Physiological mechanism governing slow and fast development in predatory ladybirds

Aphidophagous and coccidophagous ladybirds, similar to their prey, show marked differences in their pace of life (Dixon, 2000), in particular in their rate of development, with all stages of aphidophagous species developing much faster than those of coccidophagous species. Two hypotheses are proposed to account for the large difference in the pace of life of these two groups. These are that differences in the rate of development are a result of differences in lower temperature thresholds for development or the quality of their respective prey as food (Dixon et al., 2011). Analysis of published results on the rates of development of the eggs of ladybirds indicates that the inverse relationships between the number of day‐degrees required for development (K) and the lower temperature threshold for development (td_min) of these two groups are significantly different. In particular, the respective td_min overlap and K of the aphidophagous and coccidophagous species with a similar td_min are, on average, 38 and 117 day‐degrees (D^o). The relationship between the rate of development (R) and temperature (T) for aphids reared on poor‐ or high‐quality foods indicates that, although the value of td_min of a species depends on food quality, K does not, showing that it is unlikely that K is governed by food quality. Thus, there is little support for differences in either the td_min or food quality governing the difference in the pace of life of these two groups of ladybirds. The results indicate that the physiological mechanism that may govern the difference in the pace of life between these two groups is the number of day‐degrees (K) needed to complete their development. The possible evolutionary reason for this is discussed.     

天敌瓢虫种群遗传学及其在生物防治中的应用

天敌瓢虫常在农业生产的生物防治中被广泛引进和使用。这种使用过程往往会导致天敌瓢虫在遗传水平的改变,并最终影响到其生物防治的使用效果和本地生态多样性。对天敌瓢虫种群遗传学的研究,有助于我们认识甚至预测这种遗传改变的规律和生态机制,从而为生物防治计划的优化提供理论基础。本文介绍了种群遗传学的研究方法,分析在生物防治中促进天敌瓢虫遗传改变的因素,以及几种常见的天敌瓢虫种群演变情况,并讨论种群遗传学在天敌瓢虫研究的未来方向。     

Survival, development and life tables of two congeneric ladybirds in aphidophagous guilds

Two congeneric aphidophagous ladybirds, Coccinella septempunctata and Coccinella transversalis, were reared on three aphid species, Lipaphis erysimi, Myzus persicae and Aphis nerii, to estimate the effect of prey quality and intra- and interspecific interactions on their survival and development of life stages. Mortality of first instar ladybirds of both species was highest feeding on A. nerii. Preimaginal mortafity was lowest when feeding on L. erysimi （C. septempunctata, 1.6% and C. transversalis, 3.2%）, and highest when feeding on A. nerii （ C. septempunctata, 6.2% and C. transversalis, 8.2%）. Comparatively higher weight and larger size of C. septempunctata along with the lower levels of mortality recorded suggested that it is more likely to have acted as an intraguild predator than C. transversalis. High recorded mortality of C. transversalis is attributed to probable intraguild predation on account of its smaller size. The major sources of mortality were probably cannibalism, intraguild predation and other unknown factors. Lower prey quality increased the incidence of cannibalism and intraguild predation, especially in C. transversalis. The investigation suggests an intrinsic competitive advantage for C. septempunctata over C. transversalis in guilds of three aphid species.     

Laboratory study of cannibalism and interspecific predation in ladybirds

Abstract.

• 1 In the absence of aphids, adult females of Adalia bipunctata (L.) showed a greater reluctance to eat eggs than males.
• 2 Eggs and young larvae were more vulnerable to cannibalism than older larvae and starved larvae were more vulnerable than well-fed larvae.
• 3 Both egg and larval cannibalism is inversely related to the abundance of aphids.
• 4 Eggs are a better food, in terms of larval growth and survival, than aphids.
• 5 In the absence of aphids interspecific predation occurred, but not equally, between the coccinellids A.bipunctata, A.decempunctata (L.), Coccinella septempunctata L. and C.undecempunctata L.
• 6 Larvae and adults of A. bipunctata and C.septempunctata were reluctant to eat conspecific eggs painted with a water extract of the other species' eggs and larvae of C. septempunctata were more likely to die after eating a few eggs of A.bipunctata than vice versa.
• 7 These results indicate that cannibalism occurs mainly when aphid prey is scarce and is adaptive in that it improves the chances of survival, and coccinellids, to varying degrees, are defended against interspecific predation.

     

温度和光周期对两种瓢虫捕食习性和发育的影响

[目的]在农业上,气候变化影响作物产量、蚜虫动态以及捕食性瓢虫对蚜虫的调节作用.一些瓢虫为广布种,而另一些瓢虫种类却只在特定地理范围内分布,而有关气候因子如何影响瓢虫分布了解甚少.[方法]评价不同温度(15℃,20℃,25℃,30℃和35℃)和光周期(8L:16D,12L:12D和16L: 8D)对两种瓢虫[七星瓢虫Coccinella septempunctata (Linnaeus)(一种广布种)和狭臀瓢虫Coccinella transversalis Fabricius(一种东方/南亚瓢虫)]幼虫取食和利用豌豆蚜Acyrthosiphon pisum(Harris)的影响.[结果]尽管两种瓢虫的捕食作用和发育速率均在25℃时最高,但是随着日长的增加(即光周期从8L:16D增加到16L:8D)捕食作用进一步增强.与狭臀瓢虫相比,在所有温度和光周期下,七星瓢虫幼虫的取食量更高,生长和发育速率更快,其新羽化的成虫体重更重.非线性回归模型显示,七星瓢虫和狭臀瓢虫的最低发育温度分别为11℃和10.5℃左右,最高发育温度分别为41℃和43℃左右.狭臀瓢虫的幼虫捕食和发育速率对气候因子的依赖性强于七星瓢虫.[结论]本研究表明,广布种瓢虫对气候因子的敏感性比土著种瓢虫弱,因而可以广泛分布于不同的地理区域.因此,与土著种瓢虫相比,广布种瓢虫更适于作为生防因子.     

不同食物对甜菜夜蛾生长发育参数的影响

选择上海地区近年甜菜夜蛾Spodoptera exigua(Hübner)发生比较严重的甘蓝、豇豆、大葱、芦笋4种寄主植物,在温度(27±1)℃,相对湿度70%±5%,光照L:D=14:10的条件下研究了不同寄主植物对甜菜夜蛾发育特征的影响。结果表明,不同寄主植物饲喂甜菜夜蛾,甜菜夜蛾幼虫发育历期、存活率、蛹重、成虫寿命、净生殖力和内禀增长率等参数在寄主植物间有显著差异。用甘蓝饲喂的甜菜夜蛾雌虫产卵量明显高于取食大葱和芦笋的。同样,取食甘蓝的甜菜夜蛾蛹重、净生殖力和种群内禀增长率等参数明显高于取食大葱和芦笋的,说明甘蓝与其他寄主植物相比提供了较好质量的食物。     

多杀菌素和氟虫睛对瓢虫、寄生蜂和菜缢管蚜选择性的研究

采用药膜法研究了多杀菌素和氟虫睛对3种瓢虫、2种寄生蜂和菜缢管蚜的选择性。与菜缢管蚜相比,2种杀虫剂对3种瓢虫的毒性较低(LC50比值=1.73-41.9),尤其是氟虫睛对3种瓢虫以及多杀菌素对稻红瓢虫均具有较好的选择性(LC50比值大于5)。菜蚜茧蜂和蚜虫宽缘金小蜂对2种杀虫剂的敏感性远高于3种瓢虫和菜缢管蚜,2种杀虫剂对菜蚜茧蜂和蚜虫宽缘金小蜂均是高毒的(LC50比值=0.006-0.05)。3种瓢虫对多杀菌素和氟虫睛具有较高的耐药性,这将有助于其在生物防治中的应用。     

The influence of different aphid prey species on the biology and life table parameters of Propylaea japonica

The influence of three aphid prey species – Aphis craccivora Koch, Megoura viciae Buckton and Rhopalosiphum maidis (Fitch) – on the biology of Propylaea japonica (Thunberg) was studied under laboratory conditions. The development, survivorship, longevity, reproduction and life table parameters of P. japonica differed significantly among different treatments. The shortest developmental period of P. japonica (from first-instar larvae to adult) was observed on A. craccivora, whereas the longest was observed on M. viciae. The highest survivorship was observed on A. craccivora, and the lowest survivorship was observed on M. viciae. The highest sex ratio, fecundity and the shortest pre-oviposition period were observed when A. craccivora was used as prey. The longevities of P. japonica females and males did not differ significantly when reared on different aphid species. The highest values of net reproductive rate, intrinsic rate of increase and finite rate of increase were observed on A. craccivora. The results suggest that A. craccivora is a suitable prey for P. japonica among the three aphid species tested and can serve as a diet for the mass rearing of P. japonica under laboratory conditions for possible use in integrated pest management.     

Intrinsic prey suitability in specialist and generalist Harmonia ladybirds: a test of the trade‐off hypothesis for food specialization

The trade‐off hypothesis posits that increased performance on a given resource comes at the cost of decreased performance on other resources, and that this trade‐off is a driving force of food specialization in both predators and herbivores. In this study, we examined larval survival and performance in two sibling ladybird species, Harmonia yedoensis Takizawa and Harmonia axyridis Pallas (Coleoptera: Coccinellidae), fed on one of four prey species. Harmonia yedoensis is a specialist predator that preys mostly on pine aphids in the field, whereas H. axyridis is a generalist predator with a broad prey range. We experimentally showed in the laboratory that larval survival and performance were not higher when H. yedoensis was fed on pine aphids, compared with the other prey species. Rather, prey suitability was similar in both ladybird species, and H. yedoensis larvae developed as well or even better on prey species that they never utilize in nature. These results suggest that the host range in H. yedoensis may not be limited by the intrinsic suitability of the aphid species per se. Moreover, as shown by our previous study, the pine aphid is a highly elusive prey that is difficult for small ladybird hatchlings to capture, which means that the cost of utilizing this prey is high. Therefore, we conclude that some factor other than prey suitability is responsible for the observed food specialization in H. yedoensis.     

Evolution of avian eggshell structure

Data are presented suggesting that birds have evolved eggs with shells containing different structures (numbers of mammillae per unit of inner eggshell surface area, i.e., mammillary densities) to cope up with different calcium requirements imposed by different growth rates and modes of development. Precocial bird species grow slowly, but have high mammillary density, while altricial bird species grow rapidly, but have low mammillary density. These results suggest an adaptation associated with growth rate and mode of development and show, moreover, that the mammillary layer is indicative of the breeding biology of the bird. J. Morphol. 2012. © 2011 Wiley Periodicals, Inc.     

Development and immature survival of two aphidophagous ladybirds, Coelophora biplagiata and Micraspis discolor

The developmental parameters, namely developmental periods, larval survival, adult emergence, growth index, developmental rate and sex ratio in terms of female proportion in a population of two aphidophagous ladybirds, namely, Coelophora biplagiata （Swartz） and Micraspis discolor （Fabricius） were investigated, using three aphid species as prey. The immature stages of C. biplagiata were significantly affected by prey quality and developed fastest when fed on Aphis craccivora followed by Aphis gossypii and Rhopalosiphum maidis. The order of prey suitability in terms of developmental rate of immature stages of M. discolor was just reversed. The adult females were heavier than males, while pre-pupae were heavier than pupae in both ladybirds. The sex ratio of adult emergence, though female-biased in both ladybird species, was not significantly affected by prey quality.     

Interspecific interactions in a tri-trophic arthropod system: effects of a spider on the survival of larvae of three predatory ladybirds in relation to aphids

The effects of the crab spider, Misumenops tricuspidatus (Fabricius), on the larval survival of three ladybird species, Harmonia axyridis Pallas, Coccinella septempunctata L., and Propylea japonica L., in relation to aphids were investigated in the laboratory. Predation by the spider on the three ladybird species differed. All the larvae of C. septempunctata, none of H. axyridis, and an intermediate number of P. japonica were attacked and eaten by the spider. All the larvae of H. axyridis suffered mortality due to cannibalism or starvation in the treatments with and without a spider. In case of C. septempunctata, however, mortality in the early instars was significantly greater in the treatment with a spider than without a spider and no larvae developed into pupae due to predation. In the treatment without a spider, the majority of the larvae in the former treatment suffered mortality due to cannibalism or starvation, and only 13.3% of larvae developed into the adult stage. In the case of P. japonica, mortality was mainly attributed to predation in the treatment with a spider and only 26.7% became adult. In comparison, 86.7% of larvae survived to the adult stage in the treatment without a spider. In addition, in both H. axyridis and C. septempunctata, the development of young larvae was significantly slower in the presence of a spider, but this was not the case with the older larvae of H. axyridis, which indicates that the effect of the spider on larval development changed with the developmental stage of the larvae in this species. However, the spider had no significant effect on the developmental time of P. japonica larvae. Although both the spider and the ladybirds significantly affected the number of aphids, they did not have an additive effect on aphid abundance. The interactions between the spider and the ladybirds, such as intraguild predation or competition, caused them to reduce aphid population density less than the ladybirds did on their own. The effect of the spider on the larval performance of three predatory ladybirds was found to be unequal in terms of their vulnerability to predation and rate of larval development and it depended on the species and developmental stage of the ladybird.     

The effect of temperature on the growth,development and survival of Macropetasma africanus (Balss) (Penaeoidea:Penaeidae) larvae reared in the laboratory

Macropetasma africanus (Balss) has been successfully spawned and its larvae reared under controlled laboratory conditions. The relationship between egg number (E) and female total length (L) was E = 18.59 L^2.11. An experiment was designed to test the effect of temperature on larval development, survival and growth. Temperature effected larval development time, from 13–15 days at 25°C, to 25 days at 15°C (nauplius 1 to post-larva). Mortality was low for the naupliar stages at 25, 22 and 18°C, while at 15°C only 52% of the larvae reached nauplius 6. Mortality was highest from nauplius 6 to protozoea 1 (17, 21, and 18% at 25, 22, and 18°C, respectively), but decreased considerably for all temperatures once the mysis stage was reached. Overall survival rates from nauplius 1 to post-larva decreased with decreasing temperature (65, 54, 48, and 39% at 25, 22, 18, and 15°C respectively). Temperature also significantly affected larval growth. At 25°C mean total length was significantly (P < 0.05) larger than at 15°C (protozoea 2 to post-larva), while from protozoea 3 to post-larva total length differences were significantly different (P < 0.05) between 18 and 25°C. M. africanus has a major spawning peak in summer, suggesting that there may be a selective advantage to reproducing during the warmer months.     

Evolution of the self-pollinating flower in Clarkia xantiana (Onagraceae). I. Size and development of floral organs

Clarkia xantiana has two subspecies that differ in breeding system: ssp. xantiana, which is outcrossing, and ssp. parviflora, which is self-fertilizing. Outcrossing is the ancestral breeding system for the genus Clarkia. Flowers of ssp. parviflora have characteristics commonly associated with selfing taxa: they are smaller and have little temporal and spatial separation between mature anthers and stigma (dichogamy and herkogamy, respectively). Flower morphology and development were studied in four populations of each subspecies to establish the developmental changes that occurred in the evolution of selfing. In particular, we sought to evaluate the hypothesis that the selfing flower may have arisen as a byproduct of selection for rapid maturation in the arid environment occupied by ssp. parviflora. This hypothesis predicts that development time should be reduced in spp. parviflora relative to ssp. xantiana. We also sought to compare the pattern of covariation of flower morphology and development between subspecies to that within subspecies. Similar within vs. between patterns of covariation could be indicative of developmental or functional constraints on the independent evolution of floral parts. In spite of significant variation among populations within subspecies, the subspecies clearly differ in flower morphology and development. All floral organs, except ovaries, are smaller in ssp. parviflora than in ssp. xantiana. The flower plastochron, the duration of flower development from bud initiation to anthesis, and the duration of protandry are all shorter in ssp. parviflora than in ssp. xantiana. Maximum relative growth rates are higher for all organs in ssp. parviflora than in ssp. xantiana. Thus, progenesis (i.e., via a reduction in development time) is combined with growth acceleration in the evolution of the selfing flower. Since reduced development time and growth acceleration both allow selfing flowers to mature earlier than outcrossing ones, selection for early maturation may have contributed to the evolution of the selfing flower form. The pattern of trait covariation differs within spp. parviflora relative to the patterns within spp. xantiana and between the two subspecies, suggesting that floral parts can and have evolved independently of one another.     

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.     

Evolution of cannibalism and female's response to oviposition-deterring pheromone in aphidophagous predators

1.  Egg cannibalism by larvae is common in Coccinellidae and is known to be advantageous for the cannibals. Furthermore, larvae of aphidophagous ladybirds usually produce an oviposition-deterring pheromone (ODP), which inhibits oviposition by adult females. It has been proposed that the response to ODP has evolved because of the high costs of cannibalism. However, this has never been formally proved.
2.  In this paper, we study the theoretical evolution of this system. We first look at the conditions under which cannibalism and the response to ODP can evolve. Subsequently, we examine the occurrence of polymorphism both in the production of larval tracks and in the sensitivity of females to specific pheromones.
3.  The models predict that the amount of cannibalism should not depend on prey density and that evolution should lead to a continuous increase in cannibalism, and consequently larvae should always cannibalize eggs when possible. In response to the cost of cannibalism, ODP recognition can evolve, so that females avoid laying eggs in patches of prey already occupied by conspecific larvae. The result is an arms race between larvae and adult females, which favours a diversification of ODP pheromones. Our models show that: (i) females should be able to recognize mixtures of hydrocarbons rather than a single molecule; and (ii) females should be more sensitive to the tracks of their own offspring than those of non-related larvae.     

4种蚜虫对异色瓢虫生长发育和繁殖的影响

为明确桃蚜、萝卜蚜、胡萝卜微管蚜、瓜蚜对异色瓢虫种群扩繁的适宜性,在实验室条件下,以4种蚜为食料构建异色瓢虫两性生命表,研究不同蚜虫对异色瓢虫生长发育和繁殖的影响。结果表明: 取食不同蚜虫,异色瓢虫未成熟期由长到短依次为萝卜蚜(18.18 d)、瓜蚜(17.48 d)、胡萝卜微管蚜(16.23 d)、桃蚜(15.77 d);成虫前期存活率由高到低依次为胡萝卜微管蚜(88.3%)、桃蚜(86.7%)、萝卜蚜(55.0%)、瓜蚜(55.0%);成虫繁殖力由高到低依次为胡萝卜微管蚜(1750.5粒)、桃蚜(1441.5粒)、瓜蚜(1006.3粒)、萝卜蚜(965.2粒);成虫寿命由长到短依次为胡萝卜微管蚜(78.8 d)、桃蚜(63.1 d)、瓜蚜(54.3 d)、萝卜蚜(48.4 d);异色瓢虫种群内禀增长率r_m由高到低依次为桃蚜(0.19)、胡萝卜微管蚜(0.18)、萝卜蚜(0.14)、棉蚜(0.14);净增值率R₀由高到低依次为胡萝卜微管蚜(895.83)、桃蚜(600.62)、萝卜蚜(273.47)、棉蚜(268.33)。4种蚜虫中,胡萝卜微管蚜和桃蚜更有利于异色瓢虫种群扩繁。     

Green leaves enhance the growth and development of a stream macroinvertebrate shredder when senescent leaves are available

1. Freshly fallen green leaves and flowers of terrestrial plants enter temperate streams in spring and summer, when senescent leaf litter is often scarce. These resources appear to provide good supplementary food for macroinvertebrate shredders, but have some potential shortcomings as food or case material for caddisflies. 2. To compare suitability of green leaves or flowers and senescent leaves for the growth and development of stream shredders, we reared the caddisfly Lepidostoma complicatum in the laboratory with treatments that provided larvae with senescent (oak) and green (oak or maple) leaves separately, and also together, in case the combined use of both types of leaf may benefit the shredder. 3. Larvae supplied with green leaves alone grew at 65% of the rate of those provided with senescent leaves alone, due to their lower consumption rate. No individuals given green leaves alone developed into adults, whereas 70% of the individuals given senescent leaves alone did. Green leaves may inhibit larval consumption due to their high phenol content, or they may be unsuitable for case material because they are less tough than senescent leaves. 4. Larvae supplied with both senescent and green leaves (or flowers) had a higher growth rate and developed faster, than those given senescent leaves alone, whereas the proportions of successfully emerged individuals did not differ. Lepidostoma probably benefits from the higher nitrogen content of the green leaves when used together with senescent leaves. 5. These results suggest that green leaves (or flowers) cannot serve as an alternative food resource to senescent leaves, but that they can enhance the growth and development of a Lepidostoma stream shredder if senescent leaves are also available.