Genetic and phenotypic variation in diet breadth in a generalist herbivore

Summary The genetic and plastic components of polyphagy were investigated in a population ofLymantria dispar, the gypsy moth. A simple genetic experiment assessed the expression of (1) genetic variability in life history traits within each of four environments, (2) genetic variability in diet breadth, expressed as a change in the ranks of family performance across hosts, and (3) homeostasis (equivalent performance by a family across hosts) versus phenotypic plasticity (variable performance by a family across hosts). Sibs from each of 14 families, randomly selected from a single population, were reared on four diets: two natural hosts — chestnut and red oak, and two synthetic hosts — a standard laboratory diet and a low-protein version of this diet. Average population performance, measured in terms of development time and pupal weight, was better on standard laboratory diet than on low-protein diet, and was equal on chestnut and red oak for pupal weight, but better on chestnut oak for development time. Average population performance provided no information about the genetic component of host use ability. The gypsy moth expressed genetic variation in development time within each host environment and in pupal weight within natural host environments. Phenotypic plasticity was expressed by a significant number of families in development time and pupal weight across synthetic hosts and, to a lesser extent, across natural hosts. It was only across natural hosts that genetic variation in diet breadth was expressed, and this was confined to females. Genetic variability in diet breadth may be maintained in this species as a consequence of the unpredictability of its food sources.     

Rapid evolution and specialization following host colonization in a yucca moth

Host shifts followed by specialization can result in sympatric genetic differentiation, and may have fuelled the diversification of phytophagous insects. This study examines a recent colonization of a non‐native host by Prodoxus quinquepunctellus (Lepidoptera: Prodoxidae). Allozyme differentiation was detected among different host feeding populations, yet was nearly absent among similar host feeding populations in sympatry. Geographical patterns of allozyme variation showed a much higher level of population structure among populations feeding on the derived host. Conversely, mtDNA haplotype frequencies were nearly homogeneous in the derived populations compared to the ancestral populations, suggesting a bottleneck and/or rapid fixation of haplotypes following host colonization. Moth emergence coincided with host plant flowering, and phenological differences between host species translated into allochronic isolation between populations feeding on different hosts. Derived moth populations also differed significantly in three ovipositor characters from ancestral populations. These findings suggest rapid host‐specific genetic differentiation, and specialization of moth emergence time and ovipositor morphology following host colonization.     

Field-based insights to the evolution of specialization: plasticity and fitness across habitats in a specialist/generalist species pair

Factors promoting the evolution of specialists versus generalists have been little studied in ecological context. In a large-scale comparative field experiment, we studied genotypes from naturally evolved populations of a closely related generalist/specialist species pair (Polygonum persicaria and P. hydropiper), reciprocally transplanting replicates of multiple lines into open and partially shaded sites where the species naturally co-occur. We measured relative fitness, individual plasticity, herbivory, and genetic variance expressed in the contrasting light habitats at both low and high densities. Fitness data confirmed that the putative specialist out-performed the generalist in only one environment, the favorable full sun/low-density environment to which it is largely restricted in nature, while the generalist had higher lifetime reproduction in both canopy and dense neighbor shade. The generalist, P. persicaria, also expressed greater adaptive plasticity for biomass allocation and leaf size in shaded conditions than the specialist. We found no evidence that the ecological specialization of P. hydropiper reflects either genetically based fitness trade-offs or maintenance costs of plasticity, two types of genetic constraint often invoked to prevent the evolution of broadly adaptive genotypes. However, the patterns of fitness variance and herbivore damage revealed how release from herbivory in a new range can cause an introduced species to evolve as a specialist in that range, a surprising finding with important implications for invasion biology. Patterns of fitness variance between and within sites are also consistent with a possible role for the process of mutation accumulation (in this case, mutations affecting shade-expressed phenotypes) in the evolution and/or maintenance of specialization in P. hydropiper.     

Predictability, toxicity, and trophic niche breadth in fungus-feeding Drosophilidae (Diptera)

Abstract. 1. The mycophagous Drosophilidae flies in eastern North America display a range of ecological responses to their fungal trophic resources. Some ( Drosophilu duncani Sturtevant, and two species labelled as Mycodrosophila cluytonae Wheeler and Takada) specialize on the bracket fungi of the family Polyporaceae. Other species ( Drosophila falleni Wheeler, D.recens Wheeler, D.putridu Sturtevant and D.tripunctuta Loew) are broad generalists, non-selectively consuming a diverse array of Basidioniycete mushrooms. D.restaceu von Roser and D.ordinuria Coquillett utilize a broad subset of the Basidio-mycetes, while M.dimidiata hew feeds on some Basidioniycetes, including a variety of Clavariaceae (coral fungi) and Heterobasidiomycetes (jelly fungi) not commonly used by other flies, and Ascomycetes (cup fungi).
2. Unlike some phytophagous insects, host chemistry seems to have little effect on host preferences. The mycophagous drosophilids do not avoid fungi thought to be toxic or undigestible to other Diptera.
3. Resource predictability appears to delimit trophic niches in the mycophagous guild. The average duration of individual host mushroom caps is negatively correlated with three measures of trophic niche breadth. Specialist fly species utilize long-lasting resources, while more generalized mycophagous species include progressively ephemeral fungi in their diets.     

Determinants of predation on phytophagous insects: the importance of diet breadth

To evaluate the role of predation in the evolution of diet specialization and to determine the effectiveness of various larval defenses, we offered lepidopteran larvae to colonies of the tropical ant Paraponera clavata. We recorded behavioral and physical characteristics of prey items and used log-linear models to analyze their importance as deterrents to predation by P. clavata. The most important determinant of probability of prey rejection by P. clavata was a prey's diet breadth; specialists were rejected by the ants significantly more than generalists. Other less important, but significant, predictors of prey rejection included ontogeny, morphology and chemistry. Late instar caterpillars were rejected more frequently than early instars, hairy caterpillars were rejected more frequently than caterpillars with other morphologies, and one caterpillar species with an unpalatable extract was rejected more frequently than two species with palatable extracts.     

Dietary niche breadth in a local community of passerine birds: an analysis using phylogenetic contrasts

The analysis of a local community of forest passerines (13 species) using phylogenetic contrasts shows a correlation between body size of bird species and mean prey size, minimum prey size, maximum prey size and the size range of dietary items. This suggests that larger birds drop small prey taxa from their prey list, because of the difficulty of capturing very small prey, for energetic reasons or because of microhabitat usage. We find some support for the third hypothesis. Dietary niche breadth calculated across prey taxa is not related to body size. Dietary niche breadth, however, is correlated with size-corrected measurements of the bill and locomotor apparatus. Long and slender bills increase the dietary niche breadth. Thus subtle differences constrain foraging and the techniques of extracting certain prey taxa form crevices. Dietary niche breadth and foraging diversity are positively correlated with population density: at least locally dietary generalists occur at higher breeding densities than specialists.     

Low parasitoid success on a myrmecophilous host is maintained in the absence of ants

Abstract.  1. Studies of Dinocampus coccinellae , a parasitoid of ladybird beetles, have generally shown congruence between field parasitism rates of different host species and parasitoid preference and/or host suitability in the laboratory, suggesting that host intrinsic factors rather than habitat-related extrinsic factors are of greatest importance in determining D. coccinellae occurrence.
2. The myrmecophilous Coccinella magnifica exhibits much lower D. coccinellae prevalence in the field than most other Coccinella species: it has been suggested that this is a manifestation of enemy-free space provided by the predatory Formica rufa group ants with which the C. magnifica occurs.
3. Coccinella magnifica collected at the same time and locality as parasitised Coccinella septempunctata were unparasitised by D. coccinellae . In the laboratory, in the absence of ants, although the parasitoid attacked C. magnifica as readily as C. septempunctata , C. magnifica was not parasitised successfully.
4. Such results are consistent with those from other ladybirds and C. magnifica does not now benefit directly from any putative D. coccinellae -free space provided by aggressive ants. Because its close relatives exhibit high levels of D. coccinellae parasitism, C. magnifica may be useful in determining some elements important in the evolution of host protection against parasitoid attack.     

Geographical variation in genet (Genetta genetta L.) diet: a literature review

The geographical variation in the Genet Genetta genetta L. diet was analysed from 12 locations covering its entire range. Data were obtained from the available literature on food studies in this species. We studied the general food spectrum of this species and compared the importance of different prey items in each area through PCA analysis. The possible influence of some large-scale environmental factors (latitude, altitude, Mediterraneity) on diet were studied by means of regression analysis and anova . The study assessed the frequency of occurrence of each prey group and diet diversity. Results show the existence of two ‘food’ groups: (i) Genets which feed on a wide food spectrum, especially arthropods and (ii) Genets which feed on small mammals at a high frequency, while the remaining prey items are scarce or absent. At the intraspecific level, Genets behave as generalist species, with the small mammals (especially, the Woodmouse) as the most outstanding prey item. However, when compared with the diet of other medium-sized Palaearctic carnivores, we can say that the Genet is intermediate between typical generalists (Martens Martes spp., Red Foxes Vulpes vulpes and Badgers Meles meles) and specialists (Otters Lutra lutra, Stoat Mustela erminea and Weasel Mustela nivalis). Finally, anova shows the existence of a relationship between diet diversity and Mediterraneity (associated with taxa such as arthropods, reptiles and amphibians), but no relationship was found for latitude or altitude. The importance of interspecific competition, based on island data, is discussed.     

Niche-tracking migrants and niche-switching residents: evolution of climatic niches in New World warblers (Parulidae)

Differences in life-history traits between tropical and temperate lineages are often attributed to differences in their climatic niche dynamics. For example, the more frequent appearance of migratory behaviour in temperate-breeding species than in species originally breeding in the tropics is believed to have resulted partly from tropical climatic stability and niche conservatism constraining tropical species from shifting their ranges. However, little is known about the patterns and processes underlying climatic niche evolution in migrant and resident animals. We evaluated the evolution of overlap in climatic niches between seasons and its relationship to migratory behaviour in the Parulidae, a family of New World passerine birds. We used ordination methods to measure seasonal niche overlap and niche breadth of 54 resident and 49 migrant species and used phylogenetic comparative methods to assess patterns of climatic niche evolution. We found that despite travelling thousands of kilometres, migrants tracked climatic conditions across the year to a greater extent than tropical residents. Migrant species had wider niches than resident species, although residents as a group occupied a wider climatic space and niches of migrants and residents overlapped extensively. Neither breeding latitude nor migratory distance explained variation among species in climatic niche overlap between seasons. Our findings support the notion that tropical species have narrower niches than temperate-breeders, but does not necessarily constrain their ability to shift or expand their geographical ranges and become migratory. Overall, the tropics may have been historically less likely to experience the suite of components that generate strong selection pressures for the evolution of migratory behaviour.     

Trade-offs and the evolution of host specialization

Summary Trade-offs in performance on different hosts are thought to promote the evolution of host specificity by blocking host shifts. Yet, in contrast, most experiments using phytophagous insects have shown performance on alternative hosts to be uncorrelated or positively correlated. Recent quantitative genetic models based on mutation—selection balance indicate that underlying constraints on the simultaneous maximization of different components of fitness may not always generate negative genetic correlations. We suggest an alternative or additional explanation for the lack of observed negative genetic correlations. If performance is polygenically controlled and some performance loci possess only antagonistically pleiotropic alleles, then the expression of trade-offs in performance will vary over time in populations. Consequently, a trade-off will be seen only in populations that have adapted to two hosts and are at or close to genetic equilibrium. Therefore, studies testing performance on a novel as compared with a normal host will generally yield non-negative genetic correlations between performance on the two hosts. The results of published studies are consistent with the predictions of this hypothesis.     

Role of larval host plants in the climate-driven range expansion of the butterfly Polygonia c-album

1. Some species have expanded their ranges during recent climate warming and the availability of breeding habitat and species' dispersal ability are two important factors determining expansions. The exploitation of a wide range of larval host plants should increase an herbivorous insect species' ability to track climate by increasing habitat availability. Therefore we investigated whether the performance of a species on different host plants changed towards its range boundary, and under warmer temperatures. 2. We studied the polyphagous butterfly Polygonia c-album, which is currently expanding its range in Britain and apparently has altered its host plant preference from Humulus lupulus to include other hosts (particularly Ulmus glabra and Urtica dioica). We investigated insect performance (development time, larval growth rate, adult size, survival) and adult flight morphology on these host plants under four rearing temperatures (18-28.5 degrees C) in populations from core and range margin sites. 3. In general, differences between core and margin populations were small compared with effects of rearing temperature and host plant. In terms of insect performance, host plants were generally ranked U. glabra > or = U. dioica > H. lupulus at all temperatures. Adult P. c-album can either enter diapause or develop directly and higher temperatures resulted in more directly developing adults, but lower survival rates (particularly on the original host H. lupulus) and smaller adult size. 4. Adult flight morphology of wild-caught individuals from range margin populations appeared to be related to increased dispersal potential relative to core populations. However, there was no difference in laboratory reared individuals, and conflicting results were obtained for different measures of flight morphology in relation to larval host plant and temperature effects, making conclusions about dispersal potential difficult. 5. Current range expansion of P. c-album is associated with the exploitation of more widespread host plants on which performance is improved. This study demonstrates how polyphagy may enhance the ability of species to track climate change. Our findings suggest that observed differences in climate-driven range shifts of generalist vs. specialist species may increase in the future and are likely to lead to greatly altered community composition.     

The genetic architecture of a niche: variation and covariation in host use traits in the Colorado potato beetle

The genetic basis of host plant use by phytophagous insects can provide insight into the evolution of ecological niches, especially phenomena such as specialization and phylogenetic conservatism. We carried out a quantitative genetic analysis of multiple host use traits, estimated on five species of host plants, in the Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). Mean values of all characters varied among host plants, providing evidence that adaptation to plants may require evolution of both behavioral (preference) and post-ingestive physiological (performance) characteristics. Significant additive genetic variation was detected for several characters on several hosts, but not in the capacity to use the two major hosts, a pattern that might be caused by directional selection. No negative genetic correlations across hosts were detected for any 'performance' traits, i.e. we found no evidence of trade-offs in fitness on different plants. Larval consumption was positively genetically correlated across host plants, suggesting that diet generalization might evolve as a distinct trait, rather than by independent evolution of feeding responses to each plant species, but several other traits did not show this pattern. We explored genetic correlations among traits expressed on a given plant species, in a first effort to shed light on the number of independent traits that may evolve in response to selection for host-plant utilization. Most traits were not correlated with each other, implying that adaptation to a novel potential host could be a complex, multidimensional 'character' that might constrain adaptation and contribute to the pronounced ecological specialization and the phylogenetic niche conservatism that characterize many clades of phytophagous insects.     

Phylogenetic evidence for the evolution of ecological specialization in Timema walking‐sticks

We used phylogenetic and ecological information to study the evolution of host‐plant specialization and colour polymorphism in the genus Timema, which comprises 14 species of walking‐sticks that are subject to strong selection for cryptic coloration on their host‐plants. Phylogenetic analysis indicated that this genus consists of three main lineages. Two of the lineages include highly generalized basal species and relatively specialized distal species, and one of the lineages comprises four specialized species. We tested for phylogenetic conservatism in the traits studied via randomizing host‐plant use, and the four basic Timema colour patterns, across the tips of the phylogeny, and determining if the observed number of inferred changes was significantly low compared to the distribution of numbers of inferred changes expected under the null model. This analysis showed that (1) host‐plant use has evolved nonrandomly, such that more closely related species tend to use similar sets of hosts and (2) colour pattern evolution exhibits considerable lability. Inference of ancestral states using maximum parsimony, under four models for the relative ease of gain and loss of plant hosts or colour morphs, showed that (1) for all models with gains of host‐plants even marginally more difficult than losses, and for most optimizations with gains and losses equally difficult, the ancestral Timema were generalized, feeding on the chaparral plants Ceanothus and Adenostoma and possibly other taxa, and (2) for all models with gains of colour morphs more difficult than losses, the ancestral Timema were polymorphic for colour pattern. Generation of null distributions of inferred ancestral states showed that the maximum‐parsimony inference of host‐plant generalization was most robust for the most speciose of the three main Timema lineages. Ancestral states were also inferred using maximum likelihood, after recoding host‐plant use and colour polymorphism as dichotomous characters. Likelihood analyses provided some support for inference of generalization in host‐plant use at ancestral nodes of the two lineages exhibiting mixtures of generalists and specialists, although levels of uncertainty were high. By contrast, likelihood analysis did not estimate ancestral colour morph patterns with any confidence, due to inferred rates of change that were high with respect to speciation rates. Information from biogeography, floristic history and the timing of diversification of the genus are compatible with patterns of inferred ancestral host‐plant use. Diversification in the genus Timema appears to engender three main processes: (1) increased specialization via loss of host‐plants, (2) retention of the same, single, host‐plant and (3) shifts to novel hosts to which lineages were ‘preadapted’ in colour pattern. Our evidence suggests that the radiation of this genus has involved multiple evolutionary transitions from individual‐level specialization (multiple‐niche polymorphism) to population‐level and species‐level specialization. Ecological studies of Timema suggest that such transitions are driven by diversifying selection for crypsis. This paper provides the first phylogeny‐based evidence for the macroevolutionary importance of predation by generalist natural enemies in the evolution of specialization.     

Trophic specialization influences the rate of environmental niche evolution in damselfishes (Pomacentridae)

The rate of environmental niche evolution describes the capability of species to explore the available environmental space and is known to vary among species owing to lineage-specific factors. Trophic specialization is a main force driving species evolution and is responsible for classical examples of adaptive radiations in fishes. We investigate the effect of trophic specialization on the rate of environmental niche evolution in the damselfish, Pomacentridae, which is an important family of tropical reef fishes. First, phylogenetic niche conservatism is not detected in the family using a standard test of phylogenetic signal, and we demonstrate that the environmental niches of damselfishes that differ in trophic specialization are not equivalent while they still overlap at their mean values. Second, we estimate the relative rates of niche evolution on the phylogenetic tree and show the heterogeneity among rates of environmental niche evolution of the three trophic groups. We suggest that behavioural characteristics related to trophic specialization can constrain the evolution of the environmental niche and lead to conserved niches in specialist lineages. Our results show the extent of influence of several traits on the evolution of the environmental niche and shed new light on the evolution of damselfishes, which is a key lineage in current efforts to conserve biodiversity in coral reefs.     

Egg-laying tactic in Phyllomorpha laciniata in the presence of parasitoids

Female insects are expected to choose oviposition sites that have the best conditions for offspring development and survival. Natural enemies, such as predators and parasitoids, may have a strong influence on the selection of oviposition substrates by phytophagous insects. The golden egg bug, Phyllomorpha laciniata (Villers) (Heteroptera: Coreidae) has an unusual reproductive strategy. Females mainly use conspecifics, both males and other females, as egg-laying substrates, but occasionally they oviposit on plants as well. Survival of the eggs is higher when eggs are carried by conspecifics than when they are laid on plants, due to predation and parasitism. We investigated egg-laying behavior in the forced presence of the egg parasitoid Gryon bolivari (Giard) (Hymenoptera: Scelionidae). Specifically, we studied whether females provide egg protection by avoiding oviposition under the risk of egg parasitization. We expected a lower oviposition rate under parasitoid presence, and the eggs, if any, to be placed preferably on conspecifics and not on plants, thus ensuring higher survival of the progeny. The results show that P. laciniata 's egg-laying rate was lower when they were enclosed with parasitoids than when parasitoids were absent, especially when plants were the only substrate to oviposit on. Moreover, females showed strong preference for laying eggs on conspecifics rather than on plants. Egg-laying in P. laciniata appears to be not only influenced by the availability of conspecifics, but also by the presence of egg parasitoids. This indicates that females may be able to detect G. bolivari and avoid oviposition when parasitoids are present. We discuss the possibility of conspecifics as enemy-free space.     

The distribution and density of a lycaenid butterfly in relation to Lasius ants

Summary Larvae and pupae of lycaenid butteflies are often associated with ants: this is usually a mutualism in which ants guard the lycaenids from natural enemies, and the lycaenid larvae and pupae provide sugars and amino acids for the ants. A possible consequence of the interaction is spatially correlated ant and lycaenid distributions, but the phenomenon is poorly documented. We examined the lycaenid Plebejus argus, which is tended by Lasius ants. Within habitat patches, P. argus eggs, larvae and pupae were all spatially associated with Lasius. On a larger scale, the densities of butterflies in different habitat patches and populations, and whether the butterfly was present or not, were correlated with Lasius ant densities. The association of P. argus with Lasius ants is consistent among populations, and occurs at several spatial scales. Other aspects of the ecology of P. argus are more variable.     

Use it or lose it: reproductive implications of ecological specialization in a haematophagous ectoparasite

Using experimentally induced disruptive selection, we tested two hypotheses regarding the evolution of specialization in parasites. The 'trade-off' hypothesis suggests that adaptation to a specific host may come at the expense of a reduced performance when exploiting another host. The alternative 'relaxed selection' hypothesis suggests that the ability to exploit a given host would deteriorate when becoming obsolete. Three replicate populations of a flea Xenopsylla ramesis were maintained on each of two rodent hosts, Meriones crassus and Dipodillus dasyurus, for nine generations. Fleas maintained on a specific host species for a few generations substantially decreased their reproductive performance when transferred to an alternative host species, whereas they generally did not increase their performance on their maintenance host. The results support the 'relaxed selection' hypothesis of the evolution of ecological specialization in haematophagous ectoparasites, while suggesting that trade-offs are unlikely drivers of specialization. Further work is needed to study the extent by which the observed specializations are based on epigenetic or genetic modifications.     

The relationship between diet breadth and geographic range size in the butterfly subfamily Nymphalinae--a study of global scale

The "oscillation hypothesis" has been proposed as a general explanation for the exceptional diversification of herbivorous insect species. The hypothesis states that speciation rates are elevated through repeated correlated changes--oscillations--in degree of host plant specificity and geographic range. The aim of this study is to test one of the predictions from the oscillation hypothesis: a positive correlation between diet breadth (number of host plants used) and geographic range size, using the globally distributed butterfly subfamily Nymphalinae. Data on diet breadth and global geographic range were collected for 182 Nymphalinae butterflies species and the size of the geographic range was measured using a GIS. We tested both diet breadth and geographic range size for phylogenetic signal to see if species are independent of each other with respect to these characters. As this test gave inconclusive results, data was analysed both using cross-species comparisons and taking phylogeny into account using generalised estimating equations as applied in the APE package in R. Irrespective of which method was used, we found a significant positive correlation between diet breadth and geographic range size. These results are consistent for two different measures of diet breadth and removal of outliers. We conclude that the global range sizes of Nymphalinae butterflies are correlated to diet breadth. That is, butterflies that feed on a large number of host plants tend to have larger geographic ranges than do butterflies that feed on fewer plants. These results lend support for an important step in the oscillation hypothesis of plant-driven diversification, in that it can provide the necessary fuel for future population fragmentation and speciation.     

Maintenance of narrow diet breadth in the monarch butterfly caterpillar: response to various plant species and chemicals

In order to better understand the maintenance of a fairly narrow diet breadth in monarch butterfly larvae, Danaus plexippus L. (Lepidoptera: Nymphalidae: Danainae), we measured feeding preference and survival on host and non-host plant species, and sensitivity to host and non-host plant chemicals. For the plant species tested, a hierarchy of feeding preferences was observed; only plants from the Asclepiadaceae were more or equally preferred to Asclepias curassavica, the common control. The feeding preferences among plant species within the Asclepiadaceae are similar to published mean cardenolide concentrations. However, since cardenolide data were not collected from individual plants tested, definitive conclusions regarding cardenolide concentrations and plant acceptability cannot be made. Although several non-Asclepiadaceae were eaten in small quantities, all were less preferred to A. curassavica. Additionally, these non-Asclepiadaceae do not support continued feeding, development, and survival of first and fifth-instar larvae. Preference for a host versus a non-host (A. curassavica versus Vinca rosea) increased for A. curassavica reared larvae as compared to diet-reared larvae suggesting plasticity in larval food preferences. Furthermore, host species were significantly preferred over non-host plant species in bioassays using a host plant or sucrose as a common control. Larval responses to pure chemicals were examined in order to determine if host and non-host chemicals stimulate or deter feeding in monarch larvae. We found that larvae were stimulated to feed by some ubiquitous plant chemicals, such as sucrose, inositol, and rutin. In contrast, several non-host plant chemicals deterred feeding: caffeine, apocynin, gossypol, tomatine, atropine, quercitrin, and sinigrin. Additionally the cardenolides digitoxin and ouabain, which are not in milkweed plants, were neutral in their influence on feeding. Another non-milkweed cardenolide, cymarin, significantly deterred feeding. Extracts of A. curassavica leaves were tested in bioassays to determine which components of the leaf stimulate feeding. Both an ethanol extract of whole leaves and a hexane leaf-surface extract are phagostimulatory, suggesting the involvement of both polar and non-polar plant compounds. These data suggest that the host range of D. plexippus larvae is maintained by both feeding stimulatory and deterrent chemicals in host and non-host plants.