Egg dumping by predatory insects

Synovigenic insects resorb oocytes when food is scarce and mature oocytes when food is plentiful. These two antagonistic processes allow an optimal allocation of resources to reproduction and somatic functions. Unlike hymenopteran parasitoids, ladybirds cannot resorb mature oocytes present in the oviducts. Is the energy contained in these oocytes lost or is there a mechanism for recovering it when needed? Females of two species of ladybird beetles Adalia bipunctata (L.) and Adalia decempunctata (L.) that are starved for >24 h lay single infertile eggs, which they immediately eat, and these eggs comprise the mature oocytes in the oviducts at the onset of starvation. This behaviour has some similarities to egg dumping reported in herbivorous insects and is part, in ladybird beetles, of a process to retrieve energy invested in reproduction. Such behaviour may exist in other predatory synovigenic insects species that do not invest in maternal care.     

Beyond fecundity and longevity: trade-offs between reproduction and survival mediated by behavioural responses of the seed beetle, Callosobruchus maculatus

Abstract.  Many studies of life-history traits have failed to find trade-offs where they are predicted by theory. A hypothesis that explains the lack of trade-offs between fecundity and longevity in the seed beetle, Callosobruchus maculatus , is proposed. By manipulating host availability time and host size, trade-offs mediated by behavioural responses of the female to adapt to environmental change are tested. Females show no decrease in lifetime fecundity when host availability time is limited to only 4 h on each day. However, longevity significantly increases when the female is provided with small beans after host deprivation. Because neither acquisition, nor utilization by females of these four manipulation treatments significantly differs, studies are carried out to demonstrate whether the energy shifted from increased longevity without decreasing fecundity. Providing abundant small or large beans each day directly after host deprivation, significantly increases the number of daily eggs laid by the female for several days, whereas the female decreases the uniformity of her egg dispersion only when small beans are provided. Therefore, the female shows a response to a change in the environment by adjusting egg-laying rate and/or egg-dispersion pattern. This may change the traits of reproduction and survival. Because energy allocations can be shifted between components of reproduction (e.g. host-selection behaviour and fecundity) or between reproduction and survival, fecundity and longevity may be inappropriate indices for trade-off analyses in this study. A framework for exploring the costs of reproduction mediated by physiological and behavioural changes in C. maculatus is proposed and discussed.     

Egg maturation and oviposition strategy of a capital breeder,Cleorodes lichenaria,feeding on lichens at the larval stage

Abstract 1. Capital breeders rely solely on resources acquired before breeding, whereas income breeders may obtain considerable amounts of resources following the commencement of reproduction. In income breeders oviposition occurs over a longer time period with a small number of eggs ready to be laid at the start of adult life, whereas capital breeders reproduce more rapidly and contain higher numbers of mature eggs at eclosion relative to potential fecundity. 2. This study explored egg maturation and oviposition strategy of a nocturnal geometrid moth, Cleorodes lichenaria. It was predicted, based on the long larval period of C. lichenaria and the known biology of other geometrids, that this lichen‐feeding geometrid moth concentrates egg production at the beginning of its breeding period and that external nutrients have a limited role on its fecundity. 3. Approximately 46% of eggs were ready to be laid in newly hatched females and approximately 40% of their potential fecundity was realised during the first night, after which the number and the weight of eggs decreased steadily. Feeding or drinking did not increase the fecundity of females. Pupal mass correlated positively with realised fecundity and ovigeny index. 4. Reproduction of C. lichenaria is based solely on larval‐derived resources. It is suggested that females are able to resorb thoracic tissues to increase fecundity and that this ability is probably linked to relatively long lifespan and low ovigeny index of this species, characteristics rarely observed in other geometrid moths.     

Factors affecting egg maturation in the bean weevil Callosobruchus maculatus

Abstract. This paper concerns the effects of mate and seed availability on the rate of egg maturation in the bean weevil Callosobruchus maculatus (F.). Egg maturation starts before emergence from the seed and, provided that both oviposition sites and mates are available, eggs are laid at a rate determined by the number of oviposition sites, and mature at a similar rate. If seeds or mates are absent then a small number of eggs are laid, but oocytes continue to mature until the oviduct-capacity is approached. The number of eggs that a female can store is dependent on her body weight and does not correlate with the number of ovarioles. If, after a period in which oocyte development has been halted, conditions for egg-laying become suitable, then egg maturation can be re-started, but only after the oviducts have been emptied of eggs. The rate of egg maturation is then similar to that for females of the same age which have been maturing eggs since emergence.     

Host preference of Callosobruchus maculatus: a comparison of life history characteristics for three strains of beetles on two varieties of cowpea

Abstract: The reproductive success of Callosobruchus maculatus Fabricius, the main insect pest of stored cowpea, may vary between strains of this beetle and between varieties of the host seeds. Life history parameters of beetle strains from three different origins in West Africa were compared on two susceptible varieties of cowpea, Vigna unguiculata (L.) Walp. All beetle strains were assayed in a no‐choice and a two‐choice test. No major differences were found between the beetle strains. In a no‐choice situation, the developmental period from egg to adult was prolonged on the bean variety Kpodjiguegue. In a two‐choice situation, the beetles showed a strong preference for the Californian blackeyed bean variety to oviposit on. Here again the development took longer on Kpodjiguegue beans and the intrinsic rate of increase of the beetle population was lower. Using either equal numbers of beans of the same size or equal weights of beans of undetermined size of the two bean varieties did not affect the outcome of the test.     

The function of multiple mating in oviposition and egg maturation in the seed beetle Callosobruchus maculatus

Abstract.  To comprehend the significance of multiple mating in female Callosobruchus maculatus , the material-benefits hypothesis is tested, as well as the potential function and mechanism of additional mating for females. The results show that longevity and lifetime fecundity are significantly higher in doubly-mated females, whereas females subject to an interrupted second copulation (which transfers no ejaculate) show resemblance with singly-mated females in lifetime fecundity and daily fecundity, supporting the material-benefits hypothesis. Female lifetime fecundity increases with ejaculate size in doubly-, but not singly-mated females. Doubly-mated females exhibit an immediate increase in fecundity on the day of remating. Moreover, dissection of ovaries after remating shows that the number of unlaid eggs is lower in doubly-mated females in comparison with singly-mated females, whereas the total number of unlaid and laid eggs shows no significant difference. This suggests that the second mating increases the oviposition rate, but not the egg maturation rate in female C. maculatus . Thus, ejaculate serves as a stimulus for oviposition, at least on the remating day. However, because doubly-mated females lay more eggs in total than singly-mated females, doubly-mated females appear to mature more eggs than the singly-mated counterparts. This is attributed to the 'indirect-driven' hypothesis, which states that the reduction in the number of mature eggs in the oviducts brought about by oviposition stimulates the maturation of oocytes in ovarioles to replenish the number of mature eggs.     

Egg and time limitation mediate an egg protection strategy

The number of mature eggs remaining in the ovaries and the time left for oviposition determine the reproductive decisions of the hyperdiverse guild of insects that require discrete and potentially limiting resources for oviposition (such as seeds, fruits or other insects). A female may run out of eggs before all available oviposition sites are used (egg limitation), or die before using all of her eggs (time limitation). Females are predicted to change clutch size depending on whether eggs or time is the limiting resource. We extend this framework and ask whether the same constraints influence a strategy in which females modify eggs into protective shields. In response to egg parasitism cues, female seed beetles (Mimosestes amicus) lay eggs in vertical groups of 2–4, modifying the top 1–3 eggs into shields in order to protect the bottom egg from attack by parasitoids. We made contrasting predictions of how egg and time limitation would influence egg size and the incidence and level of egg protection. By varying access to seed pods, we manipulated the number of remaining eggs a female had at the time she received a parasitism cue. Although egg size was not affected, our results confirm that egg‐limited females protected fewer eggs and time‐limited females protected more eggs. Female body size explained the number of eggs in a stack rather than host deprivation or the timing of parasitoid exposure. Our results clearly show that host availability relative to female age influences the incidence of egg protection in M. amicus. Furthermore, our study represents a novel use of life history theory to explain patterns in an unusual but compelling defensive behaviour.     

Seasonal life history trade-offs in two leafwing butterflies: Delaying reproductive development increases life expectancy

Surviving inhospitable periods or seasons may greatly affect fitness. Evidence of this exists in the prevalence of dormant stages in the life cycles of most insects. Here I focused on butterflies with distinct seasonal morphological types (not a genetic polymorphism) in which one morphological type, or form, delays reproduction until favorable conditions return, while the other form develops in an environment that favors direct reproduction. For two butterflies, Anaea aidea and A. andria, I tested the hypothesis that the development of each seasonal form involves a differential allocation of resources to survival at eclosion. I assayed differences in adult longevity among summer and winter forms in either a warm, active environment or a cool, calm environment. Winter form adults lived 40 times longer than summer form but only in calm, cool conditions. The magnitude of this difference provided compelling evidence that the winter form body plan and metabolic strategy (i.e. resource conservatism) favor long term survival. This research suggests that winter form adults maintain lowered metabolic rate, a common feature of diapause, to conserve resources and delay senescence while overwintering.     

Why resource history matters: age and oviposition history affect oviposition behaviour in exploiters of a mutualism

1. Acceptance of hosts for oviposition is often hardwired in short‐lived insects, but can be dynamic at the individual level due to variation in physiological state determinants such as ageing and prior oviposition. However, the effect of the oviposition history of resources together with time taken to accept less preferred hosts in ageing insects has rarely been investigated. 2. The time taken by parasitic fig wasps to accept resources with different oviposition histories was recorded in order to investigate the effect of wasp physiological state and resource oviposition history on oviposition behaviour. These wasps, which differ in life‐history traits, oviposit at specific developmental stages of enclosed fig inflorescences called syconia. 3. Behavioural assays were performed with naive wasps and wasps aged with and without prior oviposition experience. Syconia at the same developmental stage but differing in oviposition history were offered in no‐choice assays and the time taken to first oviposition attempt was recorded. 4. One short‐lived pro‐ovigenic galler species exhibited a decline with age in time taken to accept a syconium for oviposition. The exact timing of the transition from non‐acceptance to acceptance of less preferred syconia was determined in terms of the proportion of elapsed life span at the transition; this occurred at 25% of elapsed life span. 5. Longer‐lived parasitoids did not show any decline in specificity despite being aged for 50% of their life span. Therefore, host quality, trophic position, egg load and age may individually affect oviposition decisions or have interaction effects.     

Egg maturation dynamics of the parasitoid Microplitis rufiventris: starvation speeds maturation in early life

The number of mature eggs carried by a female parasitoid at any given moment (egg load) is a fitness‐related parameter affecting reproductive potential and impacting upon host population dynamics. Microplitis rufiventris Kokujev (Hymenoptera: Braconidae) is a solitary koinobiont endoparasitoid wasp of several noctuid pests, including Spodoptera littoralis. The number of mature eggs carried by females at emergence is approximately 50. The rate of egg maturation is strongly affected both by feeding status and access to host larvae. In early adult life, egg maturation rates are lower for 6–72 h in fed wasps compared with food‐deprived wasps. When given access to hosts, honey‐fed wasps live for approximately 9 days with high lifetime fecundity (226 eggs). By contrast to early adult life, the total realized fecundity is positively affected by feeding status, where water‐fed and starved females have 140 and 107 eggs, respectively. Egg resorption is most pronounced in the later life of females. The results suggest, in addition to confirming the effect of honey‐feeding on total fecundity, that fecundity of starved wasps includes rapid egg maturation early in life, which potentially could improve the performance of the parasitoid as a biological control agent.     

Functional anatomy of the egg pore in Callosobruchus maculatus: a trade-off between gas-exchange and protective functions?

Abstract. The chorion of the egg in Callosobruchus maculatus (Coleoptera: Bruchidae) provides protection but also has to allow exchange of respiratory gases. A single opening in the chorion (the egg pore) allows diffusion of gases, and young larvae die if the egg pore is blocked. Comparison of respiration rates using a Gilson respirometer showed an increase in oxygen uptake from the first to the seventh day after the egg was laid; the increase was from 6.8 to 11.2 pl/egg/day in a Brazil strain and from 8.6 to 23.5 pl/egg/day in a Yemen strain. The Yemen strain had double the metabolic rate of Brazil-strain insects in the larvae, which are exposed to seed toxins, but not in the non-feeding adults. It is suggested that the higher metabolic rate in the Yemen larvae is related to their ability to develop in seeds that are toxic to the Brazil strain. The egg pore also differed in size and shape between the two strains. The egg pore was cylindrical in the Brazil strain, but shorter and funnel-shaped with an increased external diameter in the Yemen strain. Calculation showed that the different shape and size of the Yemen egg pore would allow a 2–3-fold increase in gas conductance compared with the Brazil strain, and this increase was consistent with the enhanced oxygen requirement of the Yemen larvae, which was a little over twice that of the Brazil larvae at 7 days. The anatomy of egg pores in Cmaculatus thus seems to represent a trade-off between two functions, allowing adequate gas exchange while maintaining the protective function of the chorion.     

Environment-dependent reversal of a life history trade-off in the seed beetle Callosobruchus maculatus

Environmental manipulations have consistently demonstrated a cost of reproduction in the capital-breeding seed beetle, Callosobruchus maculatus, as females deprived of seeds or mates lay fewer eggs and thereby increase their longevity. Yet fecundity and longevity tend to be positively correlated within populations, perhaps as a consequence of individual differences in resource acquisition. We conducted a split-brood experiment that combined a manipulation of seed availability (seeds present or absent) with a quantitative-genetic analysis of fecundity and lifespan in each environment. Each trait was significantly heritable in each environment. Seed availability not only altered mean fecundity and longevity between environments, but also modified how the traits were correlated within environments. The signs of both the phenotypic and genetic correlations switched from positive when seeds were present to negative when seeds were absent. This reversal persisted even after the effect of body mass (a potential indicator of resource acquisition) was statistically controlled. Cross-environment genetic correlations were positive but significantly less than one for each trait. We suggest that the reversal of the fecundity-longevity relationship depends on a shift in the relative importance of resource-acquisition and resource-allocation loci between environments. In particular, a cost of reproduction may be apparent at the individual level only when seeds are scarce or absent because differences in reproductive effort become large enough to overwhelm differences in resource acquisition. Despite their common dependence on resources acquired during larval stages, fecundity and lifespan in C. maculatus do not appear to be tightly coupled in a physiological or genetic sense.     

Why do host‐deprived seed beetles ‘dump’ their eggs?

Abstract.  If host seeds are absent, females of the seed beetle Callosobruchus maculatus sometimes 'dump' eggs on unsuitable substrates, which causes complete larval mortality and decreases female lifespan. To understand the possible function of this behaviour, the present study examines genetic variation in dumping behaviour between and within populations. When deprived of hosts, females from an African population are much more prone to dump eggs than Asian females, most of which dump no eggs over their lifetimes. Egg dumping therefore cannot be explained as a simple, species-wide constraint imposed by the accumulation of mature oocytes. A transfer experiment tested the hypothesis that dumping eggs improves a female's ability to exploit the subsequent availability of seeds, perhaps by preventing a disruption of oocyte maturation. Contrary to prediction, the number of eggs laid after females are transferred to seeds is inversely related to the number dumped during deprivation. Two bidirectional selection experiments revealed heritable variation in egg-dumping behaviour within the African population. Deprived females from the dumper line dump more than twice as many eggs as do females from the nondumper line. Pre-adult development time is significantly longer in the nondumper line, which suggests that trade-offs with other fitness components could maintain variation in egg-laying behaviour within populations. The divergent responses to host availability by African and Asian females may represent a pleiotropic effect of similarly divergent responses to host quality.     

The fecundity and oviposition behavior of a South Indian strain of Callosobruchus maculatus

The fecundity of the South India strain of Callosobruchus maculatus (Fab.) (Bruchidae: Coleoptera) is 73 eggs for females developing in 31 days and increases to 94 eggs for females emerging after 41 days. Fed females lay 45–55 more eggs which is the egg-equivalent of the reserves unfed females had to sacrifice for activity. Competition as a larva did not reduce a female's fecundity. Oviposition is inhibited when females are given 25 or fewer beans for oviposition and it is reversed if females find fresh beans. Oviposition is generally released by beans carrying a below average number of eggs, thus, the eggs are almost uniformly distributed over the beans. Eggs are preferentially placed on the largest egg-free beans as expected if oviposition is released when a beetle goes from a smaller to a larger bean. The cues resulating host preferences, cowpea > mung pigeonpea chickpea, are secondary to the cues producing hyperdispersion and the selection of larger beans. We present a model for the oviposition behavior that accounts for the observed responses to species, size, and egg load of oviposition sites.

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Résumé La fécondité de femelles isolées avec un mâle, sans aliment ni eau, et laissées sans perturbation avec 50 graines au moins sert de fécondité standard. Elle augmente avec le temps de développement de l'uf à l'adulte de 73 ufs (32 j) à 94 ufs (44 j). La fécondité potentielle, obtenue avec des femelles alimentées sur sucre est de 125 ufs. Des femelles maintenues avec moins de 25 graines réduisent leur fécondité.Plus de la moitié des femelles distribuent leurs ufs uniformément jusqu'à une densité de 1,5 uf. L'uniformité peut provenir de l'induction de réactions de ponte par les graines portant moins d'ufs que l'effectif moyen. L'hyperdistribution au dessus des densités de 3 ufs/graine révèle l'aptitude à distinguer les graines avec 3 ufs des graines avec 4 ufs. Il y a moins d'écarts par rapport à l'uniformité (erreurs) avec des graines à surface rugueuse. Des femelles avec 10 graines font moins d'erreurs que des femelles avec 50 ou 100 graines.Les ufs sont ajoutés aux plus grosses graines sans ufs; ainsi, plus il y a de graines qui portent des ufs, plus le poids des graines avec ufs diminue. Un tel schéma est prévisible si la ponte est induite par des graines plus grosses que les graines rencontrées précédemment. Les femelles présentent un ordre de préférence clair quand elles sont en rapport avec leurs hôtes classiques: Vigna unguiculata > V. radiata > Cajanus cajan > Cicer arietinum Les caractères spécifiques de l'hôte, plutôt que la taille ou la texture de la surface servent à la distinction entre les graines d'un même hôte.Ces résultats expérimentaux peuvent être utilisés pour la modélisation du comportement de ponte d'une femelle.

     

Effects of temperature on life history variables in perch

Population structure, recruitment, mortality, life expectancy, age at maturity, gonad development, fecundity, net reproductive rate and energy storage were studied in a perch population exposed to thermal discharge. The experiment was conducted during 1978–1990 in an artificial enclosure at a nuclear power plant on the Baltic Sea coast. Abundance increased after the plant started operating, although the proportion of larger perch dropped to a low level. Recruitment improved, but adult mortality increased. This effect was accentuated as the fishes matured very early and at a very small size, reducing their ability to support a growing gonad in high winter temperatures. During the spawning period, condition dropped to very low levels accompanied by increased mortality. Surviving fish delayed the next spawning by one or more years and reduced their fecundity. Body condition thus improved in large perch at the end of the study period. Life-time fecundity was reduced, and reproductive performance shifted to youner ages.     

Genetic architecture of population differences in oviposition behaviour of the seed beetle Callosobruchus maculatus

Few studies have examined the genetic architecture of population differences in behaviour and its implications for population differentiation and adaptation. Even fewer have examined whether differences in genetic architecture depend on the environment in which organisms are reared or tested. We examined the genetic basis of differences in oviposition preference and egg dispersion between Asian (SI) and African (BF) populations of the seed beetle, Callosobruchus maculatus. We reared and tested females on each of two host legumes (cowpea and mung bean). The two populations differed in mean oviposition preference (BF females preferred cowpea seeds more strongly than did SI females) and egg dispersion (SI females distributed eggs more uniformly among seeds than did BF females). Observations of hybrid and backcross individuals indicated that only the population difference in oviposition preference could be explained by complete additivity, whereas substantial dominance and epistasis contributed to the differences in egg dispersion. Both rearing host and test host affected the relative magnitude of population differences in egg dispersion and the composite genetic effects. Our results thus demonstrate that the relative influence of epistasis and dominance on the behaviour of hybrids depends on the behaviour measured and that different aspects of insect oviposition are under different genetic control. In addition, the observed effect of rearing host and oviposition host on the relative importance of dominance and epistasis indicates that the genetic basis of population differences depends on the environment in which genes are expressed.     

Insect life history and the evolution of bacterial mutualism

Bacterial symbiosis has played a fundamental role in the evolution of eukaryotes. However, we still know little about how cooperative relationships with bacteria originate, and why they form in some host species but not others. Facultative symbionts that are beneficial, but not essential, provide unique insights into these processes. We use data from over a hundred aphid species to test if host life history is associated with the presence of facultative symbionts. We find that aphid species that have mutualistic associations with ants that protect them from natural enemies are less likely to carry symbionts that provide similar benefits. We also find one symbiont species occurs more frequently in unrelated aphid species that specialise on certain plant genera. In addition, aphid species that attack multiple plants often carry different symbiont complements. Our findings provide evidence of the ecological conditions that facilitate stable, mutually beneficial relationships between microbes and eukaryotic hosts.     

Habitat quality ranking depends on habitat-independent environmental factors: a model and results from Callosobruchus maculatus

1. Survival and reproduction in different habitats of a heterogeneous environment are affected by intrinsic habitat properties as well as by habitat-independent extrinsic factors. It is argued that the ranking of habitats according to quality may depend on extrinsic environmental factors that act with the same intensity in all habitats. This can happen if, across habitats, high reproductive success in the absence of extrinsic mortality is coupled with late reproduction.
2. In a model of a semelparous organism with overlapping generations that uses two habitats, the effect of different patterns of extrinsic mortality on habitat quality are analysed, as measured by the reproductive value of an egg, which estimates its expected contribution to the future gene pool.
3. Measurements of fitness components in Callosobruchus maculatus (F.) (Coleoptera: Bruchidae) on four host species demonstrate that, across hosts, long development can be associated with high reproductive success in the absence of extrinsic mortality. As a result, the quality ranking of three out of the four hosts would be different in a stable population regulated by intraspecific competition and egg parasitoids than in a growing population or a stable population regulated by age-independent larval mortality.
4. It is suggested that habitat-independent mortality may play a role in the evolution of ecological niches.