Florivory by the occupants of phytotelmata in flower parts can decrease host plant fecundity

Some types of plant accumulate liquid in their inflorescences creating phytotelmata. These environments protect the flowers against florivory, although they may be colonized by aquatic or semi-aquatic florivorous insect larvae, whose effects on the fitness of the plants remain unclear. We tested the hypothesis of floral antagonism by the occupants of phytotelmata, which predicts that florivory by the occupants of the phytotelmata represents a cost to the female fitness of the plant, reducing its fecundity. We manipulated experimentally the infestation by 3 florivores larvae species occupants of phytotelmata in inflorescences of Heliconia spathocircinata (Heliconiaceae) to test for negative direct trophic effects on the fecundity of the flowering and fruiting bracts. We found that the foraging of the hoverfly (Syrphidae) and moth (Lepidoptera) larvae in the inflorescences contributed to a decline in the fecundity of the plant. While the lepidopteran impacted fecundity when foraging in both flowering and fruiting bracts, the syrphid only affected the fruiting bracts, which indicates that the nectar and floral tissue are the principal resource exploited by the hoverfly. By contrast, soldier fly (Stratiomyidae) had a neutral effect on fecundity, while foraging in flowering or fruiting bracts. These findings corroborate our hypothesis, that herbivory by the larval occupants represents cost to the host plant having phytotelmata. The negative influence of this foraging on plant fecundity will nevertheless depend on the consequences of the exploitation of resources, which vary considerably in ephemeral habitats such as the phytotalmanta of flower parts.     

No Experimental Evidence for Host Ant Related Oviposition in a Parasitic Butterfly

The ability of adult butterflies of the genus Maculinea to locate their host ants prior to oviposition has been the subject of much discussion. We studied the egg laying behavior of the dusky large blue Maculinea nausithous whose larvae parasitize colonies of the ant Myrmica rubra. Flowerheads of the initial food plant were sprinkled with soil from ant nests, which contain chemicals involved in the nest recognition behavior of ants. The experiment was conducted to determine whether ant-released chemicals may act as oviposition cues and whether intraspecific competition for suitable plants may force female butterflies to alternative decisions. Host plant choice was not influenced by the presence of nest-derived host-ant cues. Density dependent shifts to less suitable host plants could not be ascertained nor changes in egg laying behavior across the flight period. The observed egg distribution could be primarily explained by host plant characteristics and environmental variability among sites. The result confirms the theory that host ant dependent oviposition appears to be a disadvantageous strategy in the face of resource limitation within ant colonies and the immobility of caterpillars.     

Importance of host feeding for parasitoids that attack honeydew-producing hosts

Insect parasitoids lay their eggs in arthropods. Some parasitoid species not only use their arthropod host for oviposition but also for feeding. Host feeding provides nutrients to the adult female parasitoid. However, in many species, host feeding destroys an opportunity to oviposit. For parasitoids that attack Homoptera, honeydew is a nutrient‐rich alternative that can be directly imbibed from the host anus without injuring the host. A recent study showed that feeding on host‐derived honeydew can be an advantageous alternative in terms of egg quantity and longevity. Here we explore the conditions under which destructive host feeding can provide an advantage over feeding on honeydew. For 5 days, Encarsia formosa Gahan (Hymenoptera: Aphelinidae) parasitoids were allowed daily up to 3 h to oviposit until host feeding was attempted. Host feedings were either prevented or allowed and parasitoids had ad libitum access to honeydew between foraging bouts. Even in the presence of honeydew, parasitoids allowed to host feed laid more eggs per hour of foraging per host‐feeding attempt than parasitoids that were prevented from host feeding. The higher egg‐laying rate was not compromised by survival or by change in egg volume over time. In conclusion, host feeding can provide an advantage over feeding on honeydew. This applies most likely under conditions of high host density or low extrinsic mortality of adult parasitoids, when alternative food sources cannot supply enough nutrients to prevent egg limitation. We discuss how to integrate ecological and physiological studies on host‐feeding behavior     

拟寄生蜂搜索产卵过程中对寄主的竞争

综述拟寄生蜂搜索产卵过程中对寄主竞争的最新研究进展.这类竞争具有四种方式,即标记寄主、杀卵和杀幼、守护寄主和捕食寄主.（1）标记寄主常涉及寄主标记信息素,这是由雌蜂在产卵前、产卵时或产卵后分泌的化学物质.寄主标记信息素常介导拟寄生蜂对已寄生和健康寄主的辨别、减少过寄生和多寄生、减轻种内和种间竞争压力.（2）雌蜂遇到已寄生寄主时,很多种类杀死前一雌蜂遗留的卵和幼虫,再产下自己的卵.雌蜂使用三种方法杀卵和杀幼,即产卵器穿刺、取食和使用有毒物质.通过杀卵和杀幼,产卵雌蜂清除了前一雌蜂遗留的后代,主动改善了寄主品质,从而有利于自身后代的生存.（3）守护寄主在肿腿蜂科、缘腹细蜂科、金小蜂科、缨小蜂科和茧蜂科中均有报道,守护者驱逐入侵者以保护后代及健康寄主.（4）捕食寄主不仅减少了健康寄主数量,且直接导致已寄生寄主中拟寄生蜂卵和幼虫的死亡.雌蜂一般在体内成熟卵量较少时捕食寄主.讨论了研究拟寄生蜂搜索产卵过程中竞争寄主的理论意义和实际应用价值.     

Host-size mediated trade-off in a parasitoid Sclerodermus harmandi

Optimality models of host-parasitoid relationships have traditionally assumed that host quality increases as a function of host size at parasitism. However, trade-offs may play a crucial role in species evolution and should be found in host-parasitoid interactions where the host quality may differ between different sizes. Here, we investigated the effects of host size differences of Monochamus alternatus larva on foraging decisions, parasitism and related fitness in a gregarious ectoparasitoid, Sclerodermus harmandi. Two-choice and non-choice experiments were conducted with M. alternatus larvae to evaluate preference and performance of S. harmandi, respectively. Results from two-choice test showed that adult females prefer to attack large larvae rather than small larvae. In no-choice tests, adult females needed more time to paralyze large larvae than small larvae before laying eggs on the body surface of M. alternatus larvae and had lower survival and parasitism rate on those large larvae. Foraging decisions of S. harmandi led to the selection of the most profitable host size for parasitoid development, which showed more offspring gained on large M. alternatus larvae than on small larvae and got higher body weight on those large hosts. This study demonstrates the existence of trade-off occurring during host-parasitoids interactions according to host size related quality.     

Temporal patterns of host availability, brown-headed cowbird brood parasitism, and parasite egg mass

I studied relationships between temporal patterns of host availability, brood parasitism, and egg mass for the parasitic brown-headed cowbird (Molothrus ater). At a study site consisting largely of edge habitat in north-eastern Illinois, I found 834 bird nests from 27 species. A total of 407 cowbird eggs and nestlings were found in these nests over three laying seasons. Nearly all (n= 379; 93%) were found in the nests of seven host species. For these species and all taken together, weekly nest availability generally decreased whereas parasitism frequency generally increased throughout the cowbird laying season, but the proportions of nests parasitized and the mean number of cowbird eggs in them did not. Additionally, no correlation was found between the proportions of nests parasitized and nest availability. Cowbird egg mass generally increased throughout the laying season, indicating that foraging conditions improved and that, early in the laying season, egg mass and quality may be less important than quantity. Consistently high weekly levels of parasitism indicate that cowbird reproduction was less limited by resources needed for egg production and more by the availability of suitable host nests. Fluctuating weekly host availabilities suggest that previously established, constant rates of cowbird egg laying would produce an excess of eggs during periods of low host availability. Further, the low frequency of parasitism (1%) of nests in stages too advanced for successful parasitism, and of abandoned nests, is consistent with the hypothesis that cowbirds' consistently high rate of egg production helps assure an egg is available when an appropriate nest is found. Frequently, nests were parasitized multiple times, raising the possibility that cowbirds were interfering with their own reproduction. A diverse host community increases the possibility that a decline of any one host species is unlikely to affect cowbird reproduction significantly. Received 11 July 1997 / Accepted: 31 March 1998     

The role of larval aggression and mobility in the transition between solitary and gregarious development in parasitoid wasps

Models explaining the appearance of gregariousness in insect parasitoids assume the presence of a tolerance gene that enables nonsiblicidal behaviour in gregarious larvae. Under this assumption, nontolerant individuals should attack and kill tolerant individuals when sharing a host, making this behaviour susceptible to invasion by nontolerant individuals. We propose an alternative hypothesis where gregarious larvae retain their aggressiveness but are less mobile. We tested this hypothesis with two sympatric and congeneric species of Mymaridae, Anaphes victus and Anaphes listronoti , respectively, solitary and gregarious egg parasitoids of a Curculionidae, Listronotus oregonensis . Results obtained in competition experiments and from direct observation of movement in mymariform larvae of both species support the reduced mobility hypothesis. By being immobile while retaining their fighting capacity, A. listronoti mymariform larvae appear to optimize host utilization through gregarious development, but can still defend themselves against sympatric aggressive species.     

Egg laying rather than host quality or host feeding experience drives habitat estimation in the parasitic wasp Nasonia vitripennis

In variable environments, sampling information on habitat quality is essential for making adaptive foraging decisions. In insect parasitoids, females foraging for hosts have repeatedly been shown to employ behavioral strategies that are in line with predictions from optimal foraging models. Yet, which cues exactly are employed to sample information on habitat quality has rarely been investigated. Using the gregarious parasitoid Nasonia vitripennis (Walker; Hymenoptera: Pteromalidae), we provided females with different cues about hosts to elucidate, which of them would change a wasp's posterior behavior suggesting a change in information status. We employed posterior clutch size decisions on a host as proxy for a female's estimation of habitat quality. Taking into account changes in physiological state of the foraging parasitoid, we tested whether different host qualities encountered previously change the subsequent clutch size decision in females. Additionally, we investigated whether other kinds of positive experiences—such as ample time to investigate hosts, host feeding, or egg laying—would increase a wasp's estimated value of habitat quality. Contrary to our expectations, quality differences in previously encountered hosts did not affect clutch size decisions. However, we found that prior egg laying experience changes posterior egg allocation to a host, indicating a change in female information status. Host feeding and the time available for host inspection, though correlated with egg laying experience, did not seem to contribute to this change in information status.     

Innate preference hierarchies coupled with adult experience,rather than larval imprinting or transgenerational acclimation,determine host plant use in Pieris rapae

The evolution of host range drives diversification in phytophagous insects, and understanding the female oviposition choices is pivotal for understanding host specialization. One controversial mechanism for female host choice is Hopkins’ host selection principle, where females are predicted to increase their preference for the host species they were feeding upon as larvae. A recent hypothesis posits that such larval imprinting is especially adaptive in combination with anticipatory transgenerational acclimation, so that females both allocate and adapt their offspring to their future host. We study the butterfly Pieris rapae, for which previous evidence suggests that females prefer to oviposit on host individuals of similar nitrogen content as the plant they were feeding upon as larvae, and where the offspring show higher performance on the mother's host type. We test the hypothesis that larval experience and anticipatory transgenerational effects influence female host plant acceptance (no‐choice) and preference (choice) of two host plant species (Barbarea vulgaris and Berteroa incana) of varying nitrogen content. We then test the offspring performance on these hosts. We found no evidence of larval imprinting affecting female decision‐making during oviposition, but that an adult female experience of egg laying in no‐choice trials on the less‐preferred host Be. incana slightly increased the P. rapae propensity to oviposit on Be. incana in subsequent choice trials. We found no transgenerational effects on female host acceptance or preference, but negative transgenerational effects on larval performance, because the offspring of P. rapae females that had developed on Be. incana as larvae grew slower on both hosts, and especially on Be. incana. Our results suggest that among host species, preferences are guided by hard‐wired preference hierarchies linked to species‐specific host traits and less affected by larval experience or transgenerational effects, which may be more important for females evaluating different host individuals of the same species.     

The protective role of endogenous bacterial communities in chironomid egg masses and larvae

Insects of the family Chironomidae, also known as chironomids, are distributed worldwide in a variety of water habitats. These insects display a wide range of tolerance toward metals and organic pollutions. Bacterial species known for their ability to degrade toxicants were identified from chironomid egg masses, leading to the hypothesis that bacteria may contribute to the survival of chironomids in polluted environments. To gain a better understanding of the bacterial communities that inhabit chironomids, the endogenous bacteria of egg masses and larvae were studied by 454-pyrosequencing. The microbial community of the egg masses was distinct from that of the larval stage, most likely due to the presence of one dominant bacterial Firmicutes taxon, which consisted of 28% of the total sequence reads from the larvae. This taxon may be an insect symbiont. The bacterial communities of both the egg masses and the larvae were found to include operational taxonomic units, which were closely related to species known as toxicant degraders. Furthermore, various bacterial species with the ability to detoxify metals were isolated from egg masses and larvae. Koch-like postulates were applied to demonstrate that chironomid endogenous bacterial species protect the insect from toxic heavy metals. We conclude that chironomids, which are considered pollution tolerant, are inhabited by stable endogenous bacterial communities that have a role in protecting their hosts from toxicants. This phenomenon, in which bacteria enable the continued existence of their host in hostile environments, may not be restricted only to chironomids.     

Cyclic Behavior of <Emphasis Type="Italic">Lycorma delicatula</Emphasis> (Insecta: Hemiptera: Fulgoridae) on Host Plants

Lycorma delicatula is an invasive insect species in Korea, and its populations are increasing. It sucks plant sap and can cause withering of the whole trees. To prevent damage to fruit trees, it is important to determine the behavioral characteristics of L. delicatula. We monitored migration patterns and host plant preferences, from the egg mass on the tree bark, through egg hatching to egg laying. Most eggs hatched between 0500 h and 0800 h. The nymphs ascended trees until they reached the leaves. They often fell to the ground because of physical factors, such as wind, but then ascended the trees again on the upper sides of the branches. Their falling-ascending cycle became longer as their arolia developed. The host preference changed over the course of its growth, from a broad range of plant species at the early nymph stage to a few plant species, such as Ailanthus altissima at the adult stage. We summarize the life cycle behavior as follows. Initially, there is a short-term cycle of falling and ascending, which becomes longer as the nymphs age. The other cycle is a yearly cycle of host plant preferences, with broad host preference in the early nymph stage and a narrower host preference in the adult stage. Knowledge of this cyclic behavior can be used to prevent the rapid expansion of L. delicatula populations in orchards.     

Responses of ovipositing moths to host plant deprivation: life history aspects and implications for intercropping

1. When considering intercropping as a strategy to reduce pest oviposition, knowledge about the insect’s oviposition behaviour is very important. Physiological effects on the insect because of difficulties in finding a suitable oviposition site may also be important. 2. In the present study, the effects that delays in access to host plants have on lifetime fecundity on diamondback moth and leek moth were examined. The ability to postpone egg laying, fecundity and lifetime oviposition are discussed in relation to intercrop/cover crop as a strategy to reduce oviposition on crop plants. 3. When faced with host plant deprivation, the diamondback moth is relatively more dependent upon host plant stimuli for the onset of egg production. By contrast, leek moth is able to postpone egg production for a longer time. There even appeared to be a tendency for leek moth females to extend their lifetime when faced with host plant deprivation. 4. We conclude that leek moths have the ability to postpone production of eggs and lay them later in life when finally encountering host plants after a period of host plant deprivation. Therefore, the use of intercropping as a strategy to reduce oviposition is questionable. For such an insect, use of a trap crop might be a better option because the female will lay her eggs in the trap crop and not get the opportunity to lay them later in life when finally encountering crop plants.     

Does mother know best? The preference–performance hypothesis and parent–offspring conflict in aboveground–belowground herbivore life cycles

1. A substantial amount of research on host‐plant selection by insect herbivores is focused around the preference–performance hypothesis (PPH). To date, the majority of studies have primarily considered insects with aboveground life cycles, overlooking insect herbivores that have both aboveground and belowground life stages, for which the PPH could be equally applicable. 2. This study investigated the factors influencing the performance of the root‐feeding vine weevil (Otiorhynchus sulcatus) larvae and whether this was linked to the oviposition behaviour of the maternal adult living aboveground. 3. Maternal insects feeding aboveground reduced root biomass by 34% and increased root carbon by 4%. Larvae feeding on plants subjected to aboveground herbivory had reduced mass. Irrespective of the presence of maternal herbivory, larval mass was positively correlated with root biomass. 4. Larval mass was also reduced by conspecific larvae, previously feeding on roots (19% reduction). However, the mechanism underpinning this effect remains unclear, as in contrast to maternal herbivory aboveground, prior larval feeding did not significantly affect root biomass or root carbon concentrations. 5. Maternal insects did not distinguish between plants infested with larvae and those that were free of larvae, in terms of their egg‐laying behaviour. Conversely, maternal insects tended to lay eggs on plants with smaller root systems, a behaviour that is likely to negatively affect offspring performance. 6. The PPH is not supported by our findings for the polyphagous vine weevil feeding on the host plant raspberry (Rubus idaeus), and in fact our results suggest that there is the potential for strong parent–offspring conflict in this system.     

The evolution of larval foraging behaviour in response to host plant variation in a leaf beetle

The evolutionary causes of variation in host specialization among phytophagous insects are still not well understood and identifying them is a central task in insect–host plant biology. Here we examine host utilization of the chrysomelid beetle Oreina elongata that shows interpopulation variation in the degree of specialization. We focus on larval behaviour and on what selection pressures may favour the use of two different larval host plants ( Adenostyles alliariae and Cirsium spinosissimum ) in one population as opposed to specialization onto one of them as is seen in other populations. The results suggest that the degree of exploratory foraging behaviour is higher in larvae from the two-host population than in single host populations, and a field survey of the two-host population also indicated that larvae do move between host species. A field experiment indicated that predation rates on O. elongata larvae in the two-host population are higher on one of the host species, A. alliariae , than on the alternative C. spinosissimum . In combination with earlier results this finding suggest that larvae move between hosts to obtain better food on one host, and to get better protection from predators on the other. It appears that in this two-host situation a single plant species does not provide the most beneficial conditions in all parts of O. elongata life cycle and individuals may obtain different plant-specific benefits by moving between host species. This heterogeneous host situation appears to have selected for the explorative larval foraging strategy seen in the in the two-host population. In general, the results support the notion that to understand patterns of host plant use in insects it is often vital to consider a range of host related selection pressures whose relative importance may vary between life stages of the insect.     

Larval competition within seeds: From the behaviour process to the ecological outcome in the seed beetle Callosobruchus maculatus

Abstract Environmental conditions experienced by organisms during development can have profound impacts on adult fitness and behaviour. Internally feeding larvae unable to leave the seed selected by their mother face limitations of resource suitability and competition. The host seed may guide the larval behaviour within the seed leading to differential intensity of competition and determining its process and outcome, which varies in strains of the legume seed beetle Callosobruchus maculatus (Coleoptera: Bruchidae). However, the intensity, process and outcome of larval competition in different hosts have yet to be simultaneously considered, the objective of the present study. Here we assessed the intensity, process and outcome of intrastrain larval competition as related to host type, and how they are interrelated. Larval competition was faced with two distinct strategies – scramble and contest competition depending on the insect strain and host seed species. The intensity of competition did not show any straight link with the process and outcome of competition. Only a single strain showed a contest competition process with likely interference between larvae, while the four other strains studied showed the process of scramble competition. The process of scramble competition, however, led to variable outcomes in mung beans based on larval competition curves. Such differences were not apparent on cowpea seeds and either the plateau or the peak expected on the larval fitness curves were not reached preventing the distinction of the competition outcome, a likely consequence of the egg laying behaviour of these strains limiting the maximum number of eggs laid per seed. Seed host species rather than seed size are the likely cause of the differences observed from the initial expectation. The strain showing the process of contest competition increased larval fitness with density of larvae emerged per seed regardless of the host species, an unexpected outcome based on theoretical models. In this case the egg laying behaviour of the adult female is probably the main fitness determinant of its progeny.     

Differential attraction of parasitoids in relation to specificity of kairomones from herbivores and their by‐products

Infochemicals are used by foraging parasitoids in the host selection process from habitat preference until host recognition. Kairomones from the herbivore host plays a vital role in the attraction of parasitoids, particularly in the micro‐habitat. Parasitoids are specifically attracted to their respective herbivore species even when different herbivores are present on the same plant. Chemicals emitted from different stages of host (eggs, larvae, pupae, adult), host by‐products (e.g., frass, exuviae, mandibular gland secretions, defense secretions etc.), or intra‐specific infochemicals (pheromones) can be main signals for the parasitoids. Parasitoids can differentiate between host and non‐host, between different hosts and host stages by perceiving specific volatile and contact kairomones from the host itself, host along with its by‐product, by‐products alone or intra‐specific infochemicals; of which frass (by‐product) and intra‐specific infochemicals are the most reported ones. Adult and larval parasitoids have been reported to be attracted to kairomones of their target stage or byproduct of their host. Pupal parasitoids have been found to utilize kairomones from the preceding host stage while egg parasitoids are known to exploit a variety of host infochemicals, for example, either from eggs themselves or other non‐target host stages, especially adults and adult‐related by‐products. The kairomonal chemicals identified so far include various groups, but mainly hydrocarbons. A high degree of host specificity and host acceptance is important for the parasitoids as any mistake may result in the loss of fitness.     

Does ant predation favour leaf beetle specialization on toxic host plants?

Generalist predators are frequently seen as evolutionary forces that narrow the host range in herbivorous insects. Predators may favour specialization of herbivores on host plants containing toxic chemicals (which can be used by herbivores for their own defence) if host plant‐derived defences provide better protection from enemies than do autogenously produced defences. We compared the effectiveness of these two defensive strategies in the larvae of six species of leaf beetle (Chrysomelidae) against wood ants (Formica rufa group) in field experiments. Ants were more strongly repelled by larvae with host plant‐derived, salicylaldehyde‐containing secretions than by larvae with various autogenous secretions, but collectively foraging ants ultimately overcame any type of chemical defence by social interactions, chemical signalling, and olfactory learning. As a result, ants killed all larvae of Chrysomela lapponica defended by salicylaldehyde‐containing secretions within 2 days of their introduction to willows within 15 m of ant nests. We conclude that in the field neither type of chemical defence provides complete protection against wood ants in the vicinity of their nests, and that evolutionary shifts from autogenous production of secretion to sequestration of plant allelochemicals in leaf beetles may be favoured mostly at low ant densities on the periphery of ant foraging areas.     

Ecophysiological attributes of adult overwintering in insects: insights from a field study of the nut weevil,Curculio nucum

Abstract Diapausing insect species have evolved a great diversity of life cycles, although overwintering occurs at a single development stage within most species. Understanding why diapause has evolved towards a given life stage requires investigation of both the ecological and physiological attributes. Notably, it is suggested that adult overwintering is more energy‐demanding than larval overwintering but it brings fitness gains by allowing adults to be synchronized with their seasonal requisites through an early spring emergence. This hypothesis is tested in field conditions in the nut weevil Curculio nucum, whose life cycle comprises an obligate 2‐year, nonfeeding underground phase, including a larval, followed by an adult, overwintering. In this species, adult wintering leads to an early spring emergence; at first glance, however, this does not enhance synchronization between weevils and their host because adults emerge more than 1 month before starting to breed. It is suggested that adult overwintering ultimately evolved in response to the phenology of the host, by allowing females to oviposit in nuts before their full sclerotinization. Adult overwintering appears to be costly because adults postpone reproduction for 1 year, incur a significant weight loss and require feeding before egg laying. Surprisingly, lipids are unaffected during diapause, lipogenesis even being likely in the summer metamorphosis. These results suggest that the lipids involved in egg production may entirely come from the larval stages, whereas the other nutrients are acquired through adult feeding but this remains to be tested.     

The evolution of egg clustering in butterflies: A test of the egg desiccation hypothesis

Females of many insect species cluster their eggs. Egg clustering by lepidopteran species usually results in aggregation of larvae that are more often conspicuously coloured and apparently distasteful or unpalatable than larvae of solitary species. While the costs and benefits of aggregation in terms of larval survival and growth are well documented, the evolutionary ecology of egg clustering has been long debated and is still unresolved. We tested the egg desiccation hypothesis, first proposed by Stamp (1980), which to our knowledge has never been examined experimentally. The egg desiccation hypothesis proposes that egg clustering is adaptive per se (i.e. increases fitness of females) by reducing egg mortality via desiccation.We tested this hypothesis for the Nymphalid butterfly, Chlosyne lacinia, an egg-clustering species on its sunflower host plant, Helianthus annuus. We first documented natural variation in batch size for this butterfly. We then tested experimentally hatch success of varying batch sizes and egg-layering arrangements under controlled humidity levels. Hatch success was positively related to relative humidity. Eggs in larger groups with greater number of layers had greater hatch success than smaller, monolayered egg batches, especially when relative humidity was low. Our results indicate that, not only number of eggs, but also the arrangement (i.e. layering and density), increase batch survival by protecting eggs from desiccation. However, despite increased hatch success in dense, multilayered clusters, we found wide variation in layering and density in natural populations of C. lacinia. This variation is probably maintained by trade-offs in egg survival, such as increased cannibalism of eggs by siblings, in dense clusters. Nevertheless, protection from egg desiccation provides an alternative explanation for the origin and maintenance of egg clustering in lepidopterans and possibly other insects. The pattern of egg deposition in the Nymphalidae supports this hypothesis, since most North American species cluster their eggs tightly, whereas most species in tropical regions lay eggs singly or in loose monolayers.     

SHIFT IN EGG‐LAYING STRATEGY TO AVOID PLANT DEFENSE LEADS TO REPRODUCTIVE ISOLATION IN MUTUALISTIC AND CHEATING YUCCA MOTHS

Through the process of ecological speciation, insect populations that adapt to new host plant species or to different plant tissues could speciate if such adaptations cause reproductive isolation. One of the key issues in this process is identifying the mechanisms by which adaptation in ecological traits leads directly to reproductive isolation. Here I show that within a radiation of specialist moths that pollinate and feed on yuccas, shifts in egg placement resulted in changes in female moth egg‐laying structures that led to concomitant changes in male reproductive morphology. As pollinator moths evolved to circumvent the ability of yuccas to selectively abscise flowers that contain pollinator eggs, ovipositor length became shorter. Because mating occurs through the ovipositor, shortening of the ovipositor also led to significantly shorter and wider male intromittent organs. In instances where two pollinator moth species occur in sympatry and on the same host plant species, there is one short and one long ovipositor species that are reproductively isolated. Given that many plant‐feeding insects lay eggs into plant tissues, changes in ovipositor morphology that lead to correlated changes in reproductive morphology may be a mechanism that maintains reproductive isolation among closely related species using the same host plant species.