Res. Popul. Ecol. (1989) 31, 25--34. ~) by the Society of Population Ecology

S U R V I V O R S H I P AND F E R T I L I T Y SCHEDULES OF T W O S U M A T R A N

T O R T O I S E BEETLES, ASPIDOMORPHA M I L I A R I S AND A. SANC-

TAECRUCIS ( C O L E O P T E R A : C H R Y S O M E L I D A E ) U N D E R

L A B O R A T O R Y C O N D I T I O N S 1' 2, 3

Koji NAKAMURA*, Idrus ABBAS** and Ahsol HASYIM**' r

*Ecological Laboratory, Faculty of Science, Kanazawa University, Kanazawa 920, Japan **Department of Biology, Faculty of Science, Andalas University, Padang, Sumatera Barat, Indonesia

INTRODUCTION

Lati tudinal gradients in selective forces acting to shape the life history traits of

organisms have long been argued (e.g., DOBZHANSKY, 1950; MACARTHUR, 1972). In-

ter- and intraspecific comparisons of life table parameters between temperate and

tropical populations were made by laboratory rearings in a variety of insect groups:

milkweed bugs, Oncopeltus (LANDAHL and ROOT, 1969; DINGLE, 1978, 1981; DINGLE

and BALDWIN, 1983; BALDWIN and DINGLE, 1986), fruit flies, Drosophila (BIRCH et al.,

1963; BOUL~TREAU-MERLE et al., 1982; DAVID et al., 1984) and leafeating ladybirds,

Epilachna (NAI~MUgA et al., 1984; ABBAS et al., 1985).

This article describes survivorship and fertility schedules of two species of tortoise

beetle, Aspidomorpha miliaris (FABRICIUS) (AM) and A. sanctaecrucis (FABRICIUS) (AS)

which feed on Ipomoea carnea (Convolvulaceae) under laboratory conditions in Padang,

Sumatra. The two species are large in size (9-15 mm long in AM and 9-12 mm in

AS), very common and widespread throughout Southeast Asia as pests of Ipomoea crops

such as sweet potato, L batatas (KALSHOVEN, 1981). In the preceding articles, we

presented the life table of AM (NAgAMURA and ABBAS, 1987b) and the seasonal fluctua-

tion of adult numbers and egg mortalities of field populations of the two species

(NAKAMURA and ABBAS, 1987a, in press; see these articles for relevant publications).

MATERIALS AND METHODS

Climate

Padang lies on the equator (0~ 100~ and has a predominant ly humid-

i Contributions to the knowledge of population dynamics of tortoise beetles in Sumatra 3. 2 Contribution No. 33 of Sumatra Nature Study (Entomology). 3 Partly supported by Grants from Japan Society for Promotion of Science for JSPS-DGHE Scientific

Cooperation (1980, 1982) and Grants-in-Aid for Overseas Scientific Survey from Ministry of Educa- tion, Science and Culture of Japan (Nos. 56041027 and 58041030).

4 Present address: Sukarami Research Institute for Food Crops, P.O. Box 34, Padang, Sumatera Barat, Indonesia.

26

equatorial climate: the mean monthly temperature fluctuated only between 26.7

(September to December) and 27.5~ (May) and the annual rainfall was 4764 mm

(OGINO, 1984) without distinct cycles of wet and dry seasons.

Host plant

The host plant, L carnea is a shrubby morning glory which grows as a weed in

moist soil. Its stem is robust enough to stand erect, but when its height exceeds 200-

250 cm, it tends to fall to the ground by its own weight. The plant grows very rapidly

by asexual propagation (NAKAMURA and ABBAS, 1987b).

Experiments

The following experiments were carried out from 20 December 1981 to 23

February 1982 in the Guest House of Andalas University and thereafter until 19 June

1982 in the Sumatra Nature Study Laboratory of Andalas University, both in Padang,

under room temperatures ranging from 24 to 32~

Experiment 1. Adults of the two species were collected from L carnea in Padang

and reared in the laboratory to obtain eggs for this experiment. Each egg mass was

isolated in a plastic cup (13 cm in diameter and 5 cm in depth) with fresh leaves of L

carnea, and records were kept daily of hatching and of larval molts in order to know

the developmental time. This experiment was done in the Guest House, using 15

cups for both species.

Experiment 2. A pair of newly emerged adults obtained in Experiment 1 was con-

fined in a plastic cup of the same size as in Experiment 1 to learn their survivorship

and fertility schedules. Thirteen (AM) and nine (AS) pairs were reared with daily

checking and exchange of the food plant leaves during the study period. Eggs were

laid in clusters within a paper substance (ootheca) on the undersurface of the host plant

leaves and remained there after hatching (NAKAMURA and ABBAS, 1987b). All egg

masses were dissected after hatching under a microscope to determine the number of eggs per mass.

RESULTS

Developmental times of the immature stages

The results of Experiment 1 shows that the duration of successive immature stages

(in days) was 34-39 days in AM and 30-37 days in AS (Table 1). Developmental

times of AM so far reported were: 35 days in the Philippines (SeHuLTZE, 1908) and

41 days in Malay Peninsula (CogBETT and DOVER, 1927).

Elytral color change with age

The ground color of elytra of the two species reared in Experiment 2 changed

Table 1. Duration of successive immature stages (in days) of A. miliaris and A. sanctaecrucis.

Species Egg L1 L2 L3 L4 L5 Pupa Total

A. miliaris 10-12 3 3 3-4 3-4 7 5-6 34-39 A. sanctaecrucis 7-9 4 2-3. 3 3 6-8 5-7 30-37

27

remarkably with age. Though the change proceeded gradually, four and three color

phases were recognized in AM and AS, respectively (Table 2). The color change pro-

vided a rough measure for determining the age of adults in the field study (see Discus-

sion).

Egg mass size

Figure 1 shows the frequency distribution of egg mass size, indicating that the

mean size was 37.2 (range: 12-73) in AM and 10.9 (range: 3-18) in AS. These values

were somewhat smaller than those observed in field populations (43.4 in AM,

NAI~MURA and A~BAS, 1987b; 43.4 in AM and 15.7 in AS, NAKAMURA and ABBAS,

1989), possibly because of the limited space in the cups.

Survivorship and fertility schedules

No record Was kept of sex ratio of newly emerged adults in the present rearings.

In field populations of the two species in the garden of the Sumatra Nature Study

Laboratory, newly emerged adults showed no significant deviation from the expected

1 : 1 sex ratio (binomial test, 49 females : 48 males in AM and 123 females : 138 males

in AS; NAg,~MURA et al., in preparation). Therefore, a 1 : 1 sex ratio is assumed in the

calculation below.

Figures 2 and 3 show the survivorship (lx) of female and male adults and the age

specific fertility, which is expressed as the change in the number of eggs laid per female

per day. On the basis of the expected 1 : 1 sex ratio, the age-specific fertility in these

figures is equivalent to mx • 2 (mx is usually defined as the number of living females

born per female per unit time, SOUTHWOOD, 1978). The shape of the histogram giving

the age-specific fertility shows that females produced eggs at a nearly constant rate

throughout their long reproductive period (Figs. 2 and 3). Table 3 summarizes the

Table 2. Change in ground color of elytra in A. miliaris and A. sanctaecrucis.

Age (in days) Phase Color Species after emergence

A. miliaris 0- I White

12.8- 11" Yellowish white 20.7- III Yellow 74.0- 1V Deep yellow/orange

A. sanctaecrucis 0- I Brown

23.6- 11 Golden 59.8- Ill Reddish gold

28

50-

4 0 -

30-

20-

~, l O

o , ,

o- 30- Q) L

EL. 2 0 -

10-

A. sonctoecruc is

n = 4 3

= 10.9+_1.3

I I I I I I

A. m / l i o r i s n = 1:56 = :57.2 _+ 2 . 0

I

0 80 I I I I I I I

10 20 30 4 0 50 6 0 70

No. of eggs per mass

Fig. 1. Frequency distribution in the size of egg masses of A. miliaris (bottom) and A. sanc- taecrucis (top). The number of masses examined (n) and values of mean (~) with 95% c o n - fidence limits are shown.

R S I, ,I I( 11 | | i i

, , d I

~, . . . . . ~ " 0 Z ,oo, ' , , . . . . . . ri~ o ~8~ ~ ~ m,,,oos t

J ,,,kl t 0 II I . . . . . . . . . . . . ",, 0

0 10 20 30 40 50 60 70 80 90 l O O i l O 120 130 140 150 160 170 180

No. of days after emergence

Fig. 2. Survivorship and fertility schedules ofA. miliaris under laboratory conditions. Solid and dotted lines refer to survivorship curves for females and males, respectively, and the histogram shows the number of eggs laid per female per day. Two horizontal bars depict the average longevity for both sexes, and R and S the average length of the pre- and post- reproductive period.

29

' 'c~ z 100 ~ -25 0 / . 80. :, A. sanctaecrucis .2o o.9,, ~) ,

-I-- * r

> -4o. .io ~. "~ o..

20- -5 -I0

, , l i / o , < 06 lb ab 30 4b 5b 6b 70 80 961601101kot io l , ;0~gold01i0180

No. of days after emergence

Fig. 3. Survivorship and fertility schedules of A. sanctaecruds under laboratory conditions. Explanations as in Fig. 2.

longevity, pre- and post-reproduct ive periods, fertility, intrinsic rate of natural in-

crease (r) and mean length of a generat ion (7).

Longevity, pre- and post-reproductive periods. The two species had almost the same

mean longevity with no significant difference between males (88.4 days in A M and

63.8 in AS) and females (87.9 days in A M and 83.3 in AS) (Table 3). The m a x i m u m

longevity of both species spanned 5-6 months. The mean length of the pre-reproduc-

tive period (27.2 days in A M and 33.8 in AS) was much longer than that of the post-

reproductive period (10.9 days in A M and 14.3 in AS). Although the durat ion of the

pre-reproduct ive period of the females varied considerably, i.e. 17-42 days in A M and

21-49 in AS, KENT)ALL'S rank test revealed that the female 's age at first oviposition was

not significantly correlated with either age at final oviposition or with age at death in

either species; the females with shorter pre-reproduct ive period nei ther completed their

Table 3. Survivorship and fertility schedules ofA. miliaris and A. sanctaecrucis under labOratory con- ditions.

Species A. miliaris A. sanctaecrucis

No. of replications Longevity (in days)

Pre-reproductive period (in days) Post-reproductive period (in days) Fertility

(total no. of eggs laid per female) Intrinsic rate of natural increase

(per capita per day), r Mean length of a generation

(in days), T

9 9

13 9 88.4-- + 23.5 ( 33-162)* 63.8-+31.6 (23-144) 87.9+ 17.1(43-137) 83.3--+30.4 (45-171) 27.2 + 4.3 ( 17- 42) 33.8-+ 6.5 (21- 49) 10.9 + 4.7( 4- 31) 14.3-+10.1( 2- 36)

442.9+171.7 (103-981) 80.1-+54.8 (15-201)

0.070 0.044

76.7 83.9

* Average_+ 95% confidence limits (range)

30

oviposition earlier nor died earlier than those with the longer pre-reproductive period.

Mating. In the present study, the frequency and duration of matings were not

known exactly, because the mating records were kept only in the course of daily food

exchange which took just a few minutes for each cup. The two species were observed

in copula repeatedly at various intervals. The maximum number of observed copula-

tions per pair was eight (AM) and four (AS).

The intrinsic rate of natural increase r. The r was determined by solving the equation

Z e-r%mt = 1 for r, where t is age in days (BIRCH, 1948). We need the life table of both

the immature and adult stages for the calculation of r, and in practice, we assumed that

(1) no death occurred in the immature stages because the eggs, larvae and pupae rarely

died unless we mishandled them and (2) the length of immature stage was 36.5 day~ in

AM and 33.5 in AS (i.e., the averages of total immature periods in Table 1). The r

value thus derived was 0.070 (AM) and 0.044 (AS).

The net reproductive rate Ro and mean length of generation T. The Ro (= Y. lxmx) is the

average number of female eggs produced per female adult. The mean length of

generation T was derived from T = log Ro The values of Ro and T thus derived were f

221.5 and 76.7 days in AM, and 40.0 and 83.9 days in AS (Table 3). The total

number of eggs produced per female was 442.9 in AM and 80.1 in AS, which is twice

the value of Ro. The total number of eggs produced per female was positively cor-

related with the longevity of the female; KEm)ALL'S coefficient of rank correlation (r)

14- 12~ 10-

X 8-

E 6- X 4-

2" 0

80-

IM ,I, R - - ~

. . . . . . . . . . .

6oJ x O

I ~ 40 -

20 -

o o lb 2b 3b 4b 5b 7b 8b 9b l&l;ot ol ol;,odol ol o ol o2 o2 o

Age (doys), X Fig. 4. Reproductive function (lxmx, top) and reproductive value (Vx/Vo, bottom) plotted

against age for A. miliaris, IM: duration of the immature stages. R: mean duration of the pre-reproductive period. Vertical arrow shows the extinction of the females.

31

4.

3 .

x 2-~ E X

0 50 -

25"

x" 0 20. > I~; 1 5 -

10-

I M '1( R '1

A. $onC/OOCruC/S

A ,A M A i i �9 ! w i i i , i , i ,

O , , . i

0 10 20 30 40 50 60 70 80 90 1 0 0 1 1 0 1 2 0 1 3 0 1 4 0 1 5 0 1 6 0 1 7 0 1 8 0 1 9 0 2 0 0 2 1 0

Fig. 5. against age for A. Sanctaecrucis.

Age (days), X

Reproductive function (lxrnx, top) and reproductive value (Vx/Vo, bottom) plotted Explanations as in Fig. 4.

was 0.57 in AM and 0.67 in AS (both p<0 .01) .

Age-specific fertility and reproductive value. FISHER'S (1930) reproductive value is

given by

Vx _ g* ~, e_rtl, mtAt Vo l . , = .

where t is an interval for measuring lx and mx (At = 1 in this case). The reproductive

value of temperate species usually increases until the age of first reproduction and then

rapidly decreases with age. However, the reproductive value of the two species chang-

ed with age in a different manner, as shown below. In AM the reproductive value had

a smooth peak on day 69.5 (i.e., 33 days and 5.8 days after the start of the female's

adult life and of her reproductive period, respectively), and then the value dropped

gradually, but remained at a relatively constant level until near the end of the female's

life span (Fig. 4). The reproductive value of AS (Fig. 5) had a plateau with some

peaks; the highest peak value was attained on day 84.5 of oviposition (it fell on day 51

and day 17.2 of the adult life and the reproductive period, respectively). The final

oviposition occurred on day 127 (AM) and day 135 (AS) of the female's adult life.

Thus the two species showed a high reproductive value for most of their life span, as a

result of prolonged survivorship and fertility periods (Figs. 2 and 3).

DISCUSSION

Extreme longevity of adult tortoise beetles under laboratory conditions has been

32

reported from other tropical areas: e.g., Aspidomorpha adhaerens in New Guinea can live

as long as six to seven months, with a maximum of 265 days (SIMON TI-IOMAS, 1964).

Average longevity of Gratiana spadicea in Brazil was 197.8 days for males and 83.8 days

for females (ALBUQUERQUE and BECKER, 1986).

Fluctuations in adult numbers and egg mortalities of the AM and AS were studied

under field conditions in the garden of the Sumatra Nature Study Laboratory for 27

consecutive months (1982-1985). The results indicate that (1) the two species were ac-

tive throughout the year, (2) both species suffered high egg mortality from parasitism

by wasps, (3) the extreme longevity of adults AM and AS was confirmed by mark-

recapture and elytral color change, as suggested by this article and (4) they had exten-

sive dispersal power (NAKAMURA and ABBAS , in press, and in preparation).

The host plant, L carnea grows prolifically as a weed everywhere in Padang.

Local people repeatedly clear the plants every 2-3 months because of their vigorotls

asexual propagation (NAKAMURA and ABBAS, 1987b). Adults of AM and AS which

tend to fly actively and have prolonged survivorship and reproductive schedules. Con-

sequently, they can easily colonize a newly formed or recovered food plant patch,

where they are less likely to sustain wasp parasitism than in the older patches. The

two species are obviously well adapted for living in tropical environments like Padang,

where rainfall is ample but unpredictable (INouE and NAKAMURA, in press), and food

resources are available throughout the year but rather patchily distributed.

In Padang, the phytophagous ladybird, Epilachna septima, which depends on bitter

cucumber, Momordica charantia (ABBAS and NAKAMURA, 1985) and E. vigintioctopunctata,

which survives on solanaceous crops (NAKAMUgA and ABBAS, 1988) are also exploiters

of patchily distributed food plants, and their demographic traits are similar to those of

the two tortoise beetles in this article.

SUMMARY

Two species of tortoise beetles, Aspidomorpha miliaris (AM) and A. sanctaecrucis (AS)

feeding on a shrub-like morning glory, Ipomoea carnea, were reared under laboratory

conditions to study their survivorship and fertility schedules. AM and AS required

34-39 days and 30-37, respectively, for the development of the immature stages. The

mean longevity of the males was 88.4 days in AM and 63.8 in AS, and that of females

was 87.9 days in AM and 83.3 in AS. The mean length of the pre-reproductive period

(27.2 days in AM and 33.8 in AS) was much longer than that of the post-reproductive

period (10.9 days in AM and 14.3 in AS). Females laid eggs at a nearly constant rate

throughout their reproductive period. The reproductive value Vx/Vo of the two species

remained high for most of their adult life, as a result of prolonged survivorship and fer-

tility periods. The total number of eggs produced per female was 442.9 (AM) and

80.1 (AS). The intrinsic rate of natural increase r was 0.070 (AM) and 0.044 (AS) per

capita per day. The prolonged reproductive schedules, coupled with strong dispersal

33

p o w e r , o f t h e s e spec ies n o d o u b t h a v e an a d a p t i v e v a l u e for l i v i n g in h i g h l y d i s t u r b e d

t r o p i c a l e n v i r o n m e n t s , w h e r e r a in fa l l is a m p l e b u t u n p r e d i c t a b l e a n d f o o d r e s o u r c e s

a r e a v a i l a b l e t h r o u g h o u t t h e y e a r in a w i d e a r ea , b u t d i s t r i b u t e d in w i d e l y f l u n g pa t -

ches .

ACKNOWLEDGMENTS: We thank all the members of Sumatra Nature Study, in particular Prof. S.

KAWAMURA (Kyoto University), Prof. R. OHGUSHI (Kanazawa University) and Dr. A. BAKAR (Andalas

University) for their encouragement. We are indebted to Dr. S. KIMOTO (Kurume Medical University)

for identification of the tortoise beetles. Cordial thanks are also due to Mr. M. KAWAMOTO (Kanazawa

University) for drawing the figures in this article.

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q ~ Z . �9 I d r u s A b b a s �9 Ahso l H a s y i m

y ; ~ - ~ . . z . . ; t Aspidomorpha miliaris (AM) ~ ~ = ~ -e y ~ ~ y ;~~ ~ ~ A. sanctaecrucis (AS) ~,

(1)~~;~ Fg2]~J~L~ ~'t~-~-"~Ta H ~ , , 34--39 (AM) :~J~O ~ 30~37 (AS) "~',~.o ~c.

(2),b~:.....~�9 " ( H ~ ) 1~'~-, ~[~ 88.4 (AM), 63.8 (AS) ;~ J~ ~1[~ 87.9 (AM), 83.3 (AS) "(',~.o ~:20

(3)p~I~?)]Fu~ (AM "~" 27.3, AS ~" 33.8 13) ~ , p~[;1~15r ~] (AM "1~- 10.9, AS "~" 14.3 13) ~ 9

(5) 1 ~ 7 ~ 9 J ~ , 442.9 (AM), 80.1 (AS) "I?~--~7~o

(6)~l~J~ ,~ , ,~ J l~ r (IEI~7~c 9) ~ , 0.070 (AM) ~3~fY 0.044 (AS) ~ ' ~ 7 ~ a o

1L~'c.