Prey-Dependent Life Attributes of an Aphidophagous Ladybird Beetle, Propylea dissecta (Coleoptera: Coccinellidae)

Predation potential, development, immature survival and reproduction of an aphidophagous ladybeetle, Propylea dissecta (Mulsant) was studied when fed on seven aphid prey, viz. Aphis gossypii, Aphis craccivora, Lipaphis erysimi, Uroleucon compositae, Brevicoryne brassicae, Rhopalosiphum maidis and Myzus persicae. A. gossypii was most suitable and consumed by the larvae and adults of P. dissecta, while M. persicae, the least. Pre-imaginal development of P. dissecta was fastest (0.080 day^-1) when A. gossypii was used as prey, whilst slowest (0.061 day^-1) on M. persicae. The immature survival, adult emergence, adult male and female longevity of P. dissecta was maximal (i.e., 77.10±0.04 and 93.21±0.79%, 57.10±1.62 and 62.40±1.93 days, respectively) on A. gossypii and minimal (i.e., 63.01±1.87 and 81.73±1.79%, 42.50±1.21 and 49.40±2.32 days, respectively) when M. persicae was provided as prey. Oviposition period, fecundity, percent egg viability and mean reproductive rate was maximum (i.e., 50.30±2.03 days, 856.00±30.00 eggs, 96.40±0.31% and 17.02 eggs per day) on A. gossypii, and minimum (i.e., 18.00±1.40 days, 212.00±18.21 eggs, 72.46±2.81% and 11.78 eggs per day) on M. persicae. Adult weight and developmental rate of P. dissecta have a positive correlation, which suggests that if immature stages of ladybeetle developed faster, they should grow into heavier adults. Female longevity and fecundity also have a positive correlation. The findings also reveal that all seven aphid species tested are essential food. Rank order of prey species was consistent in all experimental parameters.     

Temperature-dependent life attributes of an aphidophagous ladybird, Propylea dissecta

Reproductive attributes, age-specific fecundity and natality based life-tables of an aphidophagous ladybird, Propylea dissecta (Mulsant) were investigated at five temperatures, using aphid, Aphis gossypii as prey. Pre-oviposition period decreased significantly with increase in temperature up to 27°C and thereafter increased at a slower rate. Egg viability dropped significantly at temperature extremes. Females lived longer than males; however, longevity decreased significantly with increase in temperature. The predicted total fecundity and oviposition rate were 952.54 eggs and 35.15 eggs per day at 27°C predicted by a mathematical model compared to empirical estimates of 856.00±30.00 eggs and 39.7±2.26 eggs per day. Age-specific fecundity was triangular and temperature dependent. The peak in oviposition rate occurred earlier at higher temperatures. The highest values of net reproductive rate (R_o=431.1), intrinsic rate of increase (r_m=0.2134 day^-1) and finite rate of increase (λ=1.2379 day^-1) were recorded at 27°C, which suggests it is the optimum temperature for the mass rearing of P. dissecta.     

Mating behavior of an aphidophagous ladybird beetle,Propylea dissecta （Mulsant）

Abstract The mating behavior of Propylea dissecta (Mulsant) was investigated in detail. Protandry was evident, as males mature earlier than females. Predicted age of sexual maturation of male and female ladybirds was 7.12 and 9.33 days, respectively. Males performed courtship in six steps, viz. approach, watch, examine, embrace, mount and attempt. Embrace is possibly an appeasement act, while examine serves to recognize mate. Chemical signals initiate male attraction toward females. Visual and tactile cues appear to secondarily help in mate recognition. Mating lasted longest (275.40±12.23 min) when it occurred between unmated individuals. It was much shorter (176.60±5.60 min) when prior mated individuals copulated. There was a significant decrease in mating durations when adults were subjected to five successive matings. Fecundity and percent egg viability increased significantly with increase in the number of matings.     

Temperature-dependent development and immature survival of an aphidophagous ladybeetle, Propylea dissecta (Mulsant)

Development and survival of the immature stages of an aphidophagous ladybeetle, Propylea dissecta (Mulsant) was investigated at five constant temperatures, viz. 20, 25, 27, 30 and 35°C, using Aphis gossypii Glover as prey. Developmental period of all the life stages were significantly affected with change in constant temperature and developmental rate increased with increase in temperature. Theoretical lower thermal threshold for complete development and thermal constant was 10.39°C and 465.11 Day‐degrees, respectively. Of the various life stages, first instar larvae were most susceptible to mortality at temperatures between 20 and 30°C, whilst pre‐pupae suffered least mortality. Egg‐mortality was maximum at 35°C. Female biased sex ratios were obtained at all five temperatures tested with higher proportion of females at the extremes of temperature, thus suggesting that females are more thermal‐tolerant. Lowest mortality of immature stages with maximum larval survival and adult emergence was recorded at 27°C, while reverse was the case at 35°C. Thus, 27°C may be considered best for the laboratory rearing of P. dissecta.     

Effect of photoperiod on slow and fast developing individuals in aphidophagous ladybirds,Menochilus sexmaculatus and Propylea dissecta (Coleoptera: Coccinellidae)

The effects of environmental parameters on insect development have been studied extensively. But the reasons for 2 differential developmental rates within same cohort under varying environmental factors have not been explored. For the purpose, in this study the existence and stability of slow and fast development under 5 photoperiods (i.e., 8L: 16D, 10L : 14D, 12L : 12D, 14L : 10D and 16L : 8D; light and dark hours per day) and its effect on body mass and reproductive attributes in 2 aphidophagous ladybirds, Menochilus sexmaculatus (Fabricius) and Propylea dissecta (Mulsant) was examined on Aphis craccivora Koch at 27 ± 1 °C temperature. A clear bimodal (2 peaks, where the first peak represented the fast developing and the 2nd peak slow developing individuals) pattern of distribution at each photoperiod was found. The proportion of slow and fast developing individuals in a cohort differed with photoperiods. The slow developing individuals were more in numbers at 8L : 16D, in equal numbers at 14L : 10D and in less numbers at 16L: 8D, indicating that the variation in emergence was owing to exogenous cues influenced differential rates of mortality. Slow developing individuals had female biased sex ratio, higher longevity and lower body mass than fast developing individuals. Fast developing females laid higher numbers of eggs with higher egg viability than slow developing females. Study of such variations in development at different photoperiods is helpful to understand its role in the development of insects particularly ladybirds and permits the selection of fast developing bioagents for their use in biocontrol of pest species.     

Role of surface chemicals in egg cannibalism and intraguild predation by neonates of two aphidophagous ladybirds, Propylea dissecta and Coccinella transversalis

Abstract:  The role of surface chemicals in egg cannibalism and intraguild predation by neonates of two aphidophagous ladybirds, Propylea dissecta and Coccinella transversalis were examined. Neonates of both species prefer to eat non-sibling conspecific eggs than heterospecific eggs, with higher preference in P. dissecta . Surface chemicals appear to play a major role for the preference of conspecific eggs, as the ladybird behaviour was reversed when these chemicals were interchanged. The surface chemicals present on the eggs possibly act as feeding stimulants to conspecific neonates but not for heterospecific neonates. Egg clustering enhances the effect of surface chemical and stimulate non-sibling egg cannibalism and appears to reduce intraguild predation. Egg clustering is advantageous to ladybirds, as it can stimulate non-sibling cannibalism by neonates. It appears that aggregation of chemicals present on the conspecific egg surface possibly attracts the hungry conspecific neonate, providing the first meal for its survival, while protecting against heterospecific predators. Neonates of both ladybirds were reluctant to eat heterospecific eggs.     

龟纹瓢虫的行为学观察

通过以洋槐蚜Aphis robiniae Macchiati为食物对龟纹瓢虫Propylea japonica(Thunberg)进行室内饲养和观察,记述了龟纹瓢虫幼虫的孵化、蜕皮行为,成虫的羽化、捕食、交配、产卵行为,并对幼虫成虫的自残行为进行了初步观察。结果表明幼虫孵化、蜕皮,成虫羽化后均有取食自己蜕皮时产生的残皮的行为;龟纹瓢虫成虫捕食行为分为搜寻、捕捉、清洁、静止排泄4个步骤;交配一般由雄虫发起,雌虫背负成虫进行交配。龟纹瓢虫每产一粒卵平均耗时3.75秒,两粒卵之间产卵时间间隔约31.75秒,每次产卵量平均12枚;龟纹瓢虫存在自残行为,成虫在食物不足时会取食幼虫和虫卵,高龄幼虫在饥饿的状态下也取食低龄幼虫和卵,且同龄幼虫之间也会出现相互咬食的现象。     

Attack, escape and predation rates of larvae of two aphidophagous ladybirds during conspecific and heterospecific interactions

The attack, escape and predation rates for larvae of aphidophagous ladybird Propylea dissecta (Mulsant) and Coccinella transversalis Fabricius were quantified as a potential mechanism leading to the differences in the incidence of cannibalism and intraguild predation. These rates were compared at four larval instars within and between the species. The attack rates of larvae of C. transversalis were significantly higher than those of P. dissecta towards conspecific and heterospecific victims. For both species, third instars exhibited maximum tendency to attack. Escape rates in C. transversalis were higher than P. dissecta. In P. dissecta, the second instars made a greater number of escapes than other conspecific instars after being attacked by same stage cannibal or heterospecific predator. In P. dissecta, first instars suffered maximum mortality due to cannibalism and intraguild predation by conspecifics and heterospecifics of the same and older developmental stage. No larvae of C. transversalis were eaten by P. dissecta of the same stage. These results suggest that the larvae of P. dissecta were more often potential cannibals than intraguild predators, while the reverse was the case in C. transversalis. Based on this finding, it could be predicted that in patchy prey habitats, high rates of larval cannibalism in P. dissecta would occur with a high risk of cannibalism of first instars. Larvae of C. transversalis would respond as intraguild predators, while those of P. dissecta as intraguild prey. The greater size and walking activity of C. transversalis could be possible reason for this tendency.     

Why do neonates of aphidophagous ladybird beetles preferentially consume conspecific eggs in presence of aphids?

The neonates of aphidophagous ladybird beetles, Propylea dissecta, Coccinella transversalis and Coelophora saucia preferentially consumed conspecific eggs in presence of essential aphid prey, Aphis gossypii. This preference was strongest in P. dissecta and recorded in all density combinations of conspecific eggs and aphids. The neonates of all three species developed faster resulting in heavier second instars with lower mortality when fed on conspecific eggs as compared to mobile or defenceless aphids. In addition, they required less dry biomass of conspecific eggs than that of aphids. These results reveal that neonates are benefitted intrinsically when they feed on conspecific eggs. The neonates of P. dissecta exhibited discrimination for more nutritious food, as they contacted and ate conspecific eggs more frequently than aphids as their first meal. This discrimination was not found in Coccinella transversalis and Coelophora saucia possibly due to their bigger sizes and possible higher energy requirements. Although, the major effect of chemical cues was only noticed in P. dissecta, it could be concluded that chemicals (surface and within) of conspecific eggs possibly attract neonates for egg-cannibalism, as both eggs and aphids were consumed in greater numbers when coated with egg extracts. The reverse occurred when eggs and aphids were coated with aphid extracts.     

Locomotor Behavior of Flightless Harmonia axyridis Pallas (Col., Coccinellidae)

To improve the efficiency of the lady beetle H. axyridis as a biological control agent against aphids, a flightless population was obtained by feeding adults with a mutagen and selecting their progeny for nonflying but otherwise morphologically normal individuals. These flightless adults attempted to fly but immediately fell. They softened their fall by opening their elytrae and wings. The inability to fly could result from change in their flying behavior compared to control adults. The flight duration was very much shorter, and the wing beat frequency and, more particularly, the amplitude of the wing beats were clearly lower. More time was spent in the other components of the flight behavior such as wing rotation, wing immobility, and wing folding. The sequence of these patterns differed slightly, due mainly to change in their frequency. The locomotor behavior was not modified by the mutation, which affected only the wing muscles. Searching behavior of mutant adults differed from that of control adults only in that they took longer to encounter and ingest aphids. Nevertheless, the larval growth and reproductive rate remained unchanged. The behavioral and biological features of these flightless adults indicate that it should be possible to use them in biological control programs.     

Ovipositional Rhythmicity in Ladybirds (Coleoptera: Coccinellidae): A Laboratory Study

Complete and diel ovipositional rhythms have been observed in three aphidophagous ladybirds (Coleoptera: Coccinellidae): Coccinella septempunctata Linnaeus, Coccinella transversalis Fabricius and Propylea dissecta (Mulsant) for the first time. The complete ovipositional rhythm could be described in terms of a polynomial curve and daily fluctuations in all three species. In the diel cycle, all three ladybird species oviposited significantly in the scotophase. C. septempunctata females preferred to oviposit at the end of scotophase in the early morning hours (0300-0700), P. dissecta laid most eggs in the middle of scotophase during the night (2100-2300), and C. transversalis laid most at the beginning of the scotophase at dusk (1700-1900). While the diel ovipositional rhythm of C. septempunctata and P. dissecta did not differ between days, that of C. transversalis changed dramatically; there was a single peak on the first and second day of observation, and four oviposition peaks on the fifth day with the peaks being situated in the two hours preceding and succeeding the onset and end of the scotophase. Diel rhythms of C. septempunctata and P. dissecta appear to be endogenous in nature while that of C. transversalis is partly modified by exogenous factors.     

Predatory responses of selected lines of developmental variants of ladybird,Propylea dissecta (Coleoptera: Coccinellidae) in relation to increasing prey and predator densities

Individuals of the same species, population and generation frequently exhibit sub-maximal and significant genetic and phenotypic variation in their rate of development, showing slow and fast developers. Fast developers commonly have higher foraging and predation rates than slow developers. The consequence of such differences and foraging for the efficacy of biocontrol species remains under-explored. Slow and fast developers from a population of the ladybird, Propylea dissecta were separated and selected experimentally for F15 generations, and the predatory response of fourth instar larvae of control and experimentally selected slow and fast developers was then assessed at differing levels of prey (pea aphid, Acyrthosiphon pisum) and conspecific predator abundance. All individuals, whether slow or fast developers, showed a Type-II functional response, decrease in proportion of prey consumed with increasing prey biomass and an increase in proportion of prey consumed with increasing predator density. The proportion of prey consumed was highest in experimental fast developers and lowest in experimental slow developers. Attack rate was highest and handling time longest in slow developers of control/experimental groups. Mutual interference was least while area of discovery was highest in experimental fast developers. Thus, selection of fast developers for F15 generations led to higher functional responses, slower attack rates and faster prey consumption. This lower mutual interference and high searching efficiency indicates that they can be experimentally selected and used for better control of the pea aphids. This study is the first attempt to evaluate predatory responses of selected lines of an aphidophagous ladybird.     

Functional response of two aphidophagous ladybirds searching in tandem

The functional response of two ladybird species, Propylea dissecta (Mulsant) and Coccinella transversalis Fabricius in combination was studied in an effort to determine the impact of two predators on the increasing density of aphid, Aphis gossypii Glover infesting leaves of bottle gourd, Lagenaria vulgaris Seringe. The ladybirds were exposed to prey density in three combinations using adult females: (i) one P. dissecta and one C. transversalis together, (ii) two P. dissecta, and (iii) two C. transversalis, for 24 h. Prey consumption increased curvilinearly with prey density in all three combinations. It was maximum when heterospecifics searched together. The prey were handled and consumed significantly faster (9.51 min) by heterospecifics in combination, followed by two C. transversalis (11.32 min) and two P. dissecta (12.52 min). The attack rate was maximum (0.00274) by heterospecifics followed by the two C. transversalis (0.00166) and two P. dissecta (0.00092). The synergistic effect on the prey consumption may possibly be due to spatial distribution of P. dissecta and C. transversalis in terms of feeding sites, with the former being negatively, and the latter being positively geotactic. Field release of two predator species in tandem is recommended for biocontrol of A. gossypii because of the potential for synergistic action by these natural enemies.     

龟纹瓢虫卵黄蛋白的分子特性及发生动态

研究了龟纹瓢虫Propylea japonica (Thunberg) 卵黄蛋白的基本特性以及卵黄发生过程中卵黄蛋白的动态变化。PAGE和SDS-PAGE实验表明,龟纹瓢虫卵黄蛋白分子量为294.81±40.70 kD,并由分子量分别为144.68±0.03 kD和51.23±0.27 kD的两种亚基组成。对卵黄蛋白的氨基酸组成和含量分析发现,其必需氨基酸总量占57.48％,略高于非必需氨基酸,其中谷氨酸（Glu）含量最高,为15.26％;色氨酸（Trp）和蛋氨酸（Met）含量较低,分别为0.50％和0.11％。采用间接竞争ELISA法,系统测定了龟纹瓢虫成虫期脂肪体、血淋巴和卵巢中卵黄蛋白的动态变化,结果表明：脂肪体是卵黄原蛋白合成的场所,卵黄原蛋白的合成始于羽化后第2天;脂肪体、血淋巴和卵巢中卵黄原蛋白的滴度在羽化后第4天开始迅速上升,至成虫期的第8天左右达到高峰期。     

二双斑唇瓢虫对矢尖蚧的捕食作用

研究了二双斑唇瓢虫（Chilocorus bijugus Mulsant)对矢尖蚧（Unaspis yanonensis Kuwana)的捕食作用,结果表明,瓢虫雌成虫对矢尖蚧各虫态的功能反应呈Holling II型,瓢虫对矢尖蚧的捕食效应随捕食者个体间干扰作用的增加而下降,捕食作用率（E）随着瓢虫数增加呈幂函数下降曲线,温度对瓢虫的捕食效应具有显著的影响,寻找效率（a)和处置时间（Th)与温度之间呈二次函数关系,猎物密度对瓢虫生殖力的影响呈Logistic曲线。     

Evaluation of Serangium n. sp. (Col., Coccinellidae), a predator of Bemisia tabaci (Hom., Aleyrodidae) on cassava

Abstract  The potential of a new, previously unidentified Serangium species (Col., Coccinellidae) to control the high Bemisia tabaci (Gennadius) (Hom., Aleyrodidae) populations on cassava was evaluated. Field and laboratory studies were carried out to determine the abundance and feeding capacity of this Serangium species feeding on B. tabaci on cassava. Serangium nymphs and adults were most abundant in cassava fields late in the season, rising sharply from 5 months after planting (MAP) to a peak at 7–8 MAP. Pre-imaginal development averaged 21.2 days and was longest in eggs and shortest in the L₁ instar. Mean total prey consumption of immature Serangium increased with the stage of development with the lowest consumption in the L₁ instar and highest in the L₄ instar. Mean daily consumption was lowest on the first day after hatching in the L₁ instar and rose to a peak on the 13th day after hatching in the L₄ instar. Each Serangium larva consumed a mean of over 1000 nymphs during its entire development. These results have demonstrated the potential of this Serangium species to control B. tabaci populations on cassava.     

Evolutionary significance of promiscuity in an aphidophagous ladybird, Propylea dissecta (Coleoptera: Coccinellidae)

The short- and long-term consequences of monogamy and promiscuity in an aphidophagous ladybird, Propylea dissecta (Mulsant) in terms of reproductive performance and offspring fitness were investigated. Promiscuous females were more fecund and laid more viable eggs than monogamous ones. Amongst promiscuous females, those mated with multiple males (freedom for mate choice) had significantly higher reproductive output than those mated daily with a new unmated male. The increased reproductive output of females mated with an unmated male each day in comparison to monogamous ones might possibly be a result of their increased ejaculate size. The further increase in reproductive output, when a female was allowed to choose a mate, may be because of repeated preferential mating with the fitter males. Results on long-term consequences of promiscuity revealed faster development and increased survival in offspring of promiscuous females at 25, 27 and 30 degrees C. The percent adult emergence was more strongly influenced than development by the mating regime. There were no statistically significant differences in survival of offspring of promiscuous females (mate choice) at the three temperatures, while the percent survival of offspring of monogamous females differed significantly. Offspring of promiscuous females (mate choice) were fittest in terms of development and survival, which indicates their probable better genetic constitution and subsequently that of chosen mates. Thus, promiscuity has both short- and long-term benefits and is advantageous to ladybirds. The presence of long-term benefits in terms of fitter offspring probably indicates the evolutionary rationale behind the prevalence of promiscuity in ladybirds.     

Effects of cotton cultivars differing in gossypol content on the quality of Aphis gossypii as prey for two species of Coccinellidae

Plants can have detrimental effects on biological control agents by affecting their prey or host quality. Thus, it is important to understand the tri-trophic interactions between plants, herbivores and natural enemies when implementing biological control programmes. Studies have shown that both morphological and chemical traits of host plants can affect the third trophic level. Cotton plants are known to produce alkaloids such as gossypol, a sesquiterpene aldehyde that can confer resistance against herbivorous arthropods. Nevertheless, little is known about the effect of gossypol on biological control agents. In this study, we investigated how three cotton cultivars (BRS Rubi, BRS Safira and BRS Verde) differing in gossypol content affect development and growth of predatory coccinellids, Eriopis connexa and Harmonia axyridis, feeding on the cotton aphid Aphis gossypii reared on those cultivars. The results show that the cultivar BRS Rubi (highest gossypol content) had a sub-lethal effect on the development and growth of both Coccinellidae species compared with the other cultivars. Overall, the cultivar BRS Rubi reduced slightly fecundity, net reproductive rate and intrinsic rate of natural increase for both Coccinellidae species. However, because aphid populations stay short periods of time in the field, and adult coccinellids may supplement its diet with alternative prey and plant material this sub-lethal indirect effect of gossypol may not have a detrimental effect on field biological control of cotton aphid by either E. connexa or H. axyridis, thus suggesting a compatibility between plant resistance and biological control agents.     

Reproductive performance of four aphidophagous ladybirds on cowpea aphid, Aphis craccivora Koch

Abstract:  This study investigated prey consumption, egg production, percent progeny loss, reproductive, pre- and post-reproductive periods, reproductive time ratio, reproductive rate and bioconversion efficiency of four aphidophagous ladybirds, viz. Cheilomenes sexmaculata (Fabricius), Coccinella septempunctata Linnaeus, Coccinella transversalis Fabricius and Propylea dissecta (Mulsant) on Dolichos lablab Linnaeus infested with cowpea aphid, Aphis craccivora Koch. C. sexmaculata had the highest bioconversion efficiency, reproductive rate and reproductive time ratio followed in rank order by P. dissecta , C. transversalis and C. septempunctata . This study indicates that C. sexmaculata has a narrow ecological relationship with A. craccivora . The increased allocation of resources to reproduction as indicated through a high reproductive time ratio and high bioconversion efficiency of C. sexmaculata and P. dissecta suggest that they may be better adapted to compete for this prey with larger species like C. transversalis and C. septempunctata .     

黄玛草蛉捕食米蛾卵的功能反应与数值反应

针对天敌控害潜能评价和天敌昆虫大量饲养的基础理论问题,以一种重要的捕食性天敌黄玛草蛉Mallada besalis Walker捕食米蛾卵的功能反应和数值反应研究为例,开展研究。结果表明,黄玛草蛉幼虫对米蛾卵的捕食功能反应符合HollingⅡ型方程,其捕食量在一定范围内随着猎物密度的增加而增加,最后趋近一个稳定值,且与是否供水有关,供水时3龄幼虫的理论捕食能力最大,为3205.1粒卵/d。数值反应研究结果表明,不同密度的米蛾卵对黄玛草蛉的生长、发育、繁殖有着显著的影响。随着猎物密度增大,黄玛草蛉幼虫的发育速率、雌虫的产卵量、孵化率及雌性比增大,成虫的寿命延长,最后趋于稳定。米蛾卵密度5、10、20、30、50卵/d。分别是黄玛草蛉幼虫存活、幼虫化蛹、成虫羽化、后代卵孵化及世代延续的临界值。并讨论了功能反应和数值反应研究在天敌利用研究中的侧重点和重要性。