Effect of Temperature on Demography and Predation Rate of Menochilus sexmaculatus (Coleoptera: Coccinellidae) Reared on Phenacoccus solenopsis (Hemiptera: Pseudococcidae).
Zigzag beetle, Menochilus sexmaculatus Fabricius (Coleoptera: Coccinellidae) is an important predator of cotton mealy bug, Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae). Understanding the effect of temperature variations on its demography and predation rate is necessary to predict the population dynamics of this beetle against cotton mealy bug. Age-stage, two sex life tables of zigzag beetle were constructed at three different temperature regimes: 24 +- 0.5 AdegC and 27 +- 0.5 AdegC with 60-70% R.H. and ambient condition (32 +- 4 AdegC; 16-50% R.H.) with 14:10 h (L:D) photoperiod using cotton mealy bug as a host.
According to the results, the immature duration and adult longevity were comparatively longer at lower temperature (24 +- 0.5 AdegC) and shorter at higher temperatures. Among population dynamic parameters, net reproductive rate (R0) was 216.52, 105.99 and 27.07 off-springs per individual, intrinsic rate of increase (r) was 0.1543, 0.1600 and 0.1518 off-springs per female per day at 27 +- 0.5, 24 +- 0.5, and 32 +- 4.0 AdegC, respectively. Survival rate (sxj) and age-stage specific fecundity (fx7) were greater at 24 +- 0.5 AdegC. Among immature stages, 4th instar was the most voracious with highest predation rate. Adult females consumed more cotton mealy bug nymphs at 24 +- 0.5 AdegC. Net predation rate (C0) of the beetle was 5548, 4463.2 and 2016.90 at 24 +- 0.5 AdegC, 27 +- 0.5 AdegC and 32 +- 4 AdegC, respectively.
The values of transformational rate (Qp) exhibited that 25.62, 42.11 and 74.50 nymphs of P. solenopsis were required per female beetle to lay one egg at 24 +- 0.5 AdegC, 27 +- 0.5 AdegC and 32+-4AdegC, respectively. Finite predation rate (I) was more (7.64) at 24+-0.5AdegC followed by 27+-0.5AdegC (6.56) and the lowest at 32 +- 4 AdegC (0.66). Adult beetles proved strong natural enemies of P. solenopsis. Our study provides detailed basic information for successful rearing of M. sexmaculatus in the laboratory and use as a bio-control agent against cotton mealy bug in the field at different temperature conditions.
Age-stage two sex, Life table, Menochilus sexmaculatus, Phenacoccus solenopsis, Predation rate.
Cotton mealy bug, Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae) caused losses of 3.1 million cotton bales during 2006-07 in the Punjab, Pakistan (Mahmood et al., 2011). It is a polyphagous insect pest which infests almost 194 plant species identified as crops, fruits, vegetables and ornamental plants (Vennila et al., 2011; Hameed et al., 2012). Ladybird beetles are known to be effective bio-control agents against mealy bugs (Ali and Rizvi, 2009; Michaud, 2001). A large number of predatory coccinellids have been identified as effective bio-control agents of cotton mealy bug (Michaud, 2001; Rafi et al., 2005; Ali and Rizvi, 2009).
Among these, zigzag beetle Menochilus sexmaculatus Fabricius (Coleoptera: Coccinellidae) has been suggested to be the promising bio-control agent against cotton mealy bug (Arif et al., 2012), as well as other soft bodied insects including aphids, plant hoppers, thrips, jassids, scale insects and white flies (Rahman et al., 1993; Solangi et al., 2007). This beetle is widely distributed in South Western Asia, Indonesia, Philippines, South Africa, India and Pakistan.
Being poikilothermic organisms, the development, reproduction and predatory potential of insects is affected to a great extent by temperature variations in the environment. Insects perform best only at a certain temperature range (Roy et al., 2002; Rana, 2006; Pakyari and Enkegaard, 2012). Life table studies of a predator at varying temperatures can provide detailed information about survivorship, development, mortality and life expectancy (Ali and Rizvi, 2007, 2008). Most of the previous studies on M. sexmaculatus were focused on exploring its predatory potential against different aphid species (Mari et al., 2004; Ali and Rizvi, 2009; Saleem et al., 2014) and cotton mealy bug (Arif et al., 2011, 2012; Ali et al., 2013).
An insect with better population growth rate in a particular condition does not necessarily mean an efficient predator (Yu et al., 2013).Therefore, predation rate must be considered along with population growth potential in life table under different temperature regimes to assess the efficacy of a particular predator (Chi et al., 2011).
The present study was designed to determine the effect of temperature on demographic parameters and predation rate of M. sexmaculatus against P. solenopsis. This study was based on the hypotheses that (a) temperature influences parameters of demography and predation rate of M. sexmaculatus (b) development and predation rates among individuals and between sexes determine the variations of predatory potential of M. sexmaculatus, and (c) finite predation rate of (both male and female adult) beetles also influence the predatory potential at a particular temperature.
MATERIALS AND METHODS
Demographic studies of zigzag beetle on cotton mealy bug were conducted at three sets of temperature; 24 +- 0.5 AdegC, 27 +- 0.5 AdegC and ambient temperature condition (32 +- 4 AdegC). In first two temperatures, R. H. was kept at 60-70% and for ambient laboratory condition, the temperature ranged from 28-36AdegC with 16-50% R. H. First two studies were conducted in growth chamber and the third was conducted in the laboratory at ambient conditions during the months of September and October, 2013. This is actually time of the year when cotton mealy bug population is at its peak in the cotton crop. The ambient temperature condition was used to compare development, reproduction and predation of beetle with those in controlled temperatures to get information for successful mass rearing of M. sexmaculatus.
Field collection and rearing
Cotton mealy bug was collected from Hibiscus rosa-chinensis Linnaeus (Malvales: Malvaceae) plants and reared on pumpkin Cucurbita pepo Linnaeus (Cucurbitales: Cucurbitaceae) fruits in enclosed containers to develop laboratory culture. Adult zigzag beetles were collected from Parthenium hysterophorus Linnaeus (Asterales: Asteraceae) plants near University field area and were kept in petri dishes containing cotton mealy bugs and 10% honey solution as a food. Before using for life table studies, twenty pairs of M. sexmaculatus were reared on cotton mealy bugs at each set of temperature i.e., 24 AdegC, 27 AdegC and ambient condition (32 +- 4AdegC) for two generations. To maintain genetic variability of predatory zigzag beetles, more number of adult beetles were collected from field and added in the stock culture.
Life table study of Menochilus sexmaculatus
One hundred eggs of M. sexmaculatus were shifted to 100 petri dishes each for the three temperature treatments. Moulting and survival of each larval stage from 1st to 4th instar were carefully recorded daily. The adults were paired on the basis of their body size (female larger than male) and transferred to petri-dishes in order to observe their mating behaviour (Mari et al., 2004). Survival, adult longevity and fecundity of each female beetle were recorded daily until their death.
We constructed age-stage, two sex life table to estimate life table parameters of the beetles and population dynamics parameters (r, the intrinsic rate of increase; I>> finite rate of increase; R0 the net reproductive rate, T mean generation time) using TWOSEX-MSChart (Chi, 1988, 2012a; Chi and Liu, 1985). Differences in the development time, longevity and reproduction among M. sexmaculatus at three different temperatures were analyzed using one way ANOVA followed by multiple comparison with Tukey-Kramer test (P 28AdegC) temperature on the performance of M. sexmaculatus.
Age-stage specific survival rate (sxj) of M. sexmaculatus (the probability that freshly laid eggs will survive to age x and develop to the stage j) was plotted in Figure 1, which showed overlapping curves, illustrating differences in developmental rates in both immature and mature stages. The first adults emerged at the age of 22, 16 and 12 days and the survival rates were 29.08, 50.02 and 17.64 at 24 +- 0.5 AdegC, 27 +- 0.5 AdegC and 32 +- 4 AdegC, respectively (Fig. 1).
The age-specific survival rate (lx) is the probability of newly hatched egg to survive at age x. It ignores the individual developmental rate and stage discrepancy. Age-stage specific fecundity (fxj) explains mean number of eggs laid per adult female at age x and stage j per day (adult female of M. sexmaculatus was at 7th life stage) and only counts female individuals which were able to produce eggs. Age-specific fecundity (mx) describes the fecundity of emerged larvae to adult age x; this is why age-stage specific fecundity (fx7) curves showed higher peaks than the age-specific fecundity (mx) curves (Fig. 2). First egg laying occurred on 26th, 20th and 15th day with (fx7) value of 0.1944, 1.556 and 1.000 eggs per female per day at 24 +- 0.5 AdegC, 27 +- 0.5 AdegC and 32 +- 4 AdegC, respectively.
Age-specific fecundity (mx) was 0.132, 0.304 and 0.333 females per female per day at age of 26th day, 20th day and 15th day with age specific survival (lx) was 0.53, 0.46 and 0.24 days at 24 +- 0.5 AdegC, 27 +- 0.5 AdegC and 32 +- 4 AdegC, respectively. The peak of age-stage specific fecundity (fx7) was 71.52, 69.58 and 38.84 eggs per female per day at the age of 33rd, 29th and 20th days, at 24 +- 0.5 AdegC, 27 +- 0.5 AdegC and 32 +- 4AdegC, respectively. Higher peaks of mx, lxmx and fx7 were observed at 24 +- 0.5 AdegC and the lowest at 32 +- 4 AdegC.
Traditional female age-specific life tables (Birch, 1948) do not provide information regarding stage differentiation, role of male individuals in the population, variable survival rate of different stages and their overlaps in the life history of insects (Chi ,1988; Yu et al., 2005; Chi and Su, 2006; Huang and Chi, 2012). We used age-stage, two sex life tables because it accounts variable developmental rate among individuals and stage differentiation of both sexes while calculating population parameters.
Population dynamics parameters
Population dynamic parameters, estimated by using Bootstrap method, reflected strong effect of temperature (Table II) (Efron and Tibshiramin, 1993). Intrinsic rate of increase (r) increased with increasing temperature; from 0.1543, 0.1600 and 0.1518 females per female per day at 24 +- 0.5 AdegC, 27 +- 0.5 AdegC but declined at 32 +- 4 AdegC. Due to faster development, higher daily production of eggs and earlier peaks, reproduction was higher at 27 +- 0.5 AdegC than at 24 +- 0.5 AdegC. Finite rate of increase (I>>) followed the same trend as exhibited by intrinsic rate of increase (r). The highest net reproductive rate (R0) of 216 offspring per individual was noted at 24 +- 0.5 AdegC and the lowest (27.07 offspring per individual) at 32 +- 4 AdegC. Mean generation time (T) was the longest (34.86 days) at 24 +- 0.5 AdegC and the shortest (21.73 days) at 32 +- 4AdegC. Life table studies of Harmonia dimidiata on Aphis gossypii also showed decrease in mean generation time (T) and net reproductive rate (R0) with increasing temperature Yu et al (2013).
Insects can tolerate temperature changes only up to certain limits and beyond those limits their life activities are negatively affected (Hameed et al., 2012). Same phenomena were observed in the current studies; faster rate of development, higher daily egg production and earlier peaks in reproduction at 27 +- 0.5 AdegC than at 24 +- 0.5 AdegC. However, variability in temperature as for ambient condition showed faster development but decreased survival rate and fecundity.
Predation rate of Menochilus sexmaculatus
Predation rate of M. sexmaculatus larvae increased rapidly from 1st to 4th instars. Predation increased with the progress of the larval instar (Unal et al., 2017) Total number of mealy bugs fed during immature stages was 1730, 1430.61 and 614 per larva at 24 +- 0.5 AdegC, 27 +- 0.5 AdegC and 32 +- 4 AdegC, respectively (Table III). Predation rate of both female and male adults was higher as compared to different larval instars. During whole life span, female consumed significantly more mealy bugs than those of males at all temperature regimes. M. sexmaculatus adult females consumed maximum cotton mealy bugs (5199) at 24 +- 0.5 AdegC, (3889.7) at 27 +- 0.5 AdegC and minimum (1905.83) at 32 +- 04 AdegC. Predation rate of M. sexmaculatus increased with age; the fourth instar to be the most voracious among all larval instars. Similar findings were reported by Saleem et al. (2014) while studying the predation efficacy of M. sexmaculatus against Macrosiphum rosae under laboratory conditions.
They found 4th instar consuming more preys/day of Rhopalosiphum maidis, Aphis gossypii and Therioaphis trifolii than their earlier instars. Female M. sexmaculatus with long life period showed more predatory potential than males against R. padi at varying temperature conditions Ali et al. (2012). In our study, females almost consumed more than twice the number of preys than by those by males. This voracity of female may be due to their larger body size and longevity than males, which also require more energy to meet reproduction needs (Farhadi et al., 2011). At ambient conditions, the overall period of all stages was shorter but survival rate and fecundity were lower as compared to those in controlled conditions.
Table II.- Population dynamic parameters of M. sexmaculatus reared on P. solenopsis at three different temperatures estimated by all individuals and Bootstrap technique.
Parameters###24+-0.5 AdegC###27+-0.5 AdegC###Ambient condition (32+-4
Table III.- Comparison of predation rate of M. sexmaculatus on first nymphal instar of P. solenopsis at three different temperatures.
Life stages###24+-0.5 AdegC###27+-0.5 AdegC###Ambient condition (32+-4 AdegC)
Total pre-adult###1st to 4th Instar###1730+-10.50a###1430.61+-5.55b###614.0+-4.39c
Total life span###Female###6947.0+-190a###5322.5+-110.1b###2521.67+-48.43c
Net Predation rate (C)###5548.00+-436###4463.2+-329.34###2016.90+-171.25
Transformation rate (Qp)###25.62###42.11###74.500
Stable Predation rate (I")###6.550###6.447###0.5632
Finite Predation rate (I)###7.640###6.5631###0.6559
Age-specific net predation, (qx) is the mean number of cotton mealy bugs consumed by an average individual of M. sexmaculatus during its entire life span (Chi and Yang, 2003). Both age-specific predation rate (kx) and age-specific net predation rate (qx) showed two obvious curves at egg and pupal stage because at egg and pupal stages do not consume any prey (Fig. 3). These non-predatory phases could not be reflected with traditional life table. This is crucial to decide release intervals of natural enemies for effective biological control program (Yu et al., 2013). Net predation rate (C0) calculated by taking survival rates, predation rates and longevities of the zigzag beetle into consideration was the highest (5548 cotton mealy bugs) at 24 +- 0.5 AdegC followed by that (4463.2 cotton mealy bugs) at 27 +- 0.5 AdegC.
Transformation rate (Qp) provides a demographic estimation of the relationship between the reproductive rate and predation rate of the predator (Chi an Yang, 2003). Qp reflected that M. sexmaculatus required 25.62, 42.11 and 74.50 cotton mealy bugs (1st instar) for the production of one egg at 24 +- 0.5 AdegC, 27 +- 0.5 AdegC and 32 +- 4 AdegC, respectively. More preys would be required for the beetle at 24 +- 0.5 AdegC than at 27 +- 0.5 AdegC. Contribution of males towards predation of cotton mealy bug was different as compared to females (Table III) and stage differentiation of the beetle showed clear overlaps (Fig. 1). In contrast, traditional life tables theory do not consider sex variation and stage differentiation resulting in overestimation of results regarding predation capacity (Farhadi et al., 2011). In the current studies, we calculated both population dynamics parameters and predation rate of M. sexmaculatus against cotton mealy bug to avoid such over-estimation.
Application of studies in laboratory rearing and bio-control program
Growth potential and efficacy of the rearing program like development rate of larval stage, mortality rate, weight of fresh adults and abundance in the field and life table parameters have been assessed previously (Kalushkov, 1998; Atlehan and Kaydan, 2002; Soroushmehr et al., 2008). However, the efficacy of a predator cannot be determined precisely without considering its predation rate along with population growth rate. Although intrinsic rate of increase (r) and finite rate of increase (I>>) of the beetle were higher at 27 +- 0.5 AdegC, the net predation rate was high at 24 +- 0.5 AdegC. But higher intrinsic rate of increase (r) and finite rate of increase (I>>) do not indicate higher predatory potential of the predator. Therefore, finite predation rate (I) was calculated keeping into consideration both population growth rate and predation rate to compare the predatory potential of the beetle (Chi et al., 2011).
Because faster finite predation rate (I) was observed at 24 +- 0.5 AdegC than 27 +- 0.5 AdegC, we can consider M. sexmaculatus as the more efficient predator of cotton mealy bug at 24 +- 0.5 AdegC than higher temperature regimes.
Stage-specific predation rate and life table parameters were used to assess the predator-prey relationship of M. sexmaculatus with P. solenopsis. We have demonstrated that, predation rate must be considered with reproductive rate of the beetle during life table studies to explore accurate predation capacity. At 24 +- 0.5 AdegC, M. sexmaculatus is an efficient predator as compared to other temperature regimes. In order to get the next generation earlier, the rearing at 24 +- 0.5 AdegC will save the labour expenses and consume less prey as a food.
Statement of conflict of interest
Authors have declared no conflict of interest.
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|Author:||Iftikhar, Ayesha; Aziz, Muhammad Asif; Naeem, Muhammad; Ahmad, Munir; Mukhtar, Tariq|
|Publication:||Pakistan Journal of Zoology|
|Date:||Oct 31, 2018|
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