jezs 2017; 5(3): 91-102 marginatus williams and granara de ... · imayam institute of agriculture...

~ 91 ~

Journal of Entomology and Zoology Studies 2017; 5(3): 91-102

E-ISSN: 2320-7078

P-ISSN: 2349-6800

JEZS 2017; 5(3): 91-102

© 2017 JEZS

Received: 15-03-2017

Accepted: 16-04-2017

R Nisha

Department of Crop Protection,

Imayam Institute of Agriculture

and Technology, Thuraiyur,

Trichy

JS Kennedy

Department of Agricultural

Entomology, Tamil Nadu

Agricultural University,

Coimbatore, India

Correspondence

R Nisha

Department of Crop Protection,

Imayam Institute of Agriculture

and Technology, Thuraiyur,

Trichy

Life cycle of Papaya mealybug Paracoccus

marginatus Williams and Granara de Willink on

different host plants vis-à-vis divergent natural

selection

R Nisha and JS Kennedy

Abstract Life cycle of Papaya mealybug PMB Paracoccus marginatus on different host plants was studied using

Age Specific (or Horizontal) and Stage Specific (or Vertical) Life Table. It showed enormous biological

activity of PMB on host plants. The net reproductive rate of papaya mealybug was observed to be higher

in papaya (559.48 females/female) and least in tapioca (282.53). The capacity for increase (rc) was

minimum (0.324) in tapioca and maximum in papaya (0.512). Intrinsic rate of increase (rm) has increased

with the increase in the rate of capacity for increase as it was maximum in papaya (0.570 increase per

day), minimum in tapioca (0.342/day). The cumulative K value, total generation mortality was observed

to be minimum in female than male. It was minimum in papaya 0.0325 for female and 0.0587 for male

mealybug, while tapioca recorded highest K value of among the host plants (0.1405 in female and 0.1799

in male).

Keywords: Life cycle, life table, survival, mortality, Papaya mealybug, Paracoccus marginatus, natural

selection, k value, intrinsic rate, net reproductive rate

1. Introduction

A life table is a kind of book-keeping system that ecologists often used to keep track of stage

specific mortality in the population they study [13]. A life table describes for successive age

intervals, the number of deaths, the survivors, the rate of mortality and the expectation of

further life [11]. Life table provides an important tool in understanding the changes in

population of insect pests during different developmental stages throughout their life cycle. It

is an especially useful approach in entomology, where developmental stages are discrete and

mortality rates may vary widely from one life stage to another [7]. It is very useful to analyse

the mortality of insect population, to determine key factors responsible for the highest

mortality within population. Moreover, various mathematical formulas also indicated for the

appropriate evaluation of life fecundity tables, stable age distribution and life expectancy [4].

The construction of several life tables may be possible to prepare a predictive model which can

be tested against natural population fluctuations [12]. Life table is an important analytical

technique in studying distribution, determination of age and mortality of an organism and

individuals can be calculated [11]. On being a polyphagous pest, PMB is feeding on many

plants with divergent variation in development and bioecology [14]. Hence, the present research

was conducted to study the life cycle of PMB on different host plants and it reported for first

time in the current study.

2. Materials and methods

The research work was carried out in the Insectary of Department of Crop Protection, Tamil

Nadu Agricultural University, Coimbatore and Tamil Nadu, India (Department of Agricultural

Entomology and biocontrol laboratory).

2.1. Collection and mass culturing of Paracoccus marginatus

Potato sprouts was used as an alternate food source for rearing mealybugs. Mass culturing of

potato sprouts (Fig. 1) was done in line with the reference of [17]. Papaya mealybugs collected

from different host plants like papaya, tapioca, cotton, mulberry, brinjal and hibiscus were

released on potato sprouts (Fig. 2) using camel hair brush at the rate of 3 to 5 ovisacs per

potato and mealybugs en masse were obtained within 25 to 30 days of release. Mass culturing

~ 92 ~

Journal of Entomology and Zoology Studies

was also carried out in above said host plants and used for

further experiments [2]. The observations needed for the life

cycle studies were documented and calculated as below.

Fig 1: Mass culturing of papaya mealybug Paracoccus marginatus

on potato sprouts

2.2. Description of life table statistics

Life table describes the mortality and survival patterns of a

population. On the basis of mortality ratios for each age or

age group, life tables provide information on parameters such

as the number of survivors, the number of deaths and the life

expectancy [8].

a) Papaya b) Cotton

c) Tapioca d) Mulberry

e) Brinjal f) Hibiscus

g) Potato sprouts

Fig 2: Host plants raised in pots for biology and life cycle studies of

Papaya mealybug Paracoccus marginatus

Column one of the life table gives the age of life from birth to

death. The second column shows the survivorship to each age

of life, starting out at birth (age 0), and diminishing from age

to age in accordance with the mortality. The figures in this

column are generally denoted by the symbol lx. The third

column indicates the corresponding survival fraction (Sx) at

each interval of life, being simply the fraction between lx of

the subsequent stage and lx of the current stage. The fourth

column gives the death rate in each day of life or to be more

exact, the probability at a given age of dying in an interval,

this being denoted by the symbol qx.

The figures in the fifth and sixth column are the total number

of females produced for total population and number of

females produced per female, respectively. The seventh and

eighth columns are auxiliary columns employed in computing

the ninth column, which gives the expectation of life at each

age. The ninth column gives the average number of insects

living in each age of life. The figures in this column may also

be interpreted as the number of days of life lived within a

given age of life. Column eighth is obtained by cumulating

the figures in column seventh beginning at the end. Lastly,

column ninth, gives the expectation of life or the average after

lifetime at each age class of life. It is obtained as the quotient

of the figures in column eighth and the corresponding figures

in column two, for this gives a total number of age class lived

by survivors of a cohort after a given age, divided by the

number of insects entering that age [8]. The 10th and 11th

columns are auxiliary columns used in the calculation of the

12th column denoted as intrinsic rate of natural increase (rm).

2.2.1. Construction of age and stage specific life table

The life tables for insect species were built by partitioning its

life-cycle into distinct development stages (e.g., eggs, larvae,

pupae and adults; eggs, nymphs and adults), and by

evaluating the development time and survival or mortality for

each individual stage. For females, the age-dependent total

oviposition (fecundity/reproduction) was also determined.

The different life table parameters viz., Survivorship (lx) [15],

Survivorship curves [10], Fixation of survivorship curves [16],

Survival fraction (Sx), Apparent mortality, Mortality survivor

ratio (MSR), Indispensable mortality (IM), K-values [18], Net

reproductive rate, Intrinsic rate of natural increase, Finite rate

of increase (λ), Mean generation time (T), Doubling time of

population (t) were calculated as per the earlier experiments [4]

[1, 3, 11].

3. Results and Discussion

Data on vital schedules such as survival, mortality, and

fecundity were collected for papaya mealybug on different

host plants. The observations on fecundity and total number

of females emerged were recorded from the experiment. By

using the above said observations life tables were constructed

for mealybugs from different host plants. The stage specific

life tables were constructed from the data derived from age

specific life tables.

3.1. Age specific life table of Paracoccus marginatus from

different host plants

The data on the age specific life tables of papaya mealybug

from different crops are exhibited in the Tables 1 to 7. The

results revealed that the total life span of adult mealybug was

minimum and last for 28 days in papaya, while it was

maximum in tapioca (42 days). In papaya the production of

off springs (56 females/female) started from the eighth day

and ceased on 18th day of life span with production of 26

~ 93 ~


females per female. The ninth day was the start of

reproduction period in cotton (65 females) and ended with 21

females on 19th day with 30 days of adult longevity. In potato

sprouts, mealybug started reproduction (56 females) from 9th

day and ended up with 22 females on 19th day and the total

life span was lasted for 31 days.

In mulberry PMB recorded 49 days of females on 10th day

and 10 females on 21st day. The adult longevity was 32 days.

The 10th day was the start of reproduction period in brinjal

with production of 44 females per female and ended with 11

females on 22nd day and had the life span of 35 days. In

hibiscus and tapioca, the reproduction of offsprings (31 and

21 females/female) started from the 11th and 13th day and

ceased on 23rd and 26th day of their life span (36 days and 42

days respectively) with reproduction of 5 and 2 females per

female respectively. Papaya, cotton and potato sprouts

recorded higher reproduction of offsprings with lesser

lifespan and production days. Mulberry was marginally same

as that of those crops. Whereas, tapioca, hibiscus and brinjal

recorded lesser production of offsprings with maximum life

span and production days.

Egg - 8 days

First instar (Crawler) 5 days

Total - 3.2 days

Total - 5 days

Second instar

Third instar

Adult Female16.2 days

Total

Life Cycle50 Days

Fig 3: Biology of papaya mealybug Paracocccus marginatus on

papaya

Table 1: Age specific life table of Paracoccus marginatus on papaya

x n lx mx lxmx xlxmx e-rcx e-rc

x. lxmx e-rmx rm e-rmx e-r

mx lxmx

0 100 1 0

1 94 0.94 0

2 90 0.9 0 0 0.00 0.36 0.00 0.32 0.00

3 85 0.85 0 0.00 0.00 0.22 0.00 0.18 0.00

4 79 0.79 0 0.00 0.00 0.13 0.00 0.10 0.00

5 72 0.72 0 0.00 0.00 0.08 0.00 0.06 0.00

6 69 0.69 0 0.00 0.00 0.05 0.00 0.03 0.00

7 62 0.62 0 0.00 0.00 0.03 0.00 0.02 0.00

8 58 0.58 56 32.48 259.84 0.02 0.54 0.01 0.60 0.01 0.34

9 56 0.56 92 51.52 463.68 0.01 0.51 0.00 0.59 0.01 0.30

10 55 0.55 112 61.60 616.00 0.01 0.37 0.00 0.58 0.00 0.21

11 52 0.52 135 70.20 772.20 0.00 0.25 0.00 0.58 0.00 0.13

12 50 0.5 146 73.00 876.00 0.00 0.16 0.00 0.57 0.00 0.08

13 46 0.46 176 80.96 1052.48 0.00 0.10 0.00 0.57 0.00 0.05

14 46 0.46 153 70.38 985.32 0.00 0.05 0.00 0.56 0.00 0.02

15 45 0.45 121 54.45 816.75 0.00 0.03 0.00 0.56 0.00 0.01

16 42 0.42 97 40.74 651.84 0.00 0.01 0.00 0.56 0.00 0.00

17 40 0.4 42 16.80 285.60 0.00 0.00 0.00 0.55 0.00 0.00

18 35 0.35 21 7.35 132.30 0.00 0.00 0.00 0.55 0.00 0.00

19 32 0.32 0 0.00 0.00 0.00 0.00 0.00 0.00

20 26 0.26 0 0.00 0.00 0.00 0.00 0.00 0.00

21 25 0.25 0 0.00 0.00 0.00 0.00 0.00 0.00

22 20 0.2 0 0.00 0.00 0.00 0.00 0.00 0.00

23 20 0.2 0 0.00 0.00 0.00 0.00 0.00 0.00

24 15 0.15 0 0.00 0.00 0.00 0.00 0.00 0.00

25 10 0.1 0 0.00 0.00 0.00 0.00 0.00 0.00

26 10 0.1 0 0.00 0.00 0.00 0.00 0.00 0.00

27 5 0.05 0 0.00 0.00 0.00 0.00 0.00 0.00

28 3 0.03 0 0.00 0.00 0.00 0.00 0.00 0.00

29 0 0 0 0.00 0.00 0.00 0.00 0.00 0.00

Total 559.48 6912.01 2.03 1.15

Table 2: Age specific life table of Paracoccus marginatus on cotton



mx lxmx

0 100 1 0

1 95 0.95 0

0.59

2 90 0.9 0 0.00 0.00 0.39 0.00

0.35 0.00

3 85 0.85 0 0.00 0.00 0.24 0.00

0.21 0.00

4 80 0.8 0 0.00 0.00 0.15 0.00

0.12 0.00

5 75 0.75 0 0.00 0.00 0.09 0.00

0.07 0.00

~ 94 ~


6 75 0.75 0 0.00 0.00 0.06 0.00

0.04 0.00

7 70 0.7 0 0.00 0.00 0.04 0.00

0.03 0.00

8 68 0.68 0 0.00 0.00 0.02 0.00

0.02 0.00

9 62 0.62 65 40.30 362.70 0.01 0.56 0.01 0.54 0.01 0.37

10 59 0.59 89 52.51 525.10 0.01 0.46 0.00 0.54 0.01 0.29

11 50 0.5 117 58.50 643.50 0.01 0.32 0.00 0.53 0.00 0.19

12 50 0.5 124 62.00 744.00 0.00 0.21 0.00 0.53 0.00 0.12

13 46 0.46 165 75.90 986.70 0.00 0.16 0.00 0.52 0.00 0.09

14 42 0.42 142 59.64 834.96 0.00 0.08 0.00 0.52 0.00 0.04

15 40 0.4 128 51.20 768.00 0.00 0.04 0.00 0.52 0.00 0.02

16 35 0.35 112 39.20 627.20 0.00 0.02 0.00 0.51 0.00 0.01

17 35 0.35 97 33.95 577.15 0.00 0.01 0.00 0.51 0.00 0.00

18 30 0.3 64 19.20 345.60 0.00 0.00 0.00 0.51 0.00 0.00

19 28 0.28 21 5.88 111.72 0.00 0.00 0.00 0.51 0.00 0.00

20 22 0.22 0 0.00 0.00 0.00 0.00

0.00 0.00

21 20 0.2 0 0.00 0.00 0.00 0.00

0.00 0.00

22 20 0.2 0 0.00 0.00 0.00 0.00

0.00 0.00

23 18 0.18 0 0.00 0.00 0.00 0.00

0.00 0.00

24 15 0.15 0 0.00 0.00 0.00 0.00

0.00 0.00

25 10 0.1 0 0.00 0.00 0.00 0.00

0.00 0.00

26 8 0.08 0 0.00 0.00 0.00 0.00

0.00 0.00

27 6 0.06 0 0.00 0.00 0.00 0.00

0.00 0.00

28 4 0.04 0 0.00 0.00 0.00 0.00

0.00 0.00

29 2 0.02 0 0.00 0.00 0.00 0.00

0.00 0.00

30 2 0.02 0 0.00 0.00 0.00 0.00

0.00 0.00

31 0 0 0 0.00 0.00 0.00 0.00

0.00 0.00

Total

498.28 6526.63

1.86

1.13

Table 3: Age specific life table of Paracoccus marginatus on tapioca



mx lxmx

0 100 1 0

1 86 0.86 0

2 85 0.85 0 0.00 0.00 0.52 0.00

0.50 0.00

3 80 0.8 0 0.00 0.00 0.38 0.00

0.36 0.00

4 80 0.8 0 0.00 0.00 0.27 0.00

0.25 0.00

5 75 0.75 0 0.00 0.00 0.20 0.00

0.18 0.00

6 70 0.7 0 0.00 0.00 0.14 0.00

0.13 0.00

7 70 0.7 0 0.00 0.00 0.10 0.00

0.09 0.00

8 70 0.7 0 0.00 0.00 0.07 0.00

0.06 0.00

9 68 0.68 0 0.00 0.00 0.05 0.00

0.05 0.00

10 68 0.68 0 0.00 0.00 0.04 0.00

0.03 0.00

11 66 0.66 0 0.00 0.00 0.03 0.00

0.02 0.00

12 65 0.65 0 0.00 0.00 0.02 0.00

0.02 0.00

13 60 0.6 21 12.60 163.80 0.01 0.19 0.01 0.35 0.01 0.15

14 54 0.54 35 18.90 264.60 0.01 0.20 0.01 0.34 0.01 0.16

15 50 0.5 64 32.00 480.00 0.01 0.25 0.01 0.34 0.01 0.19

16 50 0.5 91 45.50 728.00 0.01 0.25 0.00 0.34 0.00 0.19

17 46 0.46 112 51.52 875.84 0.00 0.21 0.00 0.34 0.00 0.15

18 43 0.43 89 38.27 688.86 0.00 0.11 0.00 0.34 0.00 0.08

19 40 0.4 73 29.20 554.80 0.00 0.06 0.00 0.34 0.00 0.04

20 37 0.37 52 19.24 384.80 0.00 0.03 0.00 0.34 0.00 0.02

21 35 0.35 47 16.45 345.45 0.00 0.02 0.00 0.34 0.00 0.01

22 35 0.35 23 8.05 177.10 0.00 0.01 0.00 0.34 0.00 0.00

23 30 0.3 20 6.00 138.00 0.00 0.00 0.00 0.34 0.00 0.00

24 28 0.28 11 3.08 73.92 0.00 0.00 0.00 0.34 0.00 0.00

25 22 0.22 6 1.32 33.00 0.00 0.00 0.00 0.34 0.00 0.00

26 20 0.2 2 0.40 10.40 0.00 0.00 0.00 0.33 0.00 0.00

27 20 0.2 0 0.00 0.00 0.00 0.00

0.00 0.00

28 20 0.2 0 0.00 0.00 0.00 0.00

0.00 0.00

29 15 0.15 0 0.00 0.00 0.00 0.00

0.00 0.00

30 15 0.15 0 0.00 0.00 0.00 0.00

0.00 0.00

31 14 0.14 0 0.00 0.00 0.00 0.00

0.00 0.00



mx lxmx

32 11 0.11 0 0.00 0.00 0.00 0.00

0.00 0.00

33 10 0.1 0 0.00 0.00 0.00 0.00

0.00 0.00

34 10 0.1 0 0.00 0.00 0.00 0.00

0.00 0.00

35 9 0.09 0 0.00 0.00 0.00 0.00

0.00 0.00

36 7 0.07 0 0.00 0.00 0.00 0.00

0.00 0.00

~ 95 ~


37 5 0.05 0 0.00 0.00 0.00 0.00

0.00 0.00

38 5 0.05 0 0.00 0.00 0.00 0.00

0.00 0.00

39 3 0.03 0 0.00 0.00 0.00 0.00

0.00 0.00

40 3 0.03 0 0.00 0.00 0.00 0.00

0.00 0.00

41 1 0.01 0 0.00 0.00 0.00 0.00

0.00 0.00

42 1 0.01 0 0.00 0.00 0.00 0.00

0.00 0.00

43 0 0 0 0.00 0.00 0.00 0.00

0.00 0.00

Total

282.53 4918.57

1.32

0.97

Table 4: Age specific life table of Paracoccus marginatus on mulberry



mx lxmx

0 100 1 0

1 90 0.9 0

2 90 0.9 0 0.00 0.00 0.43 0.00

0.40 0.00

3 86 0.86 0 0.00 0.00 0.28 0.00

0.25 0.00

4 81 0.81 0 0.00 0.00 0.18 0.00

0.16 0.00

5 75 0.75 0 0.00 0.00 0.12 0.00

0.10 0.00

6 74 0.74 0 0.00 0.00 0.08 0.00

0.06 0.00

7 74 0.74 0 0.00 0.00 0.05 0.00

0.04 0.00

8 65 0.65 0 0.00 0.00 0.03 0.00

0.02 0.00

9 62 0.62 0 0.00 0.00 0.02 0.00

0.02 0.00

10 56 0.56 49 27.44 274.40 0.01 0.39 0.01 0.48 0.01 0.27

11 50 0.5 85 42.50 467.50 0.01 0.39 0.01 0.47 0.01 0.26

12 50 0.5 111 55.50 666.00 0.01 0.33 0.00 0.47 0.00 0.22

13 45 0.45 135 60.75 789.75 0.00 0.24 0.00 0.46 0.00 0.15

14 40 0.4 146 58.40 817.60 0.00 0.15 0.00 0.46 0.00 0.09

15 35 0.35 129 45.15 677.25 0.00 0.08 0.00 0.46 0.00 0.04

16 32 0.32 117 37.44 599.04 0.00 0.04 0.00 0.46 0.00 0.02

17 30 0.3 97 29.10 494.70 0.00 0.02 0.00 0.46 0.00 0.01

18 30 0.3 71 21.30 383.40 0.00 0.01 0.00 0.45 0.00 0.01

19 28 0.28 64 17.92 340.48 0.00 0.01 0.00 0.45 0.00 0.00

20 22 0.22 33 7.26 145.20 0.00 0.00 0.00 0.45 0.00 0.00

21 20 0.2 10 2.00 42.00 0.00 0.00 0.00 0.45 0.00 0.00

22 17 0.17 0 0.00 0.00 0.00 0.00

0.00 0.00

23 15 0.15 0 0.00 0.00 0.00 0.00

0.00 0.00

24 15 0.15 0 0.00 0.00 0.00 0.00

0.00 0.00

25 12 0.12 0 0.00 0.00 0.00 0.00

0.00 0.00

26 10 0.1 0 0.00 0.00 0.00 0.00

0.00 0.00

27 8 0.08 0 0.00 0.00 0.00 0.00

0.00 0.00

28 6 0.06 0 0.00 0.00 0.00 0.00

0.00 0.00

29 5 0.05 0 0.00 0.00 0.00 0.00

0.00 0.00

30 3 0.03 0 0.00 0.00 0.00 0.00

0.00 0.00

31 1 0.01 0 0.00 0.00 0.00 0.00

0.00 0.00

32 1 0.01 0 0.00 0.00 0.00 0.00

0.00 0.00

33 0 0 0 0.00 0.00 0.00 0.00

0.00 0.00

Total

404.76 5697.32

1.65

1.07

Table 5: Age specific life table of Paracoccus marginatus on brinjal



mx lxmx

0 100 1 0

1 90 0.9 0

2 90 0.9 0 0.00 0.00 0.43 0.00

0.40 0.00

3 85 0.85 0 0.00 0.00 0.28 0.00

0.25 0.00

4 80 0.8 0 0.00 0.00 0.18 0.00

0.16 0.00

5 75 0.75 0 0.00 0.00 0.12 0.00

0.10 0.00

6 74 0.74 0 0.00 0.00 0.08 0.00

0.06 0.00

7 70 0.7 0 0.00 0.00 0.05 0.00

0.04 0.00

8 66 0.66 0 0.00 0.00 0.03 0.00

0.03 0.00

9 62 0.62 0 0.00 0.00 0.02 0.00

0.02 0.00

10 57 0.57 44 25.08 250.80 0.01 0.36 0.01 0.47 0.01 0.26

11 55 0.55 79 43.45 477.95 0.01 0.41 0.01 0.47 0.01 0.29

12 50 0.5 93 46.50 558.00 0.01 0.28 0.00 0.46 0.00 0.19

13 46 0.46 112 51.52 669.76 0.00 0.21 0.00 0.46 0.00 0.14

14 46 0.46 125 57.50 805.00 0.00 0.15 0.00 0.46 0.00 0.10

15 45 0.45 104 46.80 702.00 0.00 0.08 0.00 0.45 0.00 0.05

16 40 0.4 88 35.20 563.20 0.00 0.04 0.00 0.45 0.00 0.02

17 37 0.37 79 29.23 496.91 0.00 0.02 0.00 0.45 0.00 0.01

18 37 0.37 41 15.17 273.06 0.00 0.01 0.00 0.45 0.00 0.00

~ 96 ~


19 33 0.33 22 7.26 137.94 0.00 0.00 0.00 0.45 0.00 0.00

20 30 0.3 11 3.30 66.00 0.00 0.00 0.00 0.45 0.00 0.00

21 25 0.25 5 1.25 26.25 0.00 0.00 0.00 0.45 0.00 0.00

22 20 0.2 0 0.00 0.00 0.00 0.00

0.00 0.00

23 20 0.2 0 0.00 0.00 0.00 0.00

0.00 0.00

24 17 0.17 0 0.00 0.00 0.00 0.00

0.00 0.00

25 15 0.15 0 0.00 0.00 0.00 0.00

0.00 0.00

26 15 0.15 0 0.00 0.00 0.00 0.00

0.00 0.00

27 12 0.12 0 0.00 0.00 0.00 0.00

0.00 0.00

28 10 0.1 0 0.00 0.00 0.00 0.00

0.00 0.00

29 8 0.08 0 0.00 0.00 0.00 0.00

0.00 0.00

30 7 0.07 0 0.00 0.00 0.00 0.00

0.00 0.00

31 7 0.07 0 0.00 0.00 0.00 0.00

0.00 0.00

32 5 0.05 0 0.00 0.00 0.00 0.00

0.00 0.00

33 5 0.05 0 0.00 0.00 0.00 0.00

0.00 0.00

34 3 0.03 0 0.00 0.00 0.00 0.00

0.00 0.00

35 1 0.01 0 0.00 0.00 0.00 0.00

0.00 0.00

36 0 0 0 0.00 0.00 0.00 0.00

0.00 0.00

Total

362.26 5026.87

1.56

1.07

Table 6: Age specific life table of Paracoccus marginatus on hibiscus



mx lxmx

0 100 1 0

1 85 0.85 0

2 85 0.85 0 0.00 0.00 0.46 0.00 0.44 0.00

3 83 0.83 0 0.00 0.00 0.31 0.00 0.29 0.00

4 80 0.8 0 0.00 0.00 0.21 0.00 0.19 0.00

5 73 0.73 0 0.00 0.00 0.14 0.00 0.13 0.00

6 70 0.7 0 0.00 0.00 0.10 0.00 0.09 0.00

7 70 0.7 0 0.00 0.00 0.07 0.00 0.06 0.00

8 65 0.65 0 0.00 0.00 0.05 0.00 0.04 0.00

9 65 0.65 0 0.00 0.00 0.03 0.00 0.03 0.00

10 60 0.6 0 0.00 0.00 0.02 0.00 0.02 0.00

11 54 0.54 31 16.74 184.14 0.01 0.24 0.01 0.42 0.01 0.19

12 50 0.5 66 33.00 396.00 0.01 0.32 0.01 0.41 0.01 0.24

13 45 0.45 98 44.10 573.30 0.01 0.29 0.00 0.41 0.00 0.22

14 45 0.45 119 53.55 749.70 0.00 0.24 0.00 0.41 0.00 0.18

15 45 0.45 127 57.15 857.25 0.00 0.17 0.00 0.41 0.00 0.12

16 40 0.4 81 32.40 518.40 0.00 0.07 0.00 0.41 0.00 0.05

17 35 0.35 69 24.15 410.55 0.00 0.03 0.00 0.41 0.00 0.02

18 33 0.33 41 13.53 243.54 0.00 0.01 0.00 0.40 0.00 0.01

19 33 0.33 29 9.57 181.83 0.00 0.01 0.00 0.40 0.00 0.00

20 30 0.3 19 5.70 114.00 0.00 0.00 0.00 0.40 0.00 0.00

21 27 0.27 11 2.97 62.37 0.00 0.00 0.00 0.40 0.00 0.00

22 21 0.21 10 2.10 46.20 0.00 0.00 0.00 0.40 0.00 0.00

23 20 0.2 5 1.00 23.00 0.00 0.00 0.00 0.40 0.00 0.00

24 18 0.18 0 0.00 0.00 0.00 0.00 0.00 0.00

25 14 0.14 0 0.00 0.00 0.00 0.00 0.00 0.00

26 14 0.14 0 0.00 0.00 0.00 0.00 0.00 0.00

27 12 0.12 0 0.00 0.00 0.00 0.00 0.00 0.00

28 10 0.1 0 0.00 0.00 0.00 0.00 0.00 0.00

29 8 0.08 0 0.00 0.00 0.00 0.00 0.00 0.00

30 7 0.07 0 0.00 0.00 0.00 0.00 0.00 0.00

31 7 0.07 0 0.00 0.00 0.00 0.00 0.00 0.00

32 5 0.05 0 0.00 0.00 0.00 0.00 0.00 0.00

33 5 0.05 0 0.00 0.00 0.00 0.00 0.00 0.00

34 3 0.03 0 0.00 0.00 0.00 0.00 0.00 0.00

35 3 0.03 0 0.00 0.00 0.00 0.00 0.00 0.00

36 3 0.03 0 0.00 0.00 0.00 0.00 0.00 0.00

37 0 0 0 0.00 0.00 0.00 0.00 0.00 0.00

Total 295.96 4360.28 1.39 1.03

Table 7: Age specific life table of Paracoccus marginatus on potato sprouts



mx lxmx

0 100 1 0

1 92 0.92 0

2 85 0.85 0 0.00 0.00 0.39 0.00

0.36 0.00

3 80 0.8 0 0.00 0.00 0.25 0.00

0.22 0.00

~ 97 ~


4 80 0.8 0 0.00 0.00 0.15 0.00

0.13 0.00

5 77 0.77 0 0.00 0.00 0.10 0.00

0.08 0.00

6 74 0.74 0 0.00 0.00 0.06 0.00

0.05 0.00

7 74 0.74 0 0.00 0.00 0.04 0.00

0.03 0.00

8 65 0.65 0 0.00 0.00 0.02 0.00

0.02 0.00

9 62 0.62 56 34.72 312.48 0.01 0.51 0.01 0.53 0.01 0.35

10 56 0.56 79 44.24 442.40 0.01 0.41 0.01 0.53 0.01 0.26

11 50 0.5 111 55.50 610.50 0.01 0.32 0.00 0.52 0.00 0.20

12 50 0.5 124 62.00 744.00 0.00 0.23 0.00 0.52 0.00 0.13

13 45 0.45 165 74.25 965.25 0.00 0.17 0.00 0.51 0.00 0.10

14 40 0.4 143 57.20 800.80 0.00 0.08 0.00 0.51 0.00 0.04

15 35 0.35 132 46.20 693.00 0.00 0.04 0.00 0.51 0.00 0.02

16 32 0.32 112 35.84 573.44 0.00 0.02 0.00 0.50 0.00 0.01

17 30 0.3 97 29.10 494.70 0.00 0.01 0.00 0.50 0.00 0.00

18 30 0.3 59 17.70 318.60 0.00 0.00 0.00 0.50 0.00 0.00

19 28 0.28 22 6.16 117.04 0.00 0.00 0.00 0.50 0.00 0.00

20 22 0.22 0 0.00 0.00 0.00 0.00

0.00 0.00

21 20 0.2 0 0.00 0.00 0.00 0.00

0.00 0.00

22 17 0.17 0 0.00 0.00 0.00 0.00

0.00 0.00

23 15 0.15 0 0.00 0.00 0.00 0.00

0.00 0.00

24 15 0.15 0 0.00 0.00 0.00 0.00

0.00 0.00

25 12 0.12 0 0.00 0.00 0.00 0.00

0.00 0.00

26 10 0.1 0 0.00 0.00 0.00 0.00

0.00 0.00

27 8 0.08 0 0.00 0.00 0.00 0.00

0.00 0.00

28 6 0.06 0 0.00 0.00 0.00 0.00

0.00 0.00

29 5 0.05 0 0.00 0.00 0.00 0.00

0.00 0.00

30 3 0.03 0 0.00 0.00 0.00 0.00

0.00 0.00

31 1 0.01 0 0.00 0.00 0.00 0.00

0.00 0.00

32 0 0.00 0 0.00 0.00 0.00 0.00

0.00 0.00

Total

462.91 6072.21

1.80

1.12

The parameters of age specific life table on different host

plants are summarized in Table 9. Determining the life cycle

of an insect was important to understand its development,

distribution and abundance. In polyphagous insects, life cycle

can vary with the plant species it feeds on. There were

differences in the life history parameters of P. marginatus and

thereby able to develop, survive and reproduce on different

hosts. The life cycle of P. marginatus was reported for first

time in the current study and it varied accordingly to the host

plants. While observing the age specific life table of the pest

in the present study, it has been concluded that the net

reproductive rate of papaya mealybug was observed to be

higher in papaya (559.48 females/female) and followed by

cotton with 498.28 females. Whereas in tapioca it had the

least net reproductive rate of (282.53). The capacity for

increase (rc) was minimum (0.324) in tapioca and maximum

in papaya (0.512) followed by cotton (0.474) and potato

sprouts (0.427). Intrinsic rate of increase (rm) has increased

with the increase in the rate of capacity for increase in all the

host plants by following same trend as that of rc value. It was

maximum in papaya (0.570 increase per day), while recording

minimum in tapioca (0.342/day). Mealybug had the longest

doubling time in tapioca (2.028 days) followed by hibiscus

(1.696 days), while recorded shortest time in papaya (1.216

days). This is because of the innate capacity of P. marginatus

to live shorter and reproduce more in hosts, in which they are

adapted.

The results were in contrast with report of [2] Amarasekare et

al. (2008), who found out that mealybug eggs survived

similarly on the four plants plumeria, acalypha, hibiscus and

parthenium. The lower survival of the first and second instars

was reflected in plumeria and the survival for the third-instar

males and females, and the fourth-instar males were not

affected by the host species. They also concluded that adults

emerged on plumeria with a higher proportion of females than

on the other three host species.

3.1.1. Survivorship curve of Paracoccus marginatus on


The survival exhibited by P. marginatus indicated that it

belongs to type III survivorship curve. In general, survival

decreased with increasing in days. The curve indicated that

the mortality during early stage of the pest was higher at

higher in tapioca. Tapioca recorded 50 per cent mortality at

16h day whereas other crops it was on 12th day of their

longevity (Fig.1.). Using DUD’s method, survivorship curves

of P. marginatus on different host plants were smoothened.

Parameters (a and b) of the smoothened curves of different

host plants are given in the Table 8.

Table 8: Response of survival of Paracoccus marginatus on


Host plants ‘a’ (Days of 50%

mortality)

‘b’

(Intercept)

r2

value

Papaya 12 2.180 0.783

Cotton 12 0.922 0.983

Tapioca 16 0.864 0.973

Mulberry 12 0.894 0.967

Brinjal 12 0.883 0.972

Hibiscus 12 0.854 0.965

Potato sprouts 12 0.899 0.974

~ 98 ~


Fig 4: Survivorship curve of Paracoccus marginatus on different host plants

Table 9: Life table parameters of Paracoccus marginatus on different host plants

Parameter Papaya Cotton Tapioca Mulberry Brinjal Hibiscus Potato sprouts

Age of first oviposition (days) 8 9 13 10 10 11 9

Age of last oviposition (days) 18 21 26 21 22 23 22

Length of oviposition (days) 11 13 14 12 13 13 14

Net Reproductive rate (R0) (females/female) 559.48 498.28 282.53 404.76 362.26 295.96 462.91

Approx generation time (Tc) days 12.35 13.10 17.41 14.08 13.88 14.73 13.12

Capacity for increase (rc) 0.512 0.474 0.324 0.427 0.425 0.386 0.468

Intrinsic rate of increase (rm) per day 0.570 0.521 0.342 0.462 0.457 0.409 0.512

Mean generation time (T) (days) 11.10 11.92 16.51 12.98 12.91 13.92 11.98

Finite rate of increas (λ) per day 1.768 1.684 1.407 1.588 1.579 1.505 1.669

Doubling time (t) days 1.216 1.330 2.028 1.499 1.518 1.696 1.353

3.2. Stage specific life table of Paracoccus marginatus on


The data on the stage specific life table of papaya mealybug

on different crops was exhibited in the Tables (10 - 24). The

data revealed that mealybug had female biased sex ratio in the

production of offsprings. The sex ratio of mealybug was

estimated in the second instar and the ratio was back

calculated in egg and first instar stages. The stage specific life

table parameters involving survivals and mortality in each

stage of both female and male mealybug on papaya are

exhibited in the tables 10 and 11. The analysis was started

with 890 and 175 eggs in female and male mealybugs

respectively and finally 660 and 89 emerged successfully as

adults respectively as females and males on papaya host and

760 eggs in females and 205 eggs in male mealybugs in

cotton.

Subsequently it started with 299 and 246 eggs in female and

male mealybugs respectively and finally 82 and 31 were

emerged successfully as adults respectively in females and

males on tapioca and with 610 and 315 eggs in female and

male mealybugs respectively and finally 356 and 85 were

emerged successfully as adults respectively in female and

male on mulberry. It started with 526, 399 and 708 eggs in

female and 309, 296 and 242 eggs in male mealybug in

brinjal, hibiscus and potato sprouts respectively.

Survival proportion and Survival fraction (Sx) of female

mealybug was maximum 74.16 and 0.91 per cent respectively

in papaya and minimum in tapioca 27.42 and 0.76 per cent

respectively and in same male melaybugs recorded 50.86 and

0.74 per cent and 12.60 per cent and 0.75 respectively. The

highest apparent mortality was recorded in tapioca by

mealybugs than other host plants. While comparing instars in

female mealybugs, the lower apparent mortality was observed

in first instars (23.65) and the second instars had the highest

mortality of 30.32. In male mealybugs, it was maximum in

fourth instars (50.00). In female mealybugs, lowest apparent

mortality was observed in first instar (6.14) and the third

instar had the highest mortality of 9.14 among the instars in

papaya. In male mealybugs, it was maximum in fourth instar

(26.45), and minimum in first instar (5.14).

Mortality survivor ratio (MSR) was estimated to understand

the amount of population increase in a particular stage, if,

mortality in the particular stage had not occurred. In egg stage

MSR was observed to be marginally same (0.47 and 0.48

respectively) in female and male in tapioca. It was found to be

observed that MSR in second instars was 0.31 in female and

0.34 in male. MSR was more in male mealybugs (0.18 and

0.36 in third and fourth instars respectively) than females

(0.10 in third instar). In other host plants also, MSR was

recorded synchronically. Indispensable mortality (IM) was

calculated to understand the amount of mortality, which can

be avoided if the factor causing mortaliy is not allowed to

operate. The examination of mealybugs revealed that, it was

more in females than males in all the stages. In females, it was

maximum in papaya 47.71 to 67 and minimum in tapioca

25.39 to 38.78 and in males, it was maximum in tapioca 10.50

to 31.00 and minimum in papaya 2.08 to 2.24

In the stage specific life table of the current study, survival

~ 99 ~


proportion, survival fraction (Sx), apparent mortality,

mortality survivor ratio (MSR), indispensable mortality (IM)

and K – value were recorded the significant difference in all

the host plants. There was similarity observed that all the host

plants recorded the female biased sex ratio but different ratio

among the host plants, which was reported earlier by [2] the

author. The loss of first instar P. marginatus was observed in

the current study, might be due to the movement of crawlers

(first instars) away from the leaf tissues and they falling off

the plants. This movement was observed on all plants,

although it was more obviously evidenced by [2] the author,

who reported a loss of 17 to 18% of the first instars mealybug

on hibiscus, acalypha, and parthenium. Preliminary studies

demonstrated that the crawlers of P. marginatus, which were

dislodged from the leaves, were not be able to survive, unless

they moved back or were placed back on the leaf.

In the present study, there were differences observed in the

adult longevity of mealybugs on host plants. However, [2]

reported no difference in adult longevity of males and females

occurred on different hosts. However, survival proportion and

fraction of mealybug was maximum in papaya and minimum

in tapioca and hibiscus, whereas the mortality ratio was

maximum in tapioca and hibiscus and minimum in papaya.

The total generation mortality K-value was observed to be

minimum in female mealybugs than male (Table 24). It was

minimum in papaya 0.0325 for female mealybugs and 0.0587

for male mealybugs, while tapioca recorded highest K value

of 0.1799 among the host plants (0.1405 in female and 0.1799

in male) and it was followed by hibiscus in both female and

male pest (0.0959 and 0.1478 respectively).

Table 10: Stage specific life table of female Paracoccus marginatus on papaya

Stage lx dx Survival proportion Survival fraction (Sx) Apparent mortality MSR IM K- value

Egg 890 60 100 0.93 6.74 0.07 47.71 0.030

Instar I 830 51 93.26 0.94 6.14 0.07 43.21 0.028

Instar II 779 52 87.53 0.93 6.68 0.07 47.21 0.030

Instar III 727 67 81.69 0.91 9.22 0.10 67.00 0.042

Adult 660 660 74.16 0.00 100.00 0.00

Average 0.033

Table 11: Stage specific life table of male Paracoccus marginatus on papaya


Egg 175 12 100 0.93 6.86 0.07 2.24 0.031

Instar I 163 9 93.14 0.94 5.52 0.06 2.21 0.025

Instar II 154 11 88.00 0.93 7.14 0.08 2.19 0.032

Instar III 143 22 81.71 0.85 15.38 0.18 2.16 0.073

Instar IV 121 32 69.14 0.74 26.45 0.36 2.08 0.133

Adult 89 89 50.86 0.00 100.00

Average 0.059

Table 12: Stage specific life table of female Paracoccus marginatus on cotton

Stage lx dx Survival proportion Survival fraction Sx Apparent mortality MSR IM K- value

Egg 760 90 100 0.88 11.84 0.13 65.69 0.0547

Instar I 670 70 88.16 0.90 10.45 0.12 57.05 0.0479

Instar II 600 75 78.95 0.88 12.50 0.14 69.86 0.0580

Instar III 525 36 69.08 0.93 6.86 0.07 36.00 0.0309

Adult 489 489 64.34 0.00 100.00

Average 0.0479

Table 13: Stage specific life table of male Paracoccus marginatus on cotton


Egg 205 25 100 0.88 12.20 0.14 11.53 0.0565

Instar I 180 19 87.80 0.89 10.56 0.12 9.80 0.0484

Instar II 161 20 78.54 0.88 12.42 0.14 11.77 0.0576

Instar III 141 29 68.78 0.79 20.57 0.26 21.49 0.1000

Instar IV 112 29 54.63 0.74 25.89 0.35 29.00 0.1301

Adult 83 83 40.49 100.00

Average 0.0785

Table 14: Stage specific life table of female Paracoccus marginatus on tapioca


Egg 299 96 100 0.68 32.11 0.47 38.78 0.1682

Instar I 203 48 67.89 0.76 23.65 0.31 25.39 0.1172

Instar II 155 47 51.84 0.70 30.32 0.44 35.69 0.1569

Instar III 108 26 36.12 0.76 24.07 0.32 26.00 0.1196

Adult 82 82 27.42 0 100.00

Average 0.1405

~ 100 ~


Table 15: Stage specific life table of male Paracoccus marginatus on tapioca


Egg 246 80 100 0.67 32.52 0.48 14.94 0.1708

Instar I 166 42 67.48 0.75 25.30 0.34 10.50 0.1267

Instar II 124 36 50.41 0.71 29.03 0.41 12.68 0.1489

Instar III 88 26 35.77 0.70 29.55 0.42 13.00 0.1521

Instar IV 62 31 25.20 0.50 50.00 1.00 31.00 0.3010

Adult 31 31 12.60 0.00 100.00

Average 0.1799

Table 16: Stage specific life table of female Paracoccus marginatus on mulberry


Egg 610 89 100 0.85 14.59 0.17 60.81 0.0685

Instar I 521 42 85.41 0.92 8.06 0.09 31.22 0.0365

Instar II 479 75 78.52 0.84 15.66 0.19 66.09 0.0740

Instar III 404 48 66.23 0.88 11.88 0.13 48.00 0.0549

Adult 356 356 58.36 0.00 100.00

Average 0.0585

Table 17: Stage specific life table of male Paracoccus marginatus on mulberry


Egg 315 46 100 0.85 14.60 0.17 14.54 0.0686

Instar I 269 27 85.40 0.90 10.04 0.11 9.48 0.0459

Instar II 242 34 76.83 0.86 14.05 0.16 13.89 0.0658

Instar III 208 80 66.03 0.62 38.46 0.63 53.13 0.2109

Instar IV 128 43 40.63 0.66 33.59 0.51 43.00 0.1778

Adult 85 85 26.98 0.00 100.00

Average 0.1138

Table 18: Stage specific life table of female Paracoccus marginatus on brinjal


Egg 526 64 100 0.88 12.17 0.14 40.87 0.0563

Instar I 462 64 87.83 0.86 13.85 0.16 47.44 0.0648

Instar II 398 60 75.67 0.85 15.08 0.18 52.37 0.0710

Instar III 338 43 64.26 0.87 12.72 0.15 43.00 0.0591

Adult 295 295 56.08 0.00 100.00

Average 0.0628

Table 19: Stage specific life table of male Paracoccus marginatus on brinjal


Egg 309 37 100 0.88 11.97 0.14 9.79 0.0554

Instar I 272 41 88.03 0.85 15.07 0.18 12.78 0.0710

Instar II 231 33 74.76 0.86 14.29 0.17 12.00 0.0669

Instar III 198 75 64.08 0.62 37.88 0.61 43.90 0.2068

Instar IV 123 51 39.81 0.59 41.46 0.71 51.00 0.2326

Adult 72 72 23.30 0.00 100.00

Average 0.1265

Table 20: Stage specific life table of female Paracoccus marginatus on hibiscus


Egg 399 70 100 0.82 17.54 0.21 35.11 0.0838

Instar I 329 64 82.46 0.81 19.45 0.24 39.85 0.0940

Instar II 265 74 66.42 0.72 27.92 0.39 63.93 0.1422

Instar III 191 26 47.87 0.86 13.61 0.16 26.00 0.0635

Adult 165 165 41.35 0.00 100.00

Average 0.0959

Table 21: Stage specific life table of male Paracoccus marginatus on hibiscus


Egg 296 52 100 0.82 17.57 0.21 11.51 0.0839

Instar I 244 47 82.43 0.81 19.26 0.24 12.88 0.0929

Instar II 197 56 66.55 0.72 28.43 0.40 21.45 0.1452

Instar III 141 46 47.64 0.67 32.62 0.48 26.15 0.1715

Instar IV 95 41 32.09 0.57 43.16 0.76 41.00 0.2453

Adult 54 54 18.24 0.00 100.00

Average 0.1478

~ 101 ~


Table 22: Stage specific life table of female Paracoccus marginatus on potato sprouts


Egg 708 104 100 0.85 14.69 0.17 74.38 0.0690

Instar I 604 62 85.31 0.90 10.26 0.11 49.42 0.0470

Instar II 542 68 76.55 0.87 12.55 0.14 61.97 0.0582

Instar III 474 42 66.95 0.91 8.86 0.10 42.00 0.0403

Adult 432 432 61.02 100.00

Average 0.0536

Table 23: Stage specific life table of male Paracoccus marginatus on potato sprouts


Egg 242 36 100 0.85 14.88 0.17 15.90 0.0699

Instar I 206 20 85.12 0.90 9.71 0.11 9.78 0.0444

Instar II 186 24 76.86 0.87 12.90 0.15 13.48 0.0600

Instar III 162 39 66.94 0.76 24.07 0.32 28.85 0.1196

Instar IV 123 32 50.83 0.74 26.02 0.35 32.00 0.1309

Adult 91 91 37.60 100.00

Average 0.0850

Table 24: K value of Paracoccus marginatus on different host plants

Host crop

K value Cumulative k value

Egg period First instar Second instar Third instar Fourth instar

Male Female Male Female Male Female Male Female Male Male Female

Papaya 0.031 0.030 0.025 0.028 0.032 0.030 0.073 0.042 0.133 0.059 0.033

Cotton 0.057 0.055 0.048 0.048 0.058 0.058 0.100 0.031 0.130 0.079 0.048

Tapioca 0.171 0.168 0.127 0.117 0.149 0.157 0.152 0.120 0.301 0.180 0.141

Mulberry 0.069 0.069 0.046 0.037 0.066 0.074 0.211 0.055 0.178 0.114 0.059

Brinjal 0.055 0.056 0.071 0.065 0.067 0.071 0.207 0.059 0.233 0.127 0.063

Hibiscus 0.084 0.084 0.093 0.094 0.145 0.142 0.172 0.064 0.245 0.148 0.096

Potato 0.070 0.069 0.044 0.047 0.060 0.058 0.120 0.040 0.131 0.085 0.054

When comparing six different host plants, tapioca and

hibiscus recorded maximum the finite rate of mortality of egg

and I instar larvae. This may be attributed to variation in the

host nutrition, developmental conditions and vulnerability of

young ones. Host plant species have been shown to affect the

life history parameters of several mealybug species. Longer

pre-reproductive period and a higher progeny production were

observed in Rastrococcus invadens Williams reared on

different varieties of Mangifera indica L. [5]. The differences

observed in the life history of P. marginatus may be related to

nutritive factors, allelochemical compounds and physical

differences in leaf structures, etc. Mortality of the of citrus

mealybug Planococcus citri (Risso) was higher on green than

on red or yellow variegated Coleus blumei "Bellevue"

(Bentham) plants, and developed faster and had a higher

fecundity when developed on red-variegated plants [19].

Food availability and quality are other environmental factors

that we expect to affect life history traits, independent of

whether. These effects have been altered or caused by natural

selection in a specified environment [9]. The difference in the

total life cycle might be due to the type of host used for the

study [6]. Life history of P. marginatus was affected by the

host plants. However, it has the ability to develop, survive,

and reproduce on a variety of host plant species.

4. Conclusion

The information gathered from the life history study will be

important in the management of P. marginatus, by providing

a better understanding of its life cycle, and its ability to

survive on different host plant species. This information is

needed in the development of integrated pest management of

this pest. The ability of P. marginatus to develop on these

plant species demonstrates the possibility of movement,

distribution, and establishment of P. marginatus into new

areas.

5. Acknowledgement

Authors are thankful to the Department of Agricultural

Entomology, Tamil Nadu Agricultural University,

Coimbatore for the great support during the research period

and to the University Grand Commission, New Delhi for

providing fellowship for the research work.

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