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.     

No effect of larval experience on adult host preferences in Polygonia c‐album (Lepidoptera: Nymphalidae): on the persistence of Hopkins’ host selection principle

Abstract 1. The possible effect of juvenile imprinting or ‘chemical legacy’ on the subsequent oviposition – often called the ‘Hopkins’ host selection principle’– has been a controversial but recurrent theme in the literature on host‐plant preference. While it appears possible in principle, experimental support for the hypothesis is equivocal. The present study points out that it is also important to consider its theoretical implications, and asks under what circumstances, if any, it should be favoured by natural selection. 2. Following this reasoning, it is predicted that host preference in the polyphagous butterfly Polygonia c‐album L. (Lepidoptera, Nymphalidae) should not be influenced by larval environment. This was tested by rearing larvae on three natural host plants: the high‐ranked Urtica dioica and the medium‐ranked Salix cinerea and Ribes uva‐crispa, and exposing the naive females to oviposition choices involving the same set of plants. 3. It was found that larval host plant had no effect on oviposition decisions of the adult female. Hence, the Hopkins’ host selection principle does not seem to be applicable in this species. 4. Based on recent insights on how accuracy of environmental versus genetic information should affect the control of developmental switches, the conditions that could favour the use of juvenile cues in oviposition decisions are discussed. Although the Hopkins’ host selection hypothesis cannot be completely ruled out, we argue that the circumstances required for it to be adaptive are so specific that it should not be invoked as a general hypothesis for host selection in plant‐feeding insects.     

Host feeding experience affects host plant odour preference of the polyphagous leafminer Liriomyza bryoniae

Herbivorous insects use highly specific volatiles or blends of volatiles characteristic to particular plant species to locate their host plants. Thus, data on olfactory preferences can be valuable in developing integrated pest management tools that deal with manipulation of pest insect behaviour. We examined host plant odour preferences of the tomato leafminer, Liriomyza bryoniae (Kaltenbach) (Diptera: Agromyzidae), which is an economically important agricultural pest widespread throughout Europe. The odour preferences of leafminers were tested in dependence of feeding experiences. We ranked host plant odours by their appeal to L. bryoniae based on two‐choice tests using a Y‐tube olfactometer with five host plants: tomato, Solanum lycopersicum Mill.; bittersweet, Solanum dulcamara L.; downy ground‐cherry, Physalis pubescens L. (all Solanaceae); white goosefoot, Chenopodium album L. (Chenopodiaceae); and dead nettle, Lamium album L. (Lamiaceae). The results imply that ranking of host plant odours by their attractiveness to L. bryoniae is complicated due to the influence of larval and adult feeding experiences. Without any feeding experience as an adult, L. bryoniae males showed a preference for the airflow with host plant odour vs. pure air, whereas females did not display a preference. Further tests revealed that adult feeding experience can alter the odour choice of L. bryoniae females. After feeding experience, females showed a preference for host plant odour vs. pure air. Feeding experience in the larval stage influenced the choice by adults of both sexes: for males as well as females reared on bittersweet the odour of that plant was the most attractive. Thus, host feeding experience both in larval and/or adult stage of polyphagous tomato leafminer L. bryoniae influences host plant odour preference by adults.     

‘Do you remember the first time?’ Host plant preference in a moth is modulated by experiences during larval feeding and adult mating

In insects, like in other animals, experience‐based modulation of preference, a form of phenotypic plasticity, is common in heterogeneous environments. However, the role of multiple fitness‐relevant experiences on insect preference remains largely unexplored. For the multivoltine polyphagous moth Spodoptera littoralis we investigated effects of larval and adult experiences on subsequent reproductive behaviours. We demonstrate, for the first time in male and female insects, that mating experience on a plant modulates plant preference in subsequent reproductive behaviours, whereas exposure to the plant alone or plant together with sex pheromone does not affect this preference. When including larval feeding experiences, we found that both larval rearing and adult mating experiences modulate host plant preference. These findings represent the first evidence that host plant preferences in polyphagous insects are determined by a combination of innate preferences modulated by sensory feedback triggered by multiple rewarding experiences throughout their lifetime.     

Development of a positive preference–performance relationship in an oligophagous beetle: adaptive learning?

The relationship between oviposition preference and larval performance is a central topic in insect–plant biology. In this study, we investigate whether the oligophagous flea beetle, Altica fragariae Nakane (Coleoptera: Chrysomelidae), exhibits a positive preference–performance relationship, and whether oviposition preference develops over time. We tested the beetles using four sympatric plant species: Duchesnea indica (Andrews) Focke (the normal host plant), Agrimonia pilosa Ledeb. (a secondary host plant), and Potentilla chinensis Ser. and Sanguisorba officinalis L. (host plants of two related Altica species) (all Rosaceae). In no‐choice experiments, both oviposition rate and offspring fitness parameters (eclosion rate, development time, and body mass) were highest on D. indica. Oviposition rate was much lower on P. chinensis than on A. pilosa, whereas offspring fitness parameters did not differ significantly between these two host plants. Offspring fitness were lowest for S. officinalis, and adult females refused to oviposit on this acceptable non‐host in a no‐choice situation. Repeated two‐choice experiments showed that the proportion of oviposition on one of the novel host plants decreased significantly over time when the alternative host plant was D. indica. In repeated two‐choice experiments using A. pilosa and P. chinensis, females mainly fed on A. pilosa but distributed their eggs equally over the two host plants, in accordance with the lack of difference in offspring fitness on those hosts. Together, these results showed that A. fragariae females develop a positive preference–performance relationship over time. We suggest that A. fragariae achieves this through adaptive learning of oviposition preference: not only does the female learn to discriminate among the host plants when there is a fitness difference for her offspring, but the female also fails to discriminate when there is no fitness difference.     

Beyond preference and performance: host plant selection by monarch butterflies,Danaus plexippus

The connection between adult preferences and offspring performance is a long‐standing issue in understanding the evolutionary and ecological forces that dictate host associations and specialization in herbivorous insects. Indeed, decisions made by females about where to lay their eggs have direct consequences for fitness and are influenced by interacting factors including offspring performance, defence and competition. Nonetheless, in addition to these attributes of the offspring, a female's choices may be affected by her own prior experience. Here we examined oviposition preference, larval performance and the role of learning in the monarch butterfly, Danaus plexippus, which encounters diverse milkweed host species across its broad range and over the course of migration. Monarch females consistently preferred to oviposit on Asclepias incarnata subspecies pulchra. This plant, however, was associated with poor caterpillar growth, low sequestration of toxins and the highest plant defences (latex and trichomes). We examined flexibility in this apparently maladaptive preference by testing the impact of previous experience and competition on preference. Experience laying on an alternative plant species enhanced preference for that species in contrast to A. i. pulchra. In addition, presence of a (competing) conspecific caterpillar on A. i. pulchra had a strongly deterrent effect and reversed host plant preferences. Thus, monarch butterflies exhibit preferences contrary to what would be expected based on offspring development and sequestered defences, but their preferences are altered by learning and competition, which may allow butterflies to shift preferences as they encounter diverse milkweeds across the landscape. Learning and perception of threats (i.e. competition or predation) may be critical for most herbivorous insects, which universally experience heterogeneity among their potential host plants.     

Host plant utilization in the comma butterfly: sources of variation and evolutionary implications

A major challenge in the study of insect-host plant interactions is to understand how the different aspects of offspring performance interact to produce a preference hierarchy in the ovipositing females. In this paper we investigate host plant preference of the polyphagous butterfly Polygonia c-album (Lepidoptera: Nymphalidae) and compare it with several aspects of the life history of its offspring (growth rate, development time, adult size, survival and female fecundity). Females and offspring were tested on four naturally used host plants (Urtica dioica, Ulmus glabra, Salix caprea, and Betula pubescens). There was substantial individual variation in host plant preference, including reversals in rank order, but the differences were largely confined to differences in the ranking of Urtica dioica and S. caprea. Different aspects of performance on these two plants gave conflicting and complementary results, implying a trade-off between short development time on U. dioica, and larger size and higher fecundity on S. caprea. As all performance components showed low individual variation the large variation in host plant preference was interpreted as due to alternative oviposition strategies on the basis of similar performance hierarchies. This indicates that the larval performance component of host-plant utilization may be more conservative to evolutionary change than the preference of ovipositing females. Possible macro-evolutionary implications of this are discussed.     

Relationship between adult and larval host plant selection and larval performance in the generalist moth, Trichoplusia ni

Adult oviposition preferences are expected to correlate with host plant suitability for the development of their offspring. For most lepidopteran species, this is particularly important as the hatching neonate larvae of many species are relatively immobile. Thus, the site of oviposition chosen by a female adult can greatly influence the probability of survival for her offspring. In the present study, we investigated the oviposition preference of adult Trichoplusia ni moths for six plant species to determine whether they could accurately rank the suitability of the plants for larval development. We also compared oviposition preferences to neonate larval acceptance and preference to determine whether the adult host range matched that of larval diet breath. Our results indicate that in two-choice and no-choice tests adult T. ni were able to rank the plants accurately, with the exception of anise hyssop. However, when given a choice of all six plants together, they laid more eggs on a plant that was not suitable for larval survival. Larvae accepted and fed on all plants in no-choice tests, and accurately ranked them according to larval performance. We conclude that neonate larvae are better able than adults to rank plants according to larval performance, and that larval diet breadth is wider than the range of plants accepted by adults. We also provide a discussion of the reduced accuracy of adult oviposition preference with increased plant choices.     

Butterfly oviposition preference is not related to larval performance on a polyploid herb

The preference–performance hypothesis predicts that female insects maximize their fitness by utilizing host plants which are associated with high larval performance. Still, studies with several insect species have failed to find a positive correlation between oviposition preference and larval performance. In the present study, we experimentally investigated the relationship between oviposition preferences and larval performance in the butterfly Anthocharis cardamines. Preferences were assessed using both cage experiments and field data on the proportion of host plant individuals utilized in natural populations. Larval performance was experimentally investigated using larvae descending from 419 oviposition events by 21 females on plants from 51 populations of two ploidy types of the perennial herb Cardamine pratensis. Neither ploidy type nor population identity influenced egg survival or larval development, but increased plant inflorescence size resulted in a larger final larval size. There was no correlation between female oviposition preference and egg survival or larval development under controlled conditions. Moreover, variation in larval performance among populations under controlled conditions was not correlated with the proportion of host plants utilized in the field. Lastly, first instar larvae added to plants rejected for oviposition by butterfly females during the preference experiment performed equally well as larvae growing on plants chosen for oviposition. The lack of a correlation between larval performance and oviposition preference for A. cardamines under both experimental and natural settings suggests that female host choice does not maximize the fitness of the individual offspring.     

Oviposition acceptance and larval development of Chilo partellus stemborers in drought‐stressed wild and cultivated grasses of East Africa

Maternal host choices during oviposition by herbivorous insects determine the fitness of their offspring and may be influenced by environmental changes that can alter host‐plant quality. This is of particular relevance to ‘push‐pull’ cropping systems where host preferences are exploited to manage insect pest populations. We tested how drought stress in maize and companion plants that are used in these systems affect oviposition preference, larval feeding, and development of the spotted stemborer, Chilo partellus Swinhoe (Lepidoptera: Crambidae). Five host species were tested (all Poaceae): maize (Zea mays L.), Napier grass (Pennisetum purpureum Schumach), signal grass [Brachiaria brizantha (A. Rich) Stapf], Brachiaria cv. ‘Mulato’, and molasses grass [Melinis minutiflora (Beauv.)]. Under drought stress, maize experienced as much oviposition as control unstressed maize in choice and no‐choice experiments. Similarly, larval leaf damage was not significantly different in drought‐stressed vs. unstressed maize. In contrast, oviposition occurred less on drought‐stressed than on unstressed Napier and signal grass. Oviposition acceptance and leaf damage remained low in both drought‐stressed and unstressed molasses grass and Mulato. Larval survival and development remained high in drought‐stressed maize, but not in Napier, signal, and molasses grass and Mulato, where survival and development were low in both drought‐stressed and unstressed plants. Our results indicate that herbivore responses to drought‐stressed plants depend on the plant species and that drought stress can change host preference and acceptance rankings. In particular, trap‐crops such as Napier grass may not divert oviposition from the main maize crop under drought stress conditions.     

Host plant choice in the comma butterfly-larval choosiness may ameliorate effects of indiscriminate oviposition

In most phytophagous insects, the larval diet strongly affects future fitness and in species that do not feed on plant parts as adults, larval diet is the main source of nitrogen. In many of these insect-host plant systems, the immature larvae are considered to be fully dependent on the choice of the mothers, who, in turn, possess a highly developed host recognition system. This circumstance allows for a potential mother-offspring conflict, resulting in the female maximizing her fecundity at the expense of larval performance on suboptimal hosts. In two experiments, we aimed to investigate this relationship in the polyphagous comma butterfly, Polygonia c-album, by comparing the relative acceptance of low- and medium-ranked hosts between females and neonate larvae both within individuals between life stages, and between mothers and their offspring. The study shows a variation between females in oviposition acceptance of low-ranked hosts, and that the degree of acceptance in the mothers correlates with the probability of acceptance of the same host in the larvae. We also found a negative relationship between stages within individuals as there was a higher acceptance of lower ranked hosts in females who had abandoned said host as a larva. Notably, however, neonate larvae of the comma butterfly did not unconditionally accept to feed from the least favorable host species even when it was the only food source. Our results suggest the possibility that the disadvantages associated with a generalist oviposition strategy can be decreased by larval participation in host plant choice.     

Inconsistent use of host plants by the Alaskan swallowtail butterfly: adult preference experiments suggest labile oviposition strategy

Abstract 1. The Alaskan swallowtail butterfly (Papilio machaon aliaska) uses three unrelated plant species as hosts: Cnidium cnidiifolium (Apiaceae), Artemisia arctica (Asteraceae), and Petasites frigidus (Asteraceae). The research presented here investigated whether there are any consistent patterns in host choice by P. m. aliaska females. 2. The first two experiments were designed to test if P. m. aliaska host preference is constant or if it changes from day to day. If host preference is labile, the experiments were designed to also test whether a female’s diet breadth narrows or expands over time. 3. The third experiment tested the host preferences of female offspring from several wild-caught P. m. aliaska females. If P. m. aliaska individuals are specialised in their host use, then all of the offspring from a single female would likely prefer the same host-plant species. This experiment was also designed to test the Hopkins’ host selection principle; does the food plant on which a female is reared as a larva influence her future choices when she is searching for host plants for her own offspring? 4. The results from all of these experiments indicate that P. m. aliaska females vary greatly in their oviposition behaviour and in their preferences for the three host plants. Most populations appear to consist of generalists with labile oviposition behaviour. There is no evidence to support the Hopkins’ host selection principle. 5. It is suggested that the generalised selection of host plants by P. m. aliaska females may be a ‘bet-hedging’ strategy and that this strategy may maximise reproductive fitness in an unpredictable environment.     

Transgenerational inheritance of learned preferences for novel host plant odors in Bicyclus anynana butterflies

Many phytophagous insects have strong preferences for their host plants, which they recognize via odors, making it unclear how novel host preferences develop in the course of insect diversification. Insects may learn to prefer new host plants via exposure to their odors and pass this learned preference to their offspring. We tested this hypothesis by examining larval odor preferences before and after feeding them with leaves coated with control and novel odors and by examining odor preferences again in their offspring. Larvae of the parental generation developed a preference for two of these odors over their development. These odor preferences were also transmitted to the next generation. Offspring of butterflies fed on these new odors chose these odors more often than offspring of butterflies fed on control leaves. In addition, offspring of butterflies fed on banana odors had a significant naïve preference for the banana odors in contrast to the naïve preference for control leaves shown by individuals of the parental generation. Thus, butterflies can learn to prefer novel host plant odors via exposure to them during larval development and transmit these learned preferences to their offspring. This ability potentially facilitates shifts in host plant use over the course of insect diversification.     

GENETIC COVARIANCE BETWEEN OVIPOSITION PREFERENCE AND LARVAL PERFORMANCE IN AN INSECT HERBIVORE

An experimental study determined that females of the herbivorous fly species Liriomyza sativae (Diptera: Agromyzidae) preferentially oviposit on the plant species on which their female progeny attain the greatest pupal weight. A modified parent/offspring regression was used to quantify this relationship as an additive genetic covariance between host-plant preference and relative performance of female larvae on different plant species. The implications of a genetic covariance between preference and performance on the course of evolution in herbivores are discussed. Several females from one population refused to oviposit on one of the plant species; this population also suffered the only significant larval mortality on this plant. These results corroborate the avoidance of unsuitable host plants seen in the genetic analyses of individuals, but relative to the genetic data, such population-level data are of limited usefulness in the study of evolutionary mechanisms by which insect populations become adapted to their host plants.     

Butterflies and plants: preference/performance studies in relation to plant size and the use of intact plants vs. cuttings

Plants have evolved a number of defences to ameliorate herbivore attacks including chemicals induced by mechanical wounding. Such changes in plant chemical composition are potential confounding factors in experiments on plant – insect interactions, which often present cuttings of potential host plants to phytophagous insects. In particular, this could affect studies of female egg‐laying preference and larval performance, because the same plant chemicals that deter certain generalist insects can elevate attacks from more specialized insects. Furthermore, plant cuttings are by definition smaller than intact plants, and any female host size preference could thus affect experiments using plant cuttings. We first assessed female preference and larval performance of a specialist herbivore, Pieris napi (L.) (Lepidoptera: Pieridae, Pierini), confronted with either intact plants or leaf‐cuttings of four Brassicaceae host plants, Alliaria petiolata (Bieb.) Cavara & Grande, Barbarea vulgaris (L.) WT Aiton, Berteroa incana (L.) DC., and Brassica napus (L.). Egg and larval survival did not differ between intact plants and leaf‐cuttings, whereas larval growth was slightly, but significantly, faster on leaf‐cuttings. Females, however, significantly preferred to lay eggs on intact plants of all four hosts, although the preference hierarchy for the intact plants was largely mirrored by that for leaf‐cuttings. We then tested the female preference for different size‐classes of intact B. napus plants. Small individuals received more eggs than larger individuals, and follow‐up experiments showed that this difference was largely generated by a strong female preference for cotyledon leaves; there was no significant difference in female preference for large and small individuals when both carried cotyledons, and females landing on cotyledons were more likely to oviposit compared to when landing on a true leaf. Our study concludes that plant cuttings can serve as adequate proxies for live plants for preference/performance studies, but that experimentalists should be aware of the variation imposed both by plant handling and plant phenology for female oviposition preference.     

Host preference variation cannot explain microhabitat differentiation among sympatric Pieris napi and Pieris rapae butterflies

1. Often, closely related insect species feed on different host plant species, and the tremendous diversity of phytophagous insects is therefore attributed to host plant‐driven speciation. However, for most taxa, host use information comes from field observations of egg‐laying females or feeding caterpillars, which means that the underlying reason for a particular host‐affiliation is not easily determined. 2. Therefore, it is often unclear whether an insect feeds on a certain host because it prefers that plant to alternative hosts, or because the host distribution overlaps with the habitat requirements of the insect. 3. We ask to what extent a divergent host use in the field mirrors the host plant preferences of two closely related butterflies, Pieris napi and Pieris rapae (Pieridae). In nature, P. napi typically occurs in moister habitats than P. rapae. 4. We scanned several microhabitats at a field site in Southern Sweden during multiple years, and collected Pieris eggs from three different plants, Cardamine pratensis (wet meadows), Barbarea vulgaris (drier micro‐habitats) and Alliaria petiolata (intermediate areas). 5. As predicted, P. rapae eggs were more common than P. napi eggs on B. vulgaris, whereas all of the 358 individuals collected from C. pratensis were P. napi, indicating a divergence in host use between the Pieris species. However, under controlled laboratory conditions, both species had virtually identical oviposition preferences, laying eggs on all three plants, notably P. rapae also laying eggs on C. pratensis, indicating that habitat use, not plant preference, drives host plant use in nature.     

The influence of host plant variation and intraspecific competition on oviposition preference and offspring performance in the host races of Eurosta solidaginis

1. A series of experiments was conducted to measure the impact of plant genotype, plant growth rate, and intraspecific competition on the oviposition preference and offspring performance of the host races of Eurosta solidaginis (Diptera: Tephritidae), a fly that forms galls on Solidago altissima and Solidago gigantea (Asteraceae). Previous research has shown that both host races prefer to oviposit on their own host plant where survival is much higher than on the alternate host plant. In this study, neither host race showed any relationship between oviposition preference and offspring performance in choosing among plants of their natal host species. 2. The larval survival of both host races differed among plant genotypes when each host race oviposited on its natal host species. In one experiment, altissima host race females showed a preference among plant genotypes that was not correlated with offspring performance on those genotypes. In all other experiments, neither the altissima nor gigantea host race demonstrated a preference for specific host plant genotypes. 3. Eurosta solidaginis had a preference for ovipositing on rapidly growing ramets in all experiments, however larval survival was not correlated with ramet growth rate at the time of oviposition. 4. Eurosta solidaginis suffered high mortality from intraspecific competition in the early larval stage. There was little evidence, however, that females avoided ovipositing on ramets that had been attacked previously. This led to an aggregated distribution of eggs among ramets and strong intraspecific competition. 5. There was no interaction among plant genotype, plant growth rate, or intraspecific competition in determining oviposition preference or offspring performance.     

Oviposition preference,larval performance and adaptation of Trichoplusia ni on cabbage and cotton

Most female herbivores ensure to lay eggs where their offspring can develop successfully. The oviposition preferences of females affect strategies in pest management. In this study, the performance of two cohorts of Trichoplusia ni larvae on cabbage and cotton (after they had been transferred from their original host plants) were investigated. The preferences of female moth ovipositing and larval feeding on these two host plants were observed. The results indicated that plants significantly affected oviposition preference of the female adults and development and survival of larvae of T. ni. All females preferred to lay eggs on cabbage than cotton regardless from which host they originated. The detrimental effects of cotton on the development and survival of T. ni larvae originated from cabbage (CaTn) increased with the increase of the larval age when they were transferred. In addition, the host plant change did not significantly affect the development and survival of larvae of T. ni originating from cotton (CoTn). Larvae of CaTn preferred cabbage plants as compared to cotton plants, whereas larvae of CoTn did not show a significant choice. Although the adult females preferred laying eggs on cabbage, they did not show preferences between cotton and cabbage in a Y‐tube olfactometer test. The hypothesis of oviposition preference and performance of larvae was supported by the results of CaTn, whereas they not supported by those from CoTn. Based on these results, the strategy to manage this serious pest was discussed.     

A tritrophic analysis of host preference and performance in a polyphagous leafminer

The optimal oviposition theory predicts that oviposition preferences of phytophagous insects should correlate with host suitability for their offspring. As plant host suitability depends not only on its quality as food, but also on its provision of enemy‐free space, we examined the relationship between adult host preference and offspring performance for the leafminer Liriomyza huidobrensis (Blanchard) (Diptera: Agromyzidae) on various host plants, considering also the interaction with natural enemies. Preference and offspring performance were assessed through observational field data and laboratory experiments in central Argentina. Field data suggested a positive host preference – performance linkage, as the leafminer attained larger body size on the crops where it was more abundant. Laboratory trials supported these results: Vicia faba L. (Fabaceae) was the preferred host in the laboratory as well as in the field, performance of L. huidobrensis being also best on this host, with highest survival rates and shortest development time. The actively feeding larval stage showed the largest plant‐related effects. Higher overall parasitism rates were found on plants from which smaller leafminers were reared, reinforcing the preference–performance linkage. On the other hand, the main parasitoid Phaedrotoma scabriventris Nixon (Hymenoptera: Braconidae) reached larger body size, and caused higher mortality rates on crops where the leafminer was larger. Changes in abundance of particular parasitoid species could thus modify overall parasitism trends.     

Oviposition preference and larval performance of North American monarch butterflies on four Asclepias species

Monarch butterflies, Danaus plexippus L. (Lepidoptera: Nymphalidae), occur world‐wide and are specialist herbivores of plants in the milkweed family (Asclepiadaceae). In North America, two monarch populations breed east and west of the continental divide in areas populated by different host plant species. To examine the population variation in monarch responses to different Asclepias species, we measured oviposition preference and larval performance among captive progeny reared from adult butterflies collected in eastern and western North America. Host plant use was evaluated using two milkweed species widely distributed in eastern North America (A. incarnata and A. syriaca), and two species common to western North America (A. fascicularis and A. speciosa). We predicted that exposure to different host plant species in their respective breeding ranges could select for divergent host use traits, so that monarchs should preferentially lay more eggs on, and larvae should perform better on, milkweed species common to their native habitats. Results showed that across all adult female butterflies, oviposition preferences were highest for A. incarnata and lowest for A. fascicularis, but mean preferences did not differ significantly between eastern and western monarch populations. Larvae from both populations experienced the highest survival and growth rates on A. incarnata and A. fascicularis, and we again found no significant interactions between monarch source population and milkweed species. Moreover, the average rank order of larval performance did not correspond directly to mean female oviposition preferences, suggesting that additional factors beyond larval performance influence monarch oviposition behavior. Finally, significant family level variation was observed for both preference and performance responses within populations, suggesting an underlying genetic variation or maternal effects governing these traits.