a1 between polIinatiom was long enough (48 hr) at low (approximately nl~•‹C) ' .

/

or &G; *mgh'it\hr) at h@ @pproximat& 20' C) temperatures. With shorter or longer - - - - -- - - - - - - - - - -

- - J, - L-- - - - - - -

&e;valr. the cant?ibutim- of the second @Uen to seed production Qirninidys. -. . -

5

7

~ o ~ t o ~ visit times for' honey bee$ were significantly shoner outdoors .than in screened

- endosurs, possibly' bpcause: 1) honey outd&rs more -efficient at weking and -

f m a a g on receptive blossoms than honey bees restricted- %to7 weened enclosures and/or 2) Q

the high beefilossom ratio @ screened enclosures resulted in increased c~perirjon for /

- - -*- r-

individual blossoms and comequentiy a longer timi spent searching fw amrnive 1 , , , " . *

' . - + a

fl, Treatment-Control Interactions . i . a , . *,.

- .

requirement for bee poilinators * a le poll inafi~ Control B" .uar designed to lest the e n k ~ ' . - %- /

I F -

I screened endosure experiment (Table 8). includmg the absence of significant ooirelations /

-

-- a (Ta'bk Kt). As expected, percent __. - \

f&! a t and PI were substanrially b e r for Cdntrol A than Conml B, but fruit weight and -

number of viab1e"seeds per fruit were substantially higher, though not sigdifimtly different -- . .r

from Control B. In ' additim the mean number of seedless carpels per fruit was substantially

lower, though not s@iiEcantly different from Control B. Hawever, this may have- been due f

to the extremely small samp1e size in Control A; only seven percent of the blossoms used -

- - -- - - -- --- - - --- - - - - - - - -- - - - - --

for Control A set h i t , producing oaly four apples. Fruit set in the Conucd A treamem in

screened enclosures and outdoon was unexpected since these blossoms remained bagged for .

the duration of the experiment Blossoms which were still 'viable when the bags were - - --- -- -- - - - - - - - - - --

- - - - - - - -- - - --- - - -

- -

- - - - --

.removed at the termination of the experiment could have been pollinated at that time.

7

t

A comparison of treatments and controls in the screened enclosure experiment indicates *

that the overall pollination efficacy of Cantro1 B was higher thap all other treatments (Table i

8). Because Control B received unlimited pqlhation, multiple visitations could have consisted

of sequences of honey lxe and/or 0. lignario visits to receptive blossoms in the screened -

--

endos&er If SG, a pollination management system which combines honey bee ail 0. l i g w i a

visits mag result in improved7 p o h t i o n eflicacy on Red Delicous appks. Kuhn and Ambrose

(1984) @termined that pollination'6f Red Delicious apple blossoms by male and female 0.

&aria resulted in significantly higher fruit set than sideworking honey bees Since - sideworking by honey in Red Deiidous apples has been observed at levels as high as 6

4

86% of all f~ragen colleaing n m (Mayer 1983), the use of 0. l igm-a in a combined >

_ , - - , program a d mmpemate' for the potential re&aion in fruit set k o sideworkq has ever

- - ' , I

betlr &save& for either male or f&e L).-iTjgaia -(Table 7, -Tie 1986TTG h a y bge

-- - maq' EO WL syfferra byf vebfasymmetrical fruit D

Mean fruit weight fo\~ontrd B in vnened encl&urs was significantly heavier than

\

B - -- -

figrurnn " - one honey bee vlslt lhis result was likely due m the -- v \ -3 k - small sample size (n = f 3) and iarge c ce (SE & 16.9) of the 0, l igmiu- uearment - - -

+ 4

t k p a i 2 c i to me honey bee visit (n= 62;\ S E f 6.0).

of treatments and experiment indicates that the

Q of Control or two honey bec visit

. discussed preniobslly. mew results may have been flue m b r e r vigor (i.e. nee nuaition. soil

firti.li& and moisture content) of the orchard &ere the outdbq experiment took place' \

absence of significant correlations behveen, fruit weigh! and viable n m b r of seeds per fruit I '\

for all outdoor treatments (Figz 5). , . '1 '\\

\ . \ ,

d \\ -

\ The results of this stiidy s u ~ e s t that the pollination efficacy of one 0. lignaria visQ

\ . - on Red Delicious apples was equal to mat of honey bees in most respects, but resulted in '"i -

3 - \

a higher prqmrcion of aa)mrnem@ fruit The gllinator efficacies of one. to three honey \

I -

bee visits were similar. &use of me symmetry factor. I would not. recommend the use of

- - . ' O: ligmria & t& sole po3haor of Red DeBcious apples in h e 0kanaga.n Valley. However. d

a multi-species palligation system for Red Deliaous apples, ~ombining the foraging abilities of -u

honq bees and 0. Iignariq could potential& improve fruit set and quality. I . ,

- -

?ey -- a tn an orchard situation (raker and Torchio 1980); &d, 3) management system -

which Ge much less fabor intensive and expensive than honey bee &; beek TS 3? presently involved in honey bee pdharion could easily adapt their operatiom to accomodate

> -

management of an additional spsies for 'ooncpmitant use. +

, The superiority of, wveral?other 0. fignuria characteristics has been presented by other /

- - _ - - - -

- - authors (Table 7) but m y of them ZVe-not-be*-fully demons ted. For example, the P - - -

foraging behavior of 0. fignruia in orchards involves visits to few blossoms per tree but

many' trees per foraging trip (P. F. ~ o r c h i o . & , . ~ communication): This behavior pattetm -*..,. _

interplanted throughout the orchard Honey, bee foraging behavior in orchards is more limited a*

in between-tree visits than that of 0. iignaria (Horticul~~~$ Education Association 1967). but

- recent studies by DeGrandi-Hoffm et al. (1984) suggest that ' bee-to-bee transfer of, pollen

- within hives contributes greatly to the cross-pollination of apple orchards. Futthemore, b h i v e L r

polIen transfer would make the soda) behavior of honey' bees much more effective.' for ' -

/

* *

I pollination of self-incompatible Ipdes such as apples than would the' vhavior of. solitary

Nest initiation by 0. (ignaria was neghgible duriag the experiment in Summerkd - . .

Other resgrchm (5 F. May+: prsonat L u n i c a t i h ; kqhn and knbrose 1984) have also

encowrered problems in enmur&q nest initiation 8 0. lignari~ ushg the management . methuds developed by Torchio (1176. 1982b. 1984.' 1986). Since hated 0. fignuria females that'

. have iniriared nests and are provisioniq -re vigorous pollen c o l i e m p t h a n -

- - --- - - - -

nix having initiated nests (I? F. Torrhio. personal .ammlini&tion), o v d amrrilj,utim to 0

, * I

'\ 'U. S. D. A, Bee Bidcgy and Systematics Lahtwy, SEA, Utah ,84322

11 0 -

P

cross-pollination may be rkuced. A %eater impact on fruit set and quality might be

- L . at or above 15.7OC =ere r d i d e d for only 2.5 hr during the screened enclosu6 experiment

\

(8 to 15 May) which may have affected mating and nest initiation of 0. lig-a . Management systems for 0. fignark may have to be adapted to the biotic and abiotic

conditions indigenous ro various fruit growing areas. The use of. locally trapped] 0. lig-a in ,.

Torchio (1986) suggests that as few-as 618 nesting females 'could adequately pollinate'

one heqare- of apples' (247 treshectare). In his study, 0. lig&a were Weased- in an

. mention o f the abundance of wild &tors in the- orchard and their potential contribution . - f . to ,pollination in the apple orchard. Therefore, Torchio's estimate of- 618 females per hectare

mqy be an .underestimai& of tht actual number of 0. ligruuia needed to -provide adequate . '

pollination of apples. - t

D . . . - ,

E- the gsbawm e l f f , ~ d . f m - ef 0; 1-h- 4

' I '

1 w needed Red Delicious apples might be. done in an isolated orchard, similar to the

-

P -

situation Torchio (19f6) ukized but the abundance, diversity and m i b u t i o n to pollinption . - $

I by yiId polhpators should be monilljred- prior m the release of.-0. lignbia A ma% release

'

, 1 .

of 0. lignaria .would be nemsary',&ensure - that enough ipdifiduals would be present in the ' - . c

6 ' I

' orchard so that muhiple visits to receptive bbssoms could W y he achieved. Future studies I

i -

'should also investigate the e f f w of more than three bee visits on fruit quaiip. to allow a ,

4 *

more comprehensive mmparison of pHination eficades of both honey b;n and 0. ligwia .

f 7 - - - -- - -- - --

I *

CHERR'Y. PEAR, AND AF'PLE ORCHARDS

dg not have equal polhation efficacy. Therefore, apiarists in&t colonies to defermine their -

strength and potential value as pollina*tors of tree fnlit crops prior lo the commencement of - 4-

3

tree fruit pollination. Ely utilizing a basic pollination manageknt system for honey beks

which involves' the manipzllation of colony demographics. and population sir;, colony strength

can be adjusted to ensure that orchards receive adequate cross-pollination and ai~ -- - - - - - - - - - - - - - -- - - -- - - -- - - - -- -- ---

3

economic yieId of fruit. - .<-A- - . - 4

There are three essential features of a pollination colony which apiarists can manipulate i

-p-L-- - - -toemuige-xquziIiZation-of & -----;ZC--~

characteristics (i.e. waled and upsealed work; brood and adult population) most responsible , - for the foraging field force during polhation. They are the: 1) queen; 2) adult population; ,

!

and 3) brood. An inspection of the brood pattern determines whether the queen is a suitable

niatriarch for the rolony. If the brood pa&m on each frame' in the brood chamber is even

and stages of hood developrhmt (eggs to pupae) are represented, the queen is described

as& g o d egg-layer and does not have to be replaced The adult population provides the.

effective field force ultimately responsible for the transfer of pollen and: consequently L

fe&tion and fruit ser Most apiarists measure the -adult population of a colony by counting C

1

the number of frames in a super (hive M y ) covered by adults. If the colony is too strong. / /

adults may be remwed and mn' be added to weaker colohies to adjust their strength. The a * -

presence of brood, specifically unsealed brood (k& an+ larvae), stimulates foraging in general - . .

may be the sqme pheromone %at inhibits ovary development of worker honey bees (Jay - - - - - - - - - -- - - - - - - -

- s%&,,-B~ + r ~ d i n g - & ~ ; O l a n i - m o f w o * - ~ e r ~ ~ x ( ~ ~ ~ d ~ ~ ~ ~ ~ :- a,

+

apples. honey bees collecting p& are more effident pollinators than those coflecting nectar - - only, because they cany more poien on their bodies and are more likely to, transfer pollen - u, sti- (Free 1970x It i%-'therefore, important that apiarists involved in pollination services -

brood are present to. mahtain manipulate colmies so Qat adequate amounts of all stages of'

individual colany pollination efficacy. - - - - - -- - - - -- - - - - - -

i

Provincial and state hgencies, as well as local pollination - 1

--.- i

associations, have established , .

guidelines which define ,the minimum strength requirements for colonies-to be used for tree

that pollination rhe B. C. Ministry of Agriculture and Food - Apiary Program stipulate

of eight to 10 combs of adult bees with five to colonies shouId consist six combs of brood

supers are generally all stages of developdent A second super (both *

Langstmth deep supers) is required for extra mom while colonies are in the orchard. in ,. . * . I

nee fruit pollination is six frames fully covered by add5s k d at least 11 fr es of annb 4 ~.~ an1) in Wo supers (Burgen et $. 1984). In Oregon, two gad= of.

-

A and B) used for me fruit polhation are defined. These grades recognize

growth of a honey bee colony throughout the pollination

+colony should consist of two 2 m p s con- 11 frames - frames (4,000 'cml) should be occupied by live brood In addition, adult shdpld fully

cover six frames, An Oregon m e B orchard colony is one which fails to meet w, ~~~ ler &--&-A- mehard-&my- by not more- than 2 5 pe~cent-on~ W XiTKoUnt Xf L&

Washqton's and Oregods c d q smngth regulations stipulate the amount of comb

*

. The number of polliuation colonies required for adequate cross-pollination to ta& place* . 4

- a

is recommended on a "per he-ctaren basis depending on the tree fruit crop to be pollinated - -

(McGregor 196). For example, in standard apple planfin@ (250 to 400 trees per hmare),f . - 9 8 -

colonies per hectare are remmmendedtAve colonies per hectare are +mmmmded on high .*

- , - - aendq -a~k41antingS-of-semi-&d r ~ f 4 0 0 - ~ leOO E & - ~ - -6f- ,B :. I . I - - 3 -

a ' I \ ( . - ..*

.. Yj ae greater n&bjrh bf blosfms per hectare and the increased' tendency 0P hone; dees m -

< *

fo&e within a row @. C Miyeyer'; @erSOnaf mmmunicatfon). .For sweet g e .pollimtion, 3 . - * a ,

I ' unamactiv&ms of' pear blossoms to 'hone;* bees. (v-11 $42: Stephen 1958; Tufts and Philp

* 3 - - .@' L

1923). ' . - ., * - 88 0 I

\ * - . - P '

C 4 . . a + %, . ' . ' _

t

% -.. I b

_ d . i m m t l e ~ t h e ~ y ~ a n h ~ - ~ ~ . 6 *

/ manipubti'ms disrupt normal @my i@vity for. several days d;in'e prime. pollination periods. 1 A 0

- -

severely reducing the efcigacy of poUina& -x#ts. 4 method of utilkinuing fbrager entrance ." *

counts. as an indirect measure of pollinatioh ynit men& wo$d eliminak'the need co .,

0 * 0

d L

dismantle mlonies .3i M a & . during iospstion and wmld provide a m o y passive but *

-

accurate measure of colony potential:. for orchar&, apiarists and 'apiary in&emrs. . - - 1

. ,

9 $ @ t 'b

9 - Since previous studies' have indicated that tde propix& d yiriien-@ering fdragck , P - T

, ' the dmy characfcristics (ImpeaLd and .sealed bfdod and adult populatiod) of pollination

d %

. uni& &ed for 'cheny, pear ant a d e p@hation in the 0- hly; and b P b -

2. . evaluate+ the effects of tenqpturt global solax bdiatik (G. S. R) and relatiye '

. _ . . - - ,,rs . . . *; I

. - ~- - - ~

. - - -

./ < C - ~ -- -- ----L,----- A . . . . - .

' ~ -

I --

~ ~ -- --L. -- - -- - ~ - . - ~ - ~ - - - -

- 4- 8

. . MATERIAIS AND METHODS

" 16 May 1984 and 21 Ap$l to 19 May 1985. Forager entrance c q w were rrcorded for -- -

t '

honey bee' colonies employed as pollination units for three tree fruit &ops: &ma (Runro f /-- -

1 uvkun), pears (Pyrus conymmk) and apples ( M h s#re~ris). The same orchard sites were

used for cherries, (V. Jmes Orchard, Old Vernon Road, Rutland) .and pears (W. Cameron

- -4 Byms Rasd, Kdowna) in 1984 and 1985, while in 1984 forager &tran& counts in

* ' 1

pollination as set down by the Okanagan Pallination Association (O.V.P.A.) and the B. C. M.

. * ' v .A F.#-. Apiary Program. Following placement in each of the fruit crops, the pollination units

. . n. a r

= . were &wed a 24 hr reorientation p e e d before forager entrance counts commenced. I

,- \ -

. 1984, forager mtrance a m t s were determined for nine colonies in chemes. 17 in - ' .

=and_foy~- The number nf m a t e r i n g t k - - U W -

was recorded, with three consecutive one minute measurements taken Four repetitions were , i

completed over two days. An egxmce counts were recorded between 1OOO and 1406 hr. A

themo-hygmpphwas placed near the pollbtion units in each fruit crop to recbrd anibient

-temperature (OC) and relative humidity (%) for the duration of the study. !a addition to

ambient temperature and relative humidity, global solar radiation (Langkys/hr) data for. the

area were obtained @om the Atmospheric Environment Service6. Colony draracteri&a . % 9

measured wexe sealed worker brood (cm2) and area covered on hami by adults (cmz). - - - - - - - - - - - - -- - - - - - - - --- -

- -

immedizrelp fdlowing the Iast entrance counts and .prior to rething the pdlination units to - -

the apiary fdlowing pdl?nation, the total area of sealed worker hood and add; per

~ a & on the 1984 resul6 changes in the e&ental he&@ were made in 1985 to'

imlude the measuemmt of @th foragers exiting and poll&karihg fwe i s enmini per - I- - - -

wlmy instead of foragers enter@ per. admy. She +use of poll*&bearing f q e r s as an . .

- index of pollination unif strength is supported by a number af stu@es. which determined-that -'-. - * L

the of uosealed .bro&l (eggs and larvae) ~ ~ u l a t e c l ' f o ~ ~ in gene* and a I i

: +-. pollen-gathering in particular (Al-TMty et ai; 1972; .fie 1967; -1979; Fiee and Williams

'I recorded to determine ovnall adult populatioa ~ h 6 &mbei of foragers 'exiting y d the -

. ,

. ' / nm_QZ poll-- -- --

- - -- - - = t e r i n g ~ e r - & ~ ~ L k w & u e ~ ~ ~ ~ = ~ \

, - . '

alterrhtely- for thr& amsecutivc one minute psriads each. Six repetitions were,&e7or 1

colonia izsed for cherry, pear and apple 'pollination; 10 -colon& were used for' each cmp. - .

1- . b

P

*The daily remrding period and the three ~ e & a bnditions 'recorded were the 6 as in . ,

c

1984. Characteristics measured included bsealkd. anddimled worker brood and ana of adults '

-, -P

wing the 1984 method Colony characrexistfa ;ere meslsured ~ e d i a t e l g * following - l e last r f '

iurager entrance counts and prior to returning the colonies to the 'apiary. ' - , -

, * Y =. . . . Linear. regr&m analysis was &d to detc-e the relationshipD between forager"

r.

entrance a m t s and d o n y chmkmMcs. CorrWtion analy& was used m deteimine - I . ? 7 , - . - . .

relationships between f q e r entranck counts an'd --____- weather data - . * , b

: adult area v d r y ;foragers entering colonies in chem&'and pears .(Fig. 6). -b appla. --' '- - - r= >

I .

.* I, . + ' --- , /

entering per colony were w&ed w i t b temperame (r=-SU)in chema (Fig 7). In &; 1 2. .+

3.' i ..

1985, significant gegrdons @ < O.fl5) were found for d e d - -(tl= ,.546) q d

4 d - *

sealed wprLm brood (f = .695) versus p6Jlen-bearing foragers entering per cblonj. aod adults e . -----

H . (r2= 560) vex& -foragers . exiting p r @ l k y in Le coloqies used fix apple pollinatiq (Fig:

7 A

+ b ' B A

8). Signifmt ?&essiom between Iorager m 8 . and colony chara&risti& were not found for -

9 . * f \ 8

% . cherries or pars '@ > 0.05) $Fig 9 and -10). , , - c - . I

4 . * ' #

--

- * & e m ~ . significant mnelatims . (p. <- Q.05) weri . found when foragers exi& .per 0 . I - I . 1

k m y ' were amelated with R+.- (r= -287) and temperat& (r=. SW). a n 5 when ' a s . .

pollnrbearing foragers en& p(7 q l w y wefe* apelatedo with temperanire (r= -291): C

/- . . . .

G-SX (r= -368) and' RH. (r= 364) (Fig 19.' For pears, significant positive' carre~kms ips t . ' < 0.05) were. determined ivhy foragers exiting per colony ' -rre&xJ 'with G.S.R i r= ._

t- > 0.05). between %a & and the thne weather amditiohs were found ,in qplpples for C

I -

Fig. 8. R~~ analyses

APPLES

r2 = ,144 N.S.

r2 = ,317 N.S.

FORAGERS EXITING COLONY ( i COUNT MIN-1)

POLLEN BEARERS ENTERING COLONY

( 2 COUNT MIN-1)

F .q * analyses of f a a enring cdmy and pllp tearen muring pci+dony

Y characttristik, mas of adults, seakd brrood q d unsealed &ra , . 1983 while dmts were lacatcd in cherry orchard Mean. mcmt per-

- . was deririyed fTom total cwnts fox 18 one minutr inttrvair .. - i' .,

-- - - --- - t,!@-- . - i - - 1

-- -.

i h .-_

CHERRIES

. 2 = . o N.S.

r2 = .O N.S. . . . . ..

r2 = .o N.S. . . .

0 . . . . . 0 . . r2 = .o N.S.

r 2 = . 0 N.S.

r 2 = . 0 N.S.

POLLEN - BEARERS ENTERING COLONY

( 2 COUNT MIN-')

FORAGERS EXITING COLONY

( i COUNT M I N - ~ )

PEARS

.

. r 2 = . 0 N.S.

N.S.

r 2 = . 0

6 N.S.

I 1 I I I I 80 90 100 110 120 130

FORAGERS EXITING COLONY

( i COUNT MIN -1)

r 2 = .O .

N.S.

N.S.

I I 1 I I

r 2 = . 0 . N.S.

POLLEN - BEARERS ENTERING COLONY

(k COUNT MIN

- - & 5

i ,' /.---" 4%- * -2,

~i 11- ~ o n k dyrei-of fongcls a pa asmy. pailrn tprm enem a k m y md thrre wera'hn mndiliaa. b&bI rdu mliatioo Lmc hlrmidity'(RH.) e iwf lmFp2m. Data were ~CQIIW &ring tht F4Ymmjon 1985 while 10 *tdonies were l~l~itted in a cherry Xmhard. Mean cmno pr rahMc for six ~ p h j d m 4

per colony resulting in 60 data poias fur c a r e h i m 1 _ , t~ i

CHERRIES

r = ,139 I N.S. *. . 2 .

FORAGERS EXITING COLONY

( i COUNT MIN-I

POLLEN - BEARERS ENTERING COLONY

! - - ,"F 4

? I t'

$>

s:

> -- 4 l

.'a . . ' 4 +

- Fig 1L Conelation analyses of fixagexi exiting per colony. qolkm bearers entering per. coZony. and three weather, umditIons. sodar mdiation (G.S.R), relative humidity (R.H.) ;tnd remperantre. *Data wkre r e c ~ ~ G e d - dming the pdlination period in 1985 while 10 col@ies were l a t e d in pears. Mean counts pereminme were recurded f a six repetitions per colony

. resubg in 60 data @inis per,correlahicar 4 1

- - 4

9 , . , ' 's J

PEARS

8 033- 2.820

e 14.5 . 80 U. u 8 . r=-,107 I- NS. . . . 13.0 --I

I I I 1 I 1 25 60 95 130 165 200

FORAGERS EXITING COLONY

( i COUNT MIN -1)

. . . . . 8 U . ...... 02.0. . 3 ..2.2" 0 . 0 . . . .or4 r..

0 2 . . r = .329 s. P c 0.05

POLLEN - BEARERS ENTERING COLONY

(i COUNT MIN

APPLES

r = .I23 N.S. .

r = ,174 N.S.

0 .

22 022. 4230003 . . .

r = ,162 N.S.

FORAGERS EXITING COLONY

( X COUNT MIN- ' I

. r = .048 N.S.

20 I 1 I I 1

0 10 20 30 40 50

POLLEN - BEARERS ENTERING COLONY

( i COUNT M I N - ~ )

for apple

Ad. for

colonies king -used for polfination are f&Iy mong if there are more than 100 incwning

bees per minute at 18SCC and winds jess than 16 Lmh. The dififulty wilh using foragers -- - - - - - - - - - - - - --- -- - - -

arering mlolria is hi wfhaadulr am in colonies can &e prcchard, and therefore one of

the rxgijo~ components' of cniony sngtlgth. brwd area, is not rehted to tnuance m r s .

Fmagn mtrance counts which would alro pr&cr d e d and unsealed worker brmd area - - - -- - - - - - - - -

would- be more useful as a measm&~ of colony mength. -

i - . / In 1 k . significant rqmsiom for unsealed and sealed wcsrkcr brood (Fig. 8) v&ur

b

pollen-beaing fowers e n ~ x k i g per colony were found in the colonies used for apple

poUinatign. These rzets agree with a number of studies which determined that the amount .

1972; Free and Wil lhas 195; Todd and Reed 1970). -In addition. Free -f f 967) concluded -

than sealed brood (pupae). My results indicate that unsealed b r a d (TI= S46) was not as

effecdve at stimulating polen-gathe-ring as the sealed brood (?= .6%) (Fig. 8). These

My resuits aka indimre tbaf in donles used for apple pollination, foragers exiting per

*

#

' d d 'be effmvely waked by are individual dm).. If a colony. &d- not meet adult strength

- - - - - - - - - - - - -;r mpirm-gns. i t cwfd tlrw-k z3Jus-m- ac#lrdi@~:- --- ----

b I

pollen-bcarm entering per colmy in cherries and pears appear to respond to each differently.

In a mdy by &mill and Diee (19%f) it was determined that temperature and G.S.R .

honey bee fight d m In BmiB and Diee (1981) found that significant . J

i . * * * ~ - a & - + t a m + k d a . . *

-4

below 0.66 langleys, and negzhely amelated abo& this threshold In both years of my

k1985. the significant correla&ns found beween foragers exiting and the various

findmgs of BurilI and Ben (1931). A positive amelation between temperanrre* and 3 a

The s i a f i w t coneiaticms fotmd in &ems and pears ( F i e 11 and 12) when

-./' that in foraging area&& - m n t lo pears as a source ' h e n remains high. '& m n d explanation for hooey ke foragin%_Whavior jn Phemes and pears is related to the +

- - -- - - - - - - - - -- - - -- - -- - - - - - - - - -

positive conelations fohd k w m pollen-hmg foragers and R H in*'&ese wo nee fruit 4 1

Y

,/- crops (Rgr 11 and 12). Relative humidity and ,nectar secretion are positively correlated.' but i

-- - 3

su_m co~~cennation in the nectar is negatively mnelated with R.H. (reviewed by Crane 1974). '\ ,

- . Thh phenomerioq ,is due to rhe hypsapir : effect of the sugar contained in the nectar; ..

&&el 1952). Since it is the sugar ementiation in nectar which a m w honey beer to the'. '. .

bkxsoms, at a hi.@er RX bbxms woufd k m n e less amartive to fomgers. The results of. -'

t .

k *

my research suggest that, when these weather umditions occur during cheny and pear .A .- 9 -

~ c m A m e - i s a chift w - m W r n - are a ?

-. . I

considered to be as arrracrive to honey bees as other Fruit crops due to low concenuations .., ' *

of sugar in the nectar (Vaoseq ZW?; Stephen 1958; Tuf'?.s and-Philp 1923). &d on rhc t . . I

results of this study. weather condiubns during c h e q and pear -pollination which , ' 6

'i .. involve increased RH. m a y improve polli~tion efficacy of honey bees in p& by stirnularing . .

I *

-

pollen-gathering,. and therefore fadliming cross--tion.

From this study that predicting dmy pollination efficacy in -

-- - - - - - - - - - -- - L

/ uee. h i t aopr The foraging behavior of honey bees in tree fruit cmps such as cherries /

/ and pears, which Moom earlier if, the sprhg.than some apple varieties li.e. Red D&cii&)

/

in the Okaragm Valley, aze @erted more- by external weather conditions than inteTnal, colony .

-. : fiui\ @ w i n g args-*\ln the 4rrblooming apples, colony mcngtb is dependent on internal

4 ,

I b

emph&& in ~~edictive r y s $ m c ~ v e l o p d colony polltnation elficacy in this -, : 7 -

s + * . -$$ tree f i t crop;. 1 . - - - f

population of bees u> transfer the pollen while the flowers are receptive (McGregor W 6 ) .

Cmmt pdhation mzumgement imdves the use of overwintered honey k colonies whicb

are -moved back to apiaries for hoaey production

bee colonies (pllinatim units) are

a d h i . b f x k a p m thar their mamgem& syslems cwld be more productive, thus yielding

b a r income. They were particularly intexested in the following areas:

I. Honey Pr@uction - The movement of bee colonies to orchards seriously di&pts colony

growth during the critical spring population buildup ,period, since 2535% af the P I $ . .

foraging force mq- be lost due to disoientation within 7 days of a move (Nelson --- --

1985). The effect of this drmrption in colon). growth pa& is likely -to reduce the - 8

% I

oop kter in the -%but tbe extent of the 'loss of honey production i< not -, \

\ and hbney production under pdfination and MW-pollination management systems. , -\

\ / A ~

L Bte Produaion - Comiderable interest has recently developed in bee production in

candda kfudmg packages and (Wimton 1983. 1986). A packecof bees .

generally consists of 0.9 kg of wmkm @ a queen. while2 nudeus is a snall colony

the Y.S. with a %he 01- $10 million (Winston 1983). R m r l y , however. the

bees and two potenriallp serious mite pests of honey bees, Amrapis w d i (Rennie) and 4

VWTW jzcc6- (Owkmms), into the packageproducing areas of the U.S. (reviewed by

at. 1985; Winston iB6). It may atso be feasible for the Okanagan Valley b e e k e r s

to produce either packages or nudei as pan of their pollination management for an - - - -

- - - -

, additional source of k c m e . However, no data were available concerning tkie impact of

-'* such bee production on either polbation or honey production, nor had there been - r e s a x 3 involvhtg metho& of bee prodmion.

,

h e objective of this SRtdy aras to examine the ?oh and economics of 3 diff&e& t :.

bee management sysrems in tbe Okmqp Valley: 1) pollination (-en: system); 2) honey

production witbout pollination; and 3) package and nucleus prmiuction with and without

pollination. , .

2 apiary sites 6 km apan near Armstrong The apiary sites had similar honey yields in .

previotts yea^^ which were Gonsidered average for the region @=34.0 kg/colmy From 1974 tb I

1983. = 3 1 . 0 l&dmy for 1983, BC.MM. 1%). and had mmparable elevations @00-650

neatments were:

1- .

2.

3.

4.

5.-

6.

Each

No pebjge or nudeus removed but cdony moved for pollinationJ~trol - - Porntion);

Nucleus removed apd-colony not moved for pollination (Nuclens - Apiary); - Nudeus rebved and mlmy used for pollioation (Nucleus - Pollination):

Package removed and colon). moved for pdlination (Package - Pollination). - manipuIation is described below.

Controls: These donies had no packages or nuclei removed in the spring and were

managed ~ O T honey producrion done or for honey production and polkation.

/

* - Packages: Twenty of the cdorda had between 0.6 and 0.9 kg of 6 removed from

nmovcd MS that the pmtal dcmjes were left at pollination d t 'strength after dt.ak@. A -

standard polhatian unit was masidered t amain at least 8 -bs of bees- a& 5 4 combs

---

mfficiem- mength tq - +ve a M -0 -T kg cp&.TiZiw-(T I 6 - T n i i while - h - /

M Q.6 # W k g *-=-a as cht nuclei,

b - <,+:

Nuclei: Twenry colonies had 4 - h e nuclei removed from them on 18'Apxil 1983. - ,

Nuclei of 2 fmmes of brood 2 frZmes of honey an-d pollen. and enough workers

to cover all 4 frames. The parental colonies were maintained at pollination strength.

Pollination: Colonies destined for use in poIlimtion were &vdd on 24 April 1983 to a - block of Bartlett and Anjou pears (Benvoulin 0rcbard.s-Byms Road, Kelowna, RC, Owner: I

1.1

4 W. Cameron). On 2 M a y the polhation colonies were moved to a block of Red and .Y

- - - - -- - - - - - - - -- - - -

Golden Delicious. Spamns and Solmer Orchards Ltd-Pmley Road x C . .

Rose Road, East Kelowna, KC, On 11 May, all e

were moved back to the apiary horn w k h they prighated For the r 1

summer ci,lonies used for poliination were managed in the same manner those remaining PS in the apiaries. >

pp - -- - - - -- - -

A&: k colonies remained in the apiary for the entire summer, and were used

\ to produce honey alone, honey and a package, or honey and a nudeus. Management included

swarm mnW.adding extra supen whdecesrary, honey stoJe%E. regular queen checks. etc.

Colony characteristics measured were: 1) net colony w~~@L&To~$, colony w&hi - *% -.-

t -p --k.- 7, .%,"

weight of empty equipment); 2) s d e d worker brood area; pdA'3)' surplk jymq production.

,, Sealed brood area and Lnet colony weight were measurq every 21 to 25 days (approximatdy

the developmental period from egg to 'adult) from 3 March to 10 'August 1983 and prior to - -- - -- - -

major manipulations, indudhg fernoval of gadcages and nuclei, moves to and from pollination. - - - - - -- -- - -

Ad emaction of honey, Sealed &er bmod was measured by .p-kdng a dear plexiglass \ L

pried (13 ~ a y ) 1983). mid wavm (8 June 1983), and at the &d OF the season (113 August

,1983). Surplus h ~ e y prodrrtion was analfred udng&ta mil& fhm the single egtraction

' colony weights and sealed. brwd were wparated using* Studem Newman-Kuels T& * . . I

---. / ranired udng August data lo. de&* which system was the most productive. Gross -

of package or nucl'ei, rale of, honey. pollhation c6nm&) minus expmvs for each p e n t

as indicated in the Producers' Consems Report (B.C.M.AF. ,1984],~ . A a

%U

Cost breakdowns used in detemhhg profit/oolony were as follows: /-

- 1. Honey profit:

4ka+lme- x pFKe&#-ilmq+pm - frt---- --

In the Okanagan Valley in 1983, 60% of .the honey was sold k -an average price of

Sb53 kg (61.15Ab) to customers who have their own mtainers The aher 4% was

sold in conkinen at $2.75& (S125Ab) to refail outlets. Honey profit for each,

2 ,Po+ticm fee: ,

~36.00lpldliaation colonyfset x 2 sets = $7200.

the typical way of selling- nuclei in BC. - - P

-

i C o h y characteristics and profit per colony were all weighted

4 '

equally in determining ;B - the fd ranking. Colony characteristics for each managemgnt system

8

in August were ranked

on a scale of 1 b 4 (1 indicating a first place ranking). Statistically si& characferistics r - -

were giv equal rankings The gross pr fit. per colony was also ranked for each managemenr - ! g - - - - - - - - - - - - - - - - -- - -- -

for each colony characteristic and gross profit/colony were summe9. tp give a final cJJ

* than the 'Nucleus - Apiary' trekment (Table If). In August there were no significant

, A --

differences in the mean weights of any of the 6 trements, although the 3 heaviest * P 6

ueamenrs were the "Apiar)-' cdonies (Table 11).

,I+. \

In May, there were ne;i&ificant differences in the mean amount of +ed w e e r - - - - - -- - - - - - - - - - - -- -- -

brood i n any of th&&&ents except that &e ""Nucleus - Apiary' treatment mntainecf 4

significantly less sealed brmd than the "~ontroi - Pollination' rreatment (Table '11). There *

were no significant differences in the &owit of sealed brood found any of the 6 - - - - - -- - f - - -

- L-- L

management systems in Jtme (Table 11). In A u p s ~ a . mean area of sealed worker brood . -

in the "Nucleus - Pollination" and the nConml - Polhation" treatments were significantly i

greater tt&-tke "Nucleus ApiaryQ and "Cun~ol - Apiary" treatments. -' - '- - .

C- ,, 1 f i

t

i --. Ahalysis of hdde). yield indicated that the 'Conuol - PoWtidbn" management systaf~,

produced I'@ficantly less honey than the other five management systems (Tplble 12). I

\

\ ', .\

T ~ A systems involving pollination kere more 'profitable than those remaining in thq '' , \ ! tu-

apiaq (Table 12). Also, the m o sys& yielding the highest pm profit had a package or -

nucleus removed. . C

- '-L -3

Eked on bid@$ and e a m m i c Baa the finaf rankings f o ~ the six- management -1

- systems resulted in the -kents involving pollination being the most .prodwive (Table

13). The two tresr uatrnwts inducted remod of a package or nucleus from edonies in

--J Tahle 12, Econclrdc &€a for 6 honey bee mnagement- treatmerrts tes ted-&-- -L-----

the Okanagan Y&fley, B r i ~ i s h C c r l d i a i n 1983."

FIanagernen t Eonel3 Y i e . 1 d Gross T r e a t r e n t (kg) Prof i t JColony I-_- L- L- --

Cant rok-Apiary C o n t r o l - P o l l i n a t i o n

Nucleiis-Apiary 25.1 r 3.3 a 537.01 Nucleus-Pol l ina t ion 25.5 I 5 . 7 a $109.28 .

*mans were conpar-d by Dmcan's m d t i p l e range tes t (195 1') ; means w i t h i n a column f o l l o w e d by t h e sane le t ter are nor significantly d i f f k r e n t f P s 9 . 0 5 ) .

Table 13. Overa l l p r o d u c t i v i t y ranking of 6 honey bee management t r e a t m n t s t e s t e d i n €he Wcanagas Valley , B r i t i s h Cofurrtria f n H83.*-- -

biological Ecoqomic Score Score Overall

Score . , X ~ m a g e ~ w ~ E weq; Sea2 ed ;Honey erits s f B ~ ~ - T T @ c%? - - ---

Treatment , w t Brood a P r o f i t economic) A /

Xucleus-Pollina t i o n I I / 1 - 1 4 Package-Pollina t i o n 1 2 i 1 1 5 €on t r o l - ~ o l l i n a t i o n t 1 9 i 2 2 6

Package-Apiary I 3 . , 1 3 - 8 Nucleus-Apiary f 4 $ I 3 9 Control-Apiary f 3 1 4 10 % /"-

, ----A - - - ---- M p '

* Low sco re s i n d i c a t e the mre product ive t r ea tmen t s .

' colony management Tor pollination, honey producricn and bee production do not detract froin

overali colony vigor and yield the best incane. ~ u t & m o r e . the mon productive management

sysrems (Table 13) can provide a new -source of income through the sale of packages and/or - t

nuclei. C

I

Colonies involved in polhation treatments had lower values--& apiary treatments-in --- -- -- -

only one biological characterimic. colony weight in May and June; by August there were no " -

si@ficat differences in weight between pdhatim and apiary treatments In May and June. *

the educed weight of colonies used for polhati611 m a y have been due 6 worker - - -

-- - - --

disorientation and drifting (Free 1958; Jay 1x9; Nelson 1985) in the orchards resulting in

less nenar mUection during pollination compared with apiary treatment colonies. It i; also

possible that the honeyflow was more intense at the apiary sites than in the orchards, since

a p i q treatment colonies gained an average of 15.0 kg during the 3 week pollination period

compared to an average gain of 8.0 kg for the pollination ueatment colonic&. -i

- - -

'However. moving colonies -for pubmion did not diminish brwd rearing relative to *'

aplar) mlonia. possibly baause the &mxnhred pollen source avaikbfe in orcfiards during

the pollination period offset brood loss by 1) maintaini~g brwd rearing at a level comparable

to ap- colonies andlor 2) allowing for a rapid recovexy in b r d rar ing In fact, moving

'\ mionia for pollination m a y have acrualiy-nimulated brood rearing sin9 by August pollinating

- - --

p r o d ~ as ZIP& or more honk and ad braxj as the apian. amml mlogia ?he

Apd might be expecred to dhninkh subsequent brood areas and honey production. Rghtively

low lev& of stress on cohiw from IPee pr&uction, and pesticides may actually stimulate

brood rearing and. honey productiw but thi~ phenomenon has not bem investigated. A study -

showed Iowa Iongevities and ages when f o n g i q commenced in colonies from which a I

majority of the w&ke~s had tieen removed This simulated lo& in worker populations is

naturally occuring c d ~ events such as predation, disease* nest damage, S ~ ~ ~ X I I - - pp --

f and/or W g a a t =- manipulations involving padrdge and nucleus production or colony division.

Y +

i* - Winston and Feqmson (1985) suggest that shifts in lifespan and foraging age indicate that

workers may adjust their zemp0ra.I caste schedule to respond to rapid changes in colony "

mnditioor Other studies support &ae fmdings (Winston and &en 1982: ~ol rnes 1985:

These resulk have - two importanr implications for pollination management First they

indicate h a t although moving bees for po~inadm can reduce honey production. the increased

income from poilinauon offsets any 1m of honey. Further, the only significant reducuon in

honey proc$uction .was found in colonies which were not used for bee produrtion (Control - Pollination). Using pollination tmizs for bee production as weU as pollination resulted in

Secondly, the results of our research indicate that Okanagan Valley beekeepers could ,

- - - - - -- - - - - - - -

p r a d u packam and mdei as pan of their *tioa Gg-t-- ~6 *-

in reducing Canadian dependence on imponed bees An i n d m of this type w d d not only -

be advantageous to B.C., bur ma)- also be an imponant factor in the fllrvival of the \ G

Canadian beekeeping industry. In addition to rising import mns the. beekeeping industry faces

three problcms which are' moving nonhward from Latin America any of which muld result

in the limitation or cessation of bee imponations from infested areas of. the US. These

problems are the mites A C ~ Z P L ~ w a d i and Y m m- ( M m g et al. 1982) and the

highly aggressive Africanized honey ks (laylor and Spivak 1984; Winston 1983). The

economic porential for B.C-produd bees is promising, even without any resuiction of -- -- - -

impmations However, the -bIe r e d e o n or ounight ban-on importations of bees from ,

the U.S. in the future muld result in B.C. supplying &'on of Canada's ks The results of

our research indicate that beekeepers in the Okanagm Valley wuld play an imponant role in

CONCLUSIONS

d -'

1. Wild bees. al- abundant in namd habitats, are na ~chabte or predictable

pollinators for orchard crops in the O k m q m Valley; w

--- - bi- nearcticus, display a mwg tendency toward foraging a orchard uups and arc

tendency toward fm on apples and is indigenous w the region. Also, management - -- -

systems have already been developed;

3. A mulfispecies pdlination system for Red Delicious apples, combining h e foraging - abititia of boo& bees aod 0. ligmria could porntially improve fruit sa and quality;

-

marcticus and 0. iignarin for trap-neshg .and utilization in tree fruit pollination.

6. The planting of understory cmps i r t d s which would complemeni the flowering

cycles of .various fruit cmps could resulr in reraining populations of borh managed wild 0

bees and- hohey bees in and near orchards during ?he bloom period;

7 . The pollination :&aq O! 0. Irg- on Red Delicious apples was equal to honey

bees in fndr qualip charixzeristiis m d ~ d m g ' ~ 1 i i ~ t i o n i n d i m and fruit weight and set

but was lower than one, NO or riuee nmey t ee visits in fruit s)mmetxy;

. , /--

9. The foraging behavior of honey bees cfn cherries and pears was affected m e bv

1 . Beekeeping management systems which involve the most in- management for

goUination, honey produEtian, and bee p~sdlrction do mx derzact from o v d colony

vigor and yield the best h m e . F ~ ~ e , the most produaive management systems

Honey bees will continue to remain rhe primary insect polIinatq of nee fruit acps

r d t in increased efficiency and a h@a ecrrnomic re- are impfaas to bo& hefkcpe~s '' =

and orchardists.

- From rhe perspective of a beeketiper h v d v e d in the rental of honey bee colonies for

potttmlhn increased inoom:: r s u h g fnzm the sde- of packages MOT n d e i and potentially

from the rental of wild bee pdhauns milked in a multi-es sysrm would be - >-

appreciated The inc~usion of bee pmduaim in polhum . . -- man;iogmpnt- In the

0kanaga-u Valley has been well- received by beekeepen and has proven to be su(x:essful.

for wild bee pollinators beam: more zccessible to beekepen The management of 1 -

rrcidcnrofif md B. b i f i u i ncz&- &odd k initiated in order lo refme multkpecies - -- ---

From the orchardim penpcrive. @hation systems which are directed at improving

---. ~ o h f i o a deny strength devrminarion using forager entram munu is one am of

ofihardins and inspenors with a simple method of determining whether beekeegcrr have

beeketpm and orchardims is ssential for continued improvmenl in pollination management

Collection sites and t h e i r h a b i t a t designations f o r 1984 and 1985.

Defini t ions of h a b i t a t designation: A

1) Orchards: Far from Hatural = s i t e ' is located i n an orch'ard wj~ich is surfounded on all s i d e s by o the r orchards ,

2) Orchar&: Near Sa tura l = site is located i n an orchard which '

is bomd an one o r two sides by na tu ra l uncult ivated land . - - - - -- - -

- - - - -- -- - -

3) Uncultivated: %ar Orchards = s i t e is na tu ra l uncult ivated land bomd on one o r two s ides by orchards

4 ) Uncultivated: Far from Orchards = s i t e is na tu ra l uncult ivated , land completely surrounded by na tu ra l h a b i t a t and is

t - - -- - - -

>0.5 h h zke mrtzsr orchard -

- - 1

Ccllect ion S i t e Habitat Designation

I) G.O. Robertson-Mathewes Rd., &t LJneultivated: Near Orchards Kelowna, B.C. (elevation: 450 m) . .

2) W. Camron-2050,Byms Rd., Kelowna, Orchards: Far from Natural B.C. , CROP=Pears 1

+ ''a

3) Dilvorth H t . Estates-Dilvorth Mtn. Uncultivated: Fa r from Orchards '

Drive, Kelowna, B . C . (e levat ion: 600 m)

4 ) C. Day-Day Rd., ~ u t l a n d , B.C. CROP=Apples (elevation: 570 m)

t c- Orchards : -Sear Natural

5 ) Sutherland H i l l s Park-Ball Rd., East Uncultivated: Near Orchards Kelmma, B,C. (e levat ion 360 m)-

6 ) D . Claridge-Todd Rd., Gyana, B.C. € R O P = # ~ rries

Orchards: Sear %turd

(elevation: 500 m) - - -

Colle t im S i t e P - -

d - - - - -- -- -

H**=fk~tmECMdn~

7) Agriculture Research Stat ion, m*d - - - , . - 8

- 3

(eleviition: 454 m)

a ) Orchard Sites-apples , pears , che r r i e s

b) Budwood Orchard (bound on 2 s ides by uncul t ivated land) and Arboretum (bound on 2 s i d e s by -

uncul t ivated land and receives no . chemical sprays)

> - *

- 8) V. James-Old Sernon R&.-, Rut~ancf, -

B.C. CROPSAppfes, €berries, Pears -

(elevation: 420 m)

9 ) Knox Mtn. Park-Knox Mtn. Drive, Kelowna. B.C.

Orchards: Far from ~ a t u r a l

Orchards: Far from Natural

Orclia kds : F a r f rom-~a tGa1-

Uncultivated: Far from Orchards

10) Agriculture Canada Research Substation, Orchards: Far from Natural Hart Rd., East K e l m a , B.C. CROP=Appl es (elevat ion: 450 m) . i

1 abandoned Apple Orchard-Haynton C r t . Uncultivated: Far from Orchards a t westside Rd., Vestbank, B.C. f f

(e levat ion: 330 m) .' i t . --- -

- 2 - ekarragarreefi-tre ~d . , W i n f i e l d , K C . UG'ultivated: Far from Orchards 1, (e levat ion: 500 m) a . -

&.I

13) Oyama Lake Resort Road, *ma, B.C. fFncultivaeed: #ear Orchards * .' - ' (elevation: 750 m)

- = ,t

I

. . . * I

r a @ - -

? . . i

, 0 - . . - a

. II - < .. - - --

Y d . .".

APPENDIX I1

The h a b i t a t s and f l o r a l v i s i t a t i o n p a t t e r n s of wi ld bee s p e c i e s co l l ec t ed i n t h e Okanagan Val ley i n 1984 and 1985.

I

Habi ta t d e s c r i p t i o n :

Habi ta t 1 = Orchards: Far from Natura l

Habi ta t 2 = Orchards: Near Natural

Habi ta t 3 = Uncult ivated: Near Orchards

Hab i t a t 4 = Far from Orchards: Fa r from Orchards

Bee Species Year P l an t ( s ) V i s i t e d Number of Bees Col lec ted 1984 1985 (1984 & 1985 combined)

Hab. Hab. Hab. Hab. 1 2 3 4

AP IDAE

Babus terricola x x Taramcum of f ic inale occidentalis Prunus mim

(sweet cherry) Fragaria virginiana BaZsamorhiza sagittata Matus s y Zvestris

(apple) Malus sp. 1

(crabapple) Rosa nutkana Berberis aquifoZium

B . fervidus x x T. off ic inate B. sagittata Heuchera c y Zindrica PotentiZZa recta AstraZagus miser

r r A - 7 %?<

&

/ -- -- -* - - P-~-- -

- J' -'-

=& -+ * * - - - -. --- ' 2

.. ' . . 4

-- APPENDIX I1 cont.

- - - * * ,/ - --- --- - -- - L - - -------A- ----- 7 - :'** ,*' -

@ $ : - , % -F; '4- * Bee Species '

*

" * Y e ~ P l a n t ( ~ 1 Vf sf mi= . - .

1984 1985 . . I

r >

~ a b . - Hab. Hab.. Hab. -

1 2 3 h t t +3L + Y

* . I

- - 1

-B. central is x x 4. of f i c imZe 1 . * 1 3 ' f - M. s y h p s t r i s - - u

6 , . l . 2 - MaZus s p :l " 3 0 1

\ C

B. sagittata 1 4 B. cyE.indrica 2 3 -%. --

- - Sym&tm-of_ficina Ze - - -- -- - 4." - & . -( rL -

Dodecatheon pu Zche Z Zum l l . - - --- P. avim * 1

I c

Astra~ogus neglecta ' I 1 1

I mZaspi amtense ' 1 Eigerorr phi lade Zphicus -- 1

- - -- - - - - - - - G t b p m e x d e ---- - -- - -- - - - -

PotentiZZa arguta T

\

B. bi farius x T T . officinaze -: 1 G 7 nearcticus t P. a v i m

,a6 2L M . sylttestris , 5 1 4 - 36 '

- € , . I

&Zus s p . 1 6 . 1 B. sagZttata , 3 6

. D, puBhel2wn 2 "t

1

hnelanchier a ln i fo i i a 2 .

m m q e

miser.

- - - -

, I

Be'e S p e c i e s -- -

Year Plant (s) Visited - -

N u m b e r of Pees C o l l e c t e d 1984 f 9 g 5 - (1984 & 1985 combined)

kab, Hab. Hab. Hab. t j 1 2 3 4 -

B . me Zanopygus x x B. a p i ~ o Z i w n Maius sp. 1 P. a v i m

x x H. syZvaktris I&f,us s p . 1

-- -

Y. o;CficZmZe + 7

Y. mim

B. uppositus x x B. c y l i d r l c a A . m i s e r

-

E . mrfocinc tus x x 24. sylves tr is CastiZZeja miniata

F 3. s a g i t t a t a * 3) . f ,"emrot h s sanguineu s

Bee Species Year A f l, . 1984 1985

;L

(1984 & 1985 co6blne;r) - - Hab. Hab. Hab, Ha,_ -- .-

1 2 3 4 .

% -

* HALICTIDAE b

* - -+

,- .%7

' E a Z b Z u s - B, sagittrrh , 1 ' -L

x . C* i%ipcw&+s . '

. , J I '. -~~of f i - - - inaze$ - 2 1 2 . - 2 .

E,'c6nfusus - x A- x q?". o_F,f<cinate , 2 1 1 ~ 1 ~

"- - -- - - - ~Ckpse~Ia bsa--past--~ _- 5.- _ _ 4--:. - - . f', am-e ,.-/ 4 b .& k i n a . ' t e Z l a t a 1

Elab. Hab. Bab, H a b . - -

1 2 3 4 .. , x ,

E , near f d i

& I". o f , f m m k e

Bee Species Year Plant(s) V i s i t e d N d e r of Bees Collected- .

1984 1985 , (1984 6 1985 codined)

D. near

APPENDIX -- I1 cont.

$he>$des s p . x x P. z6pg<w;ana / - - -

a. officinate 2 . 4 A. negzecfa 1 A. miser 2 h-pittus sp, I '-

--- -- -- .Cnn:ai&

Year Bee Species S) V L s u of 1984 1985 (1984 b 1985 cod ined)

W Hab, bb. -~ab, -- ---- - -

1 2 3 4

1984 1985 (1984 6 1985 combined) -

APPENDIX I1 cont, - ---- - - --

Nab, Hab; Rab; -Hab. - - - -

t 1 2 3 4

0. man- rnantamz .

- -

d i e g a E lesp,_--- x 5"- ~&.&%US , 4 1 . ,

- ---- - - - - - < - * - - - - - - - - -- - - - Eop &i -b$ s x A, miser

i.

1 hflocrita

%

IFte Zf-s mmtana x x 14. q 2 v s t ~ i s -- 1 1

-- - - - -- m-

-- --

Stelis sp. I x x F* offZcinaZe 1

A. ~ L S C T I 1 T* &Eus d 1

r BNTHOrnORIDAE

J h b m d a sp. - -

1 -- - - +--"&7& - - - -

. . h

-- ---

" '9 ; L

Efigerola sp. 1

Me Z Zi ades x E, pk; ledetpfrims 5 con fisa G. m i s & ~ t a 1

Me Zectu rehtiua x E, m*-a 1 \

EpeoZua sp. x x T. cf4&imfe f 5 F - ~~~~ 7 6 . .

\

G. az6akzfx 3 -- - - - - - - -----p E- - q&fd &m 1

@. qzw**:s 1 -

- . Bee Species Year Plant(s) V i s i t e d Number of Bees Collected

19% 1985 (1984 & 1985co&i_rtedL -

s. H a b . HA. . Hab. 1 2- 3 4

" .

R t - T k Q ~ W- % lknlfm k w, Hillnras R C a - W . PC, Clarke. Jr. 1WE%e- - - - - - -- - -

&th&ip between the amount of d e d brood in hmeybei w1onits and their' e

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Fbm, S- W, I, 1 s Salitary bees Sci. Amer. 250: 12&127. - - -- --

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-

Wtnill. R M and- A Dim 1981. The rrspase oi honey bees to v..Uiatio~ls in solar '

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L p --

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6-

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4

C. 1972. Toxidty of field-weathered insecticide residues to four kinds of bees. Envirw. Ent 1: 393394.

- - - - - - --

2377. Pesticides and pdlinators Ann. Rev. Entomd 22: 177-192. - - --

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- - - - - - - - - - - -

- -- - *- - - --

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