Demographic traits of Hippodamia variegata (Goeze) (Coleoptera: Coccinellidae) reared on Aphis fabae...

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Demographic traits of Hippodamiavariegata (Goeze) (Coleoptera:Coccinellidae) reared on Aphis fabaeScopoli (Hemiptera: Aphididae)Shima Rahmani a & Ali R. Bandani aa Plant Protection Department , University College of Agricultureand Natural Resources, University of Tehran , Karaj , IranPublished online: 07 Mar 2013.

To cite this article: Shima Rahmani & Ali R. Bandani (2013) Demographic traits of Hippodamiavariegata (Goeze) (Coleoptera: Coccinellidae) reared on Aphis fabae Scopoli (Hemiptera:Aphididae), Archives Of Phytopathology And Plant Protection, 46:12, 1393-1402, DOI:10.1080/03235408.2013.768061

To link to this article: http://dx.doi.org/10.1080/03235408.2013.768061

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Demographic traits of Hippodamia variegata (Goeze) (Coleoptera:Coccinellidae) reared on Aphis fabae Scopoli (Hemiptera: Aphididae)

Shima Rahmani and Ali R. Bandani*

Plant Protection Department, University College of Agriculture and Natural Resources,University of Tehran, Karaj, Iran

(Received 1 January 2013; final version received 6 January 2013)

Lady beetles are the most important predatory species among arthropods; thus,studying their population parameters gives a clear picture about their life span. Thedemography of the lady beetle, Hippodamia variegata (Goeze) (Coleoptera: Cocci-nellidae), was studied under laboratory condition, on Aphis fabae at 27 °C based onthe age-stage, two-sex life table. The means and standard errors were estimated bythe bootstrap techniques. The results showed that the intrinsic rate of increase (r)and the finite rate of increase (λ) were 0.18 and 1.20 d�1, respectively. The netreproduction rate (R0) and the gross reproduction rate were 232.49 and 442.48 off-spring/individual, respectively. The mean generation time (T) was 29.03 days. Thefirst peak of reproductive value in females was on the 18th day, which coincideswith the total pre-oviposition period counted from birth. The life expectancy curve(exj) of a new hatched female adult was 60 days. The results demonstrated thatgrowth, survival and development of the predator can be described accurately by useof the age-stage, two-sex life table.

Keywords: variegated lady beetle; population parameters; age-stage, two-sex lifetable

Introduction

Among all beneficial insects and arthropods, lady beetles are the most well-known pre-dators all around the world (William 2002). The variegated beetle, Hippodamia (Ado-nia) variegata (Goeze), is originated from palaearctic area and now is widespread inmany countries (Wang et al. 1984; Pang 1993; Lotfalizadeh 2001; Franzmann 2002;Kontodimas and Stathas 2005). This coccinellid beetle is an effective predator of aphidsand is used as biocontrol agent against a wide range of aphid species in different cropssuch as wheat, cotton, tobacco, vegetables and orchards either in outdoor farms or ingreenhouses (Cabral et al. 2008). Due to the big size and high feeding capacity, thisgeneralist-variegated beetle is proposed as valuable biological control agents (Kontodi-mas and Stathas 2005) and is useful in insect pest management programmes. For aneffective management of pest population, it is necessary to know the population ecologyof the predatory species. In fact, the life table of a population gives the most compre-hensive description on the growth, survival and fecundity. Life table study of some

*Corresponding author. Email: abandani@ut.ac.ir

Archives of Phytopathology and Plant Protection, 2013Vol. 46, No. 12, 1393–1402, http://dx.doi.org/10.1080/03235408.2013.768061

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Coccinellid predators was a subject of some authors’ paper (e.g. Michaud 2000;Tsaganou et al. 2004; Katsarou et al. 2005; Kontodimas et al. 2008; Atlihan and Chi2008; Wu et al. 2010; Yu et al. 2013).

Because the survivorship, development and reproduction of a population aredescribed comprehensively in the cohort life table, thus it seems essential for both theo-retical and applied population ecology.

In evaluating the life table of a kind of organism obtained from field data, fitness ofa population can be influenced by various biotic and abiotic factors. However, field lifetables are often difficult to construct. Thus, life tables are constructed using laboratorydata under controlled conditions and are useful in revealing the maximal growth poten-tial of a population.

There are many ecology reviews and textbooks that discuss the theory and method-ology of population demography (Birch 1948; Price 1984; Cary 1993; Ricklefs andMiller 1999).

Traditionally, measuring the life tables, according to the Lewis–Leslie matrix andLotka–Euler’s formula, ignores the male population and the variable developmental ratesamong individuals. So due to taking only the age into consideration, the age-specific lifetable cannot describe the stage differentiation of insect population. In order to take bothsexes and the variable developmental rates into consideration, Chi and Liu (1985) andChi (1988) developed the age-stage, two-sex life table theory and data analysis.

In this study, because of usual variation in developmental rate among individualsand between sexes in a natural population, some of demographic traits of H. variegatain the laboratory based on the age-stage, two-sex life table were evaluated.

Materials and methods

Insect culture

The colony of H. variegata obtained from laboratory of insect ecology from departmentof plant protection, University of Tehran. This colony was reared on Aphis fabae andmaintained in a growth chamber set at 27 ± 1 °C, 70 ± 10% RH, and a photoperiod of16:8 (L:D) for several generations before being used for life table study. Aphids werereared on broad bean plants, Faba vulgaris, under 22 ± 1 °C and 70 ± 10% RH, and aphotoperiod of 16:8 (L:D). The broad bean plants were planted in pots filled with saw-dust and watered, and fertilised with HortiGrow (product of Hortiland, Holand) twiceper week.

Population demography

A cohort of about 92 eggs (6 hours old) was selected from the lady beetle laboratorycolony and placed into Petri dishes (90mm diameter) based on Yu et al. (2013). Eggswere kept in a growth chamber at 27 ± 1 °C, 70 ± 10% RH, and a photoperiod of 16:8(L:D). The eggs were checked every 6 h and newly emerged larvae were transferred tonew Petri dishes (60mm diam.). Petri dishes were kept in an incubator at 27 ± 1 °C, 70± 10% RH and a photoperiod of 16:8, and were supplied daily by all stages of A. fabaeas food sources. Mortality and development were checked every 24 h. After the emer-gence of adults, males and females were paired and checked daily in order to recordtheir survival and their laid eggs. The experiments continued until the death of all theindividuals.

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Data analyses

The raw data of H. variegata were analysed according to the theory of age-stage, two-sex life table (Chi and Liu 1985; Chi 1988). The following population parameters ofeach cohort were estimated:

1. Net reproductive rate (R0)

R0 ¼X1

x¼0

lxmx

2. Intrinsic rate of increase (r)

1 ¼X1

x¼0

e�rðxþ1Þlxmx

3. Mean generation time (T)

T ¼ lnRo

r

4. Gross reproductive rate (GRR)

GRR ¼Xn

x¼0

mx

5. Finite rate of increase (λ)

k ¼ er

The TWOSEX-MS computer programme was used (Chi 2012) for estimation ofparameters data. The means and standard errors of the population parameters were esti-mated by employing the bootstrap techniques (Efron and Tibshirani 1993).

Results

In this study, 64 eggs out of the 92 initially collected for the life table study, hatchedsuccessfully, and at last 58 emerged as adults (28 males and 30 females) (Table 1).Pre-adult mortality of H. variegata was 36.9%. The total developmental time for allpre-adult stages was 14.64 days. Male adults of H. variegata lived an average of64.89 days, which was slightly longer than that of the female adults survived 60.86 days(Table 1).

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The mean fecundity per female was 709.3 eggs and mated females began oviposi-tion on 3.53th day after emergence (pre-oviposition period (APOP)). The total pre-ovi-position period (TPOP) was 18.03 days. Copulation happened all around the adult life(Table 1).

The main demographic parameters including intrinsic rate of increase (r), net repro-ductive rate (R0), GRR, mean generation time (T) and the finite rate of increase (λ) areshown in Table 2. These statistics were 0.18 (day�1), 232.49 (offspring/individual),442.48 (offspring/individual), 29.03 (day) and 1.20 (day�1), respectively (Table 2).

The other biological parameters such as age-stage survival rate (sxj) (Figure 1),age-stage fecundity (fij) (Figure 2), the life expectancy curve (exj) (Figure 3) and thereproductive value (vxj) (Figure 4) have been given.

Discussion

Life table studies are an important tool in the evaluation of population ecology. In thisstudy, demography of H. variegata was measured in 27 °C reared on A. fabae. Thesurvival percentage was higher than that reported by El Hag and Zaitoon (1996) andLanzoni et al. (2004).

The present results for total pre-adult duration were 14.65 days. El Habi et al.(2000) found that the total developmental time was from 7 to 27.58 days and tempera-ture-dependent. Atlihan and Chi (2008) also showed that life span of coccinellidschanges by different temperatures. So, shorter immature developmental time at highertemperature might be due to the increasing metabolism in Scymnus subvillosus. In theirstudy, the probability that a newly laid egg will survive to the adult stage was similar

Table 1. Life statistics (mean ± SE) of H. variegata at 27 °C and 70% R.H.

Parameters Stage n Mean SEM

Developmental time (day) Egg 92 2.07 0.03First instar 64 2.23 0.05Second instar 64 1.84 0.07Third instar 64 2.18 0.08Fourth instar 60 3.16 0.07Pupa 58 3.17 0.05

Adult longevity (day) Male 28 64.89 3.88Female 30 60.86 3.00

APOP of female (days) Female 30 3.53 0.15TPOP of female (days) Female 30 18.03 0.18Fecundity (eggs/female) Female 30 709.3 59.69

Table 2. The population parameters (mean ± SE) of H. variegata at 27 °C and 70% R.H.

Population parameter Mean SEM

Intrinsic rate of increase (r) (day�1) 0.18 0.007Net reproductive rate (R0) (offspring/individual) 232.49 39.44GRR (offspring/individual) 442.48 68.63Mean generation time (T) (day) 29.03 0.49Finite rate of increase (λ) (day�1) 1.20 0.008

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for 20, 25 and 30 °C but decreased in 35 °C. Also, the pre-adult mortality at 20, 25 and30 °C (41.3, 37.5 and 43.8%, respectively) was lower than 35 °C (78.0%) and lifeexpectancy decreased by increasing the temperature. Mortality in the present study at27 °C was 36.95%. It was comparable with Atlihan and Chi (2008).

Figure 1. The age-stage survival rate (sij) of H. variegata in pre-adult (A) and adult (B) stages.

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The longevity of both sexes in the obtained results was different but El Hag andZaitoon (1996) did not find any significant difference between the longevity of malesand females of H. variegata fed on Brevicoryne brassicae and Rhopalosiphum padi.

TPOP in demography is very important because it can reflect the effect of the firstreproduction on the population parameters (Gabre et al. 2005). In the current study,TPOP was 18.03 and similar to the results of Atlihan and Chi (2008) in 30 °C.

According to Chi and Liu (1985), the population parameters should be analysedbased on the data of the entire cohort, (both sexes and the different developmentalstages of individuals). As proven by Chi (1988), based on the two-sex life table, therelationship between R0 and mean female fecundity (F) is given as follows:

R0 ¼ F � N f=N

where N is the total number of individuals included at the beginning of the life tablestudy and Nf is the number of female adults that resulted from N.

Use of the jackknife and bootstrap techniques to population parameters is discussedby Meyer et al. (1986). However, Huang and Chi (2012) mathematically invalidated theuse of the jackknife technique in estimating the net reproductive rate. In fact, when thenet reproductive rate was estimated by the bootstrap technique, its frequency distribu-tion met the assumptions of normality and this was important for further statistical anal-ysis. Moreover, by using the jackknife technique, zero-pseudo values were obtained forR0, and this means that the population cannot reproduce offspring and as a result, nointrinsic rate can be estimated.

Figure 2. The female age-stage fecundity (fi7) of H. variegata.

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Figure 3. The age-stage life expectancy (exj) of H. variegata in pre-adult (A) and adult (B)stages.

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The intrinsic rate of increase in H. variegata (0.18 day�1) in this study was morethan the ladybird fed on Aphis gossypii Glover reared on five host plants (Wu et al.2010).

Age-stage survival rate (sxj) gives the probability that a newly hatched larva willsurvive to age x and stage j. In addition to survival rate, this parameter shows stagetransitions. As shown in Figure 1, because sxj takes into account variation in develop-mental rates among individuals, overlapping occurs among stages. In this study, adecrease has been shown in survival rate by spending time in both sexes. Although,females survive until the day 110 but males remain alive until the day130.

Age-stage fecundity (fij) is shown in Figure 2. Because only females produce off-spring, there is only a single curve fx7 (i.e. the female is the seventh life stage). In thisstudy, because the egg batches were laid at discrete time intervals, 18 peaks of femalefecundity (fi7) were represented in Figure 2 where the maximal mean daily fecundity is25.8 eggs (the fourth pick in the day 38) (Figure 2). In this study, the mean of egg lay-ing in every day was 9.2 individuals.

The life expectancy curve (exj) calculated according to Chi and Su (2006) shows thetotal time that an individual of age x and stage j is expected to live (Figure 3). Theyounger individuals have more life expectancy (Figure 3(A)).When a female individualreached to the adult stage, life expectancy increased to 60 days (Figure 3(B)).

By taking the variable developmental rate between sexes and among individuals intoconsideration, this method can be used to detect the differences in life expectancy notonly between male and female but also among stages. Under controlled laboratory con-ditions, obtained results showed that the life expectancy decreased gradually with age-ing (This trend has been shown by other authors e.g. Abou Zied et al. 2003; Atlihanand Chi 2008). However after the day 95, there was a peak in both sexes due to lowmortality after this time (Figure 3).

The reproductive value (vxj) is the expectation of future offspring of individuals ofage x and stage j (Fisher 1930; Pianka 1994) (Figure 4). This parameter is meaningful

Figure 4. The age-stage reproductive value (vxj) of H. variegata.

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only for females because contribution of males in future population is not described byFisher (1930) and so the curve is not explained for males. If the pre-oviposition periodis counted as time from birth to first reproduction in females (TPOP) (Amir-Maafi andChi 2006), the mean TPOP for females was 18.03 days; it was similar to their first peakof reproduction which was at day 18. There are three main reproduction peaks in whichthe highest is on the day 35 (105 eggs) (in younger female adults). In the days 47 and90, egg production is reduced and this reduction in the day 90 is near to 0. However,females continue to reproduction and laying the eggs during their life.

Present results showed that studying life tables especially when both sexes anddevelopmental stages are included provides a comprehensive description of the survival,development and reproduction of a cohort of individuals.

AcknowledgementsThis work was funded by a grant from Biological Control Center of Excellence.

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