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AbstractIntraspecific variation in camouflage is common in animals. Sexual dimorphism in camouflage is less common and, where observed, attributed to trade-offs between natural selection for predator avoidance and sexual selection for conspicuous mating signals. Here we report on variation in putatively cryptic ventral hindwing patterns in the American snout butterfly, Libytheana carinenta . We use field surveys and crowdsourced data to characterize three morphs. One is found in both sexes, one is male specific, and one is female specific. The sex-specific morphs constitute a sexually dimorphic set whose frequencies change together in time. Field surveys indicate that butterflies in southern Arizona transition from midsummer dominance of the sexually monomorphic pattern to early-fall dominance of the sexually dimorphic set. Crowdsourced data indicate that the sexually dimorphic set dominates in early spring, transitioning later into a mixture of morphs dominated by the monomorphic pattern, with the dimorphic set rising in frequency again in late fall. We discuss this unique pattern of camouflage variation with respect to contemporary theory on animal coloration.

The evolution of flowers that offer oils as rewards and are pollinated by specialized bees represents a distinctive theme in plant-pollinator co-diversification. Some plants that offer acetylated glycerols as floral oils emit diacetin, a volatile by-product of oil metabolism, which is utilized by oil-collecting bees as an index signal for the presence of floral oil. However, floral oils in the genus Krameria (Krameriaceae) contain β-acetoxy-substituted fatty acids instead of acetylated glycerols, making them unlikely to emit diacetin as an oil-bee attractant. We analyzed floral headspace composition from K. bicolor and K. erecta, native to the Sonoran Desert of southwestern North America, in search of alternative candidates for volatile index signals. Using solid-phase microextraction, combined with gas chromatography-mass spectrometry, we identified 26 and 45 floral volatiles, respectively, from whole flowers and dissected flower parts of these two Krameria species. As expected, diacetin was not detected. Instead, β-ionone emerged as a strong candidate for an index signal, as it was uniquely present in dissected oil-producing floral tissues (elaiophores) of K. bicolor, as well as the larval cells and provisions from its oil-bee pollinator, Centris cockerelli. This finding suggests that the floral oil of K. bicolor is perfused with β-ionone in its tissue of origin and retains the distinctive raspberry-like scent of this volatile after being harvested by C. cockerelli bees. In contrast, the elaiophores of K. erecta, which are not thought to be pollinated by C. cockerelli, produced a blend of anise-related oxygenated aromatics not found in the elaiophores of K. bicolor. Our findings suggest that β-ionone has the potential to impact oil-foraging by C. cockerelli bees through several potential mechanisms, including larval imprinting on scented provisions or innate or learned preferences by foraging adults., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Organisms can often respond adaptively to a change in their environment through phenotypic plasticity in multiple traits, a phenomenon termed as multivariate plasticity. These different plastic responses could interact and affect each other's development as well as selection on each other, but the causes and consequences of these interactions have received relatively little attention. Here, we propose a new conceptual framework for understanding how different plastic responses can affect each other's development and why organisms should have multiple plastic responses. A plastic change in one trait could alter the phenotype of a second plastic trait by changing either the cue received by the organism (cue-mediated effect) or the response to that cue (response-mediated effect). Multivariate plasticity could benefit the organism either because the plastic responses work better when expressed together (synergy) or because each response is more effective under different environmental circumstances (complementarity). We illustrate these hypotheses with case studies, focusing on interactions between behavior and morphology, plastic traits that differ in their reversibility. Future empirical and theoretical research should investigate the consequences of these interactions for additional factors important for the evolution of plasticity, such as the limits and costs of plasticity., (© 2022 The Authors. Evolution published by Wiley Periodicals LLC on behalf of The Society for the Study of Evolution.)

Animals sometimes have prominent projections on or near their heads serving diverse functions such as male combat, mate attraction, digging, capturing prey, sensing or defence against predators. Some butterfly larvae possess a pair of long frontal projections; however, the function of those projections is not well known. Hestina japonica butterfly larvae have a pair of long hard projections on their heads (i.e., horns). Here we hypothesized that they use these horns to protect themselves from natural enemies (i.e., predators and parasitoids). Field surveys revealed that the primary natural enemies of H. japonica larvae were Polistes wasps. Cage experiments revealed that larvae with horns intact and larvae with horns removed and fitted with horns of other individuals succeeded in defending themselves against attacks of Polistes wasps significantly more often than larvae with horns removed. We discuss that the horns counter the paper wasps' hunting strategy of first biting the larvae's 'necks' and note that horns evolved repeatedly only within the Nymphalidae in a phylogeny of the Lepidoptera. This is the first demonstration that arthropods use head projections for physical defence against predators., (© 2022. The Author(s).)

Within-species variation in pollinator behavior is widely observed, but its causes have been minimally investigated. Pollinator sex is associated with large differences in behavior that may lead to predictable differences in flower foraging, but this expectation has not been explicitly tested. We investigate sex-associated differences in nectar-foraging behavior of the hawkmoth Hyles lineata, using pollen on the proboscis as a proxy for flower visitation. We tested two predictions emerging from the literature: (1) the sexes differ in the flower species they visit, (2) females are more specialized in flower choice. We also examined potential drivers underlying these predictions by performing field and laboratory experiments to test whether males (3) switch among flower species more frequently, or (4) fly farther and therefore encounter more species than females. Consistent with prediction (1), pollen load composition differed between the sexes, indicative of visitation differences. Contrary to prediction (2), females consistently carried more species-rich pollen loads than males. (3) Both sexes switched between flower species at similar rates, suggesting that differences in floral fidelity are unlikely to explain why females are less specialized than males. (4) Males flew longer distances than females; coupled with larger between-site differences in pollen composition for females, this result suggests that sex differences in mobility influence foraging, and that females may forage more frequently and in smaller areas than males. Together, our results demonstrate that sex-associated foraging differences can be large and consistent over time, and highlight the importance of sex as a driver of variation in pollinator behavior., (© 2021 Institute of Zoology, Chinese Academy of Sciences.)

Mimicry is common in interspecies interactions, yet conditions maintaining Batesian mimicry have been primarily tested in predator-prey interactions. In pollination mutualisms, floral mimetic signals thought to dupe animals into pollinating unrewarding flowers are widespread (greater than 32 plant families). Yet whether animals learn to both correctly identify floral models and reject floral mimics and whether these responses are frequency-dependent is not well understood. We tested how learning affected the effectiveness and frequency-dependence of imperfect Batesian mimicry among flowers using the generalist bumblebee, Bombus impatiens , visiting Begonia odorata , a plant species exhibiting intersexual floral mimicry. Unrewarding female flowers are mimics of pollen-rewarding male flowers (models), though mimicry to the human eye is imperfect. Flower-naive bees exhibited a perceptual bias for mimics over models, but rapidly learned to avoid mimics. Surprisingly, altering the frequency of models and mimics only marginally shaped responses by naive bees and by bees experienced with the distribution and frequency of models and mimics. Our results provide evidence both of exploitation by the plant of signal detection trade-offs in bees and of resistance by the bees, via learning, to this exploitation. Critically, we provide experimental evidence that imperfect Batesian mimicry can be adaptive and, in contrast with expectations of signal detection theory, functions largely independently of the model and mimic frequency. This article is part of the theme issue 'Signal detection theory in recognition systems: from evolving models to experimental tests'.

Similar to animal communication displays, flowers emit complex signals that attract pollinators. Signal complexity could lead to higher cognitive load for pollinators, impairing performance, or might benefit them by facilitating learning, memory and decision making. Here, we evaluated learning and memory in foragers of the bumble bee Bombus impatiens trained to simple (unimodal) versus complex (bimodal) signals under restrained conditions. Use of a proboscis extension response protocol enabled us to control the timing and duration of stimuli presented during absolute and differential learning tasks. Overall, we observed broad variation in performance under the two conditions, with bees trained to compound bimodal signals learning and remembering as well as, better than or more poorly than bees trained to unimodal signals. Interestingly, the outcome of training was affected by the specific colour-odour combination. Among unimodal stimuli, the performance with odour stimuli was higher than with colour stimuli, suggesting that olfactory signals played a more significant role in the compound bimodal condition. This was supported by the fact that after 24 h, most bimodal-treatment bees responded to odour but not visual stimuli. We did not observe differences in latency of response, suggesting that signal composition affected decision accuracy, not speed. We conclude that restrained bumble bee workers exhibit broad variation of responses to bimodal stimuli and that components of the bimodal signal may not be used equivalently. The analysis of bee performance under restrained conditions enables accurate control of the multimodal stimuli provided to individuals and to study the interaction of individual components within a compound., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Pollen collection is necessary for bee survival and important for flowering plant reproduction, yet if and how pollen extraction motor routines are modified with experience is largely unknown. Here, we used an automated reward and monitoring system to evaluate modification in a common pollen-extraction routine, floral sonication. Through a series of laboratory experiments with the bumblebee, Bombus impatiens , we examined whether variation in sonication frequency and acceleration is due to instrumental learning based on rewards, a fixed behavioral response to rewards, and/or a mechanical constraint. We first investigated whether bees could learn to adjust their sonication frequency in response to pollen rewards given only for specified frequency ranges and found no evidence of instrumental learning. However, we found that absence versus receipt of a pollen reward did lead to a predictable behavioral response, which depended on bee size. Finally, we found some evidence of mechanical constraints, in that flower mass affected sonication acceleration (but not frequency) through an interaction with bee size. In general, larger bees showed more flexibility in sonication frequency and acceleration, potentially reflecting a size-based constraint on the range over which smaller bees can modify frequency and acceleration. Overall, our results show that although bees did not display instrumental learning of sonication frequency, their sonication motor routine is nevertheless flexible.

Intraspecific variation in floral visitor behaviour and pollination efficiency has been much less studied than interspecific variation. Nevertheless, it is clear that large differences in these traits exist within species, and in particular between sexes within species. With the exception of a few well-studied interactions, however, the consequences of these differences in the pollinators and visited plants remain to be investigated. In this review, we document large and consistent differences in the foraging patterns of male and female pollinators that have been demonstrated to directly affect plant reproduction or that have clear potential to do so. Males and females differ in visitation frequency, type of flowers visited, and per-visit pollen transfer. Females gather more and different resources from flowers compared to males, and males generally tend to show more mobile foraging patterns than females. We argue that these sex-associated patterns have broad generality across pollinators, and that sex-associated differences can in some cases be larger than differences between species. We offer predictions about how these patterns will influence pollinator preference, specialization, and fidelity, as well as the cost, quality and quantity of pollination service to plants. In the face of increasing threats to plant-pollinator interactions, understanding their basic functioning and the variation inherent in their component parts is critical. We advocate for more attention to sex-based differences among pollinators in particular, and the consequences of intraspecific variation more broadly., (© 2019 The Authors. Journal of Animal Ecology © 2019 British Ecological Society.)

Mimics should not exist without their models, yet often they do. In the system involving queen and viceroy butterflies, the viceroy is both mimic and co-model depending on the local abundance of the model, the queen. Here, we integrate population surveys, chemical analyses, and predator behavior assays to demonstrate how mimics may persist in locations with low-model abundance. As the queen becomes less locally abundant, the viceroy becomes more chemically defended and unpalatable to predators. However, the observed changes in viceroy chemical defense and palatability are not attributable to differing host plant chemical defense profiles. Our results suggest that mimetic viceroy populations are maintained at localities of low-model abundance through an increase in their toxicity. Sharing the burden of predator education in some places but not others may also lower the fitness cost of warning signals thereby supporting the origin and maintenance of aposematism., Competing Interests: The authors declare no competing interests.

Aggregations of conspecific animals are common and have been documented in most phyla. Multispecies aggregations are less common and less well studied. Eight species of Brachinus beetles -famous for their unique, highly effective, chemical defense-regularly settle together form large diurnal multispecies aggregations in dark, moist areas in riparian habitats in the Sonoran Desert Region. Here, we document these multispecies aggregations and investigate the incidence and dynamics of aggregation behavior. Analysis of species composition of 59 field-collected aggregations revealed that 71% contained more than one species, eight species regularly co-occurred in aggregations, and no two species showed a preference to aggregate with one another. We provide the first phylogenetic analyses of participants in multispecies aggregations, and find that Brachinus species found together in aggregations are not each other's closest relatives but rather are dispersed throughout the phylogeny of the genus. Further, we find no tendency for species to aggregate with close relatives more frequently than distant relatives. Laboratory experiments on B. elongatulus showed that it chose to settle in occupied shelters over empty shelters. Experiments with B. hirsutus and B. elongatulus showed that B. hirsutus prefers to settle under shelters housing heterospecifics over conspecifics. Our findings suggest that these multispecies aggregations do not form by chance, but rather are initiated by a genus-wide aggregation cue associated with the presence of individuals already in a shelter, which is likely to be chemical and potentially tactile in nature., Competing Interests: The authors have declared that no competing interests exist.

Sexual selection on male body size in species with a female-biased sexual size dimorphism is common yet often poorly understood. In particular, in the majority of bee species, the relative contribution of intrasexual competition and female choice to patterns of male body size is unknown. In this field study, we examined two possible components of male mating success with respect to body size in the solitary bee Diadasia rinconis Cockerell (Hymenoptera: Apidae): 1) ability to procure a mate and 2) the duration of copulation. We found that larger males were better able to procure mates and copulated for shorter periods of time. Although consistent with sperm competition theory, differences in copulation duration were slight; possibly, the shorter copulations of larger males instead reflect in copulo female choice. Consistent with this notion, males engaged in complex courtship while mounted, characterized for the first time in any bee in such detail via audio recordings and high-speed, high-definition video. The number of pulses in male courtship behavior was also positively associated with copulation duration and may have stimulated females to continue copulating, thereby potentially allowing smaller males to transfer a full ejaculate. Females were shown to be potentially polyandrous and although we did not observe precopulatory rejection in the field, captive females frequently rejected copulation attempts by captive males. Our work indicates that intrasexual competition selects for increased body size in a solitary bee.

The question of when to collect new information and how to apply that information is central to much of behaviour. Theory suggests that the value of collecting information, or sampling, depends on environmental persistence and on the relative costs of making wrong decisions. However, empirical tests of how these variables interact are lacking. We tested whether bumblebee foraging decisions are indeed influenced by these two factors. We gave bees repeated choices between a resource providing a steady, mediocre reward and a resource fluctuating between a low reward and a high reward. In this paradigm, we manipulated environmental persistence by changing how long the quality of a fluctuating resource remained stable at one reward level. We manipulated the costs of decision errors by changing the relative values of the available rewards. Bees sampled the fluctuating resource more frequently when it changed quality more frequently, indicating that they measured environmental persistence and reacted to it as predicted by theory. Bees showed surprisingly suboptimal tracking, not reliably choosing the currently best resource except when the fluctuating resource was very persistent and the potential rewards high. While bees modify their choices in response to different levels of change and potential rewards, they do not always do so according to optimality predictions., Competing Interests: We declare we have no competing interests.

Having multiple plastic responses to a change in the environment, such as increased temperature, can be adaptive for two major reasons: synergy (the plastic responses perform better when expressed simultaneously) or complementarity (each plastic response provides a greater net benefit in a different environmental context). We investigated these hypotheses for two forms of temperature-induced plasticity of Battus philenor caterpillars in southern Arizona populations: color change (from black to red at high temperatures) and heat avoidance behavior (movement from host to elevated refuges at high host temperatures). Field assays using aluminum models showed that the cooling effect of the red color is greatly reduced in a refuge position relative to that on a host. Field assays with live caterpillars demonstrated that refuge seeking is much more important for survival under hot conditions than coloration; however, in those assays, red coloration reduced the need to seek refuges. Our results support the complementarity hypothesis: refuge seeking facilitates survival during daily temperature peaks, while color change reduces the need to leave the host over longer warm periods. We propose that combinations of rapid but costly short-term behavioral responses and slow but efficient long-term morphological responses may be common when coping with temperature change.

The ecological success of social insects is frequently ascribed to improvements in task performance due to division of labour amongst workers. While much research has focused on improvements associated with lifetime task specialization, members of colonies can specialize on a given task over shorter time periods. Eusocial bees in particular must collect pollen and nectar rewards to survive, but most workers appear to mix collection of both rewards over their lifetimes. We asked whether bumblebees specialize over timescales shorter than their lifetime. We also explored factors that govern such patterns, and asked whether reward specialists made more foraging bouts than generalists. In particular, we described antennal morphology and size of all foragers in a single colony and related these factors to each forager's complete foraging history, obtained using radio frequency identification (RFID). Only a small proportion of foragers were lifetime specialists; nevertheless, >50% of foragers specialized daily on a given reward. Contrary to expectations, daily and lifetime reward specialists were not better foragers (being neither larger nor making more bouts); larger bees with more antennal olfactory sensilla made more bouts, but were not more specialized. We discuss causes and functions of short and long-term patterns of specialization for bumblebee colonies., Competing Interests: The authors declare no competing financial interests.

Many animals, including insects, make decisions using both personally gathered information and social information derived from the behavior of other, usually conspecific, individuals [1]. Moreover, animals adjust use of social versus personal information appropriately under a variety of experimental conditions [2-5]. An important factor in how information is used is the information's reliability, that is, how consistently the information is correlated with something of relevance in the environment [6]. The reliability of information determines which signals should be attended to during communication [6-9], which types of stimuli animals should learn about, and even whether learning should evolve [10, 11]. Here, we show that bumble bees (Bombus impatiens) account for the reliability of personally acquired information (which flower color was previously associated with reward) and social information (which flowers are chosen by other bees) in making foraging decisions; however, the two types of information are not treated equally. Bees prefer to use social information if it predicts a reward at all, but if social information becomes entirely unreliable, flower color will be used instead. This greater sensitivity to the reliability of social information, and avoidance of conspecifics in some cases, may reflect the specific ecological circumstances of bee foraging. Overall, the bees' ability to make decisions based on both personally acquired and socially derived information, and the relative reliability of both, demonstrates a new level of sophistication and flexibility in animal, particularly insect, decision-making., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Bees are model organisms for the study of learning and memory, yet nearly all such research to date has used a single reward, nectar. Many bees collect both nectar (carbohydrates) and pollen (protein) on a single foraging bout, sometimes from different plant species. We tested whether individual bumblebees could learn colour associations with nectar and pollen rewards simultaneously in a foraging scenario where one floral type offered only nectar and the other only pollen. We found that bees readily learned multiple reward-colour associations, and when presented with novel floral targets generalized to colours similar to those trained for each reward type. These results expand the ecological significance of work on bee learning and raise new questions regarding the cognitive ecology of pollination., (© 2015 The Author(s).)

The ability of individual animals to select habitats optimal for development and survival can be constrained by the costs of moving through the environment. Animals that seek overwintering sites underground, for example, may be constrained by the energy required to burrow into the soil. We conducted field and laboratory studies to determine the relationship between individual size and overwintering site selection in the tephritid flies, Rhagoletis juglandis and Rhagoletis suavis. We also explored the effect of site selection on pupal mortality, parasitism, and the ability to emerge from overwintering sites after eclosion. In both species, and in both lab and field tests, larger pupae were found at deeper soil depths. In addition, marginally non-significant trends indicated pupae in deeper sites were 48% more likely to survive the overwintering period. Finally, larger individuals were more likely to eclose and emerge from the soil at a given depth, but flies in deep overwintering sites were less likely to emerge from those sites than flies in shallow sites. Our data indicate that overwintering site selection represents a trade-off between avoiding predators and parasites that occur at shallow sites, and the energetic and mortality costs of burrowing to, overwintering in, and emerging from, deeper sites. The size-dependent overwintering site selection demonstrated here has implications for population dynamics and pest control strategies. Some fly control measures, such as the introduction of parasites or predators, will be mitigated when the deepest and least accessible overwintering pupae represent a disproportionately large amount of the population's reproductive capacity.

Animals sometimes develop conspicuous projections on or near their heads as, e.g., weaponry, burrowing or digging tools, and probes to search for resources. The frontal projections that insects generally use to locate and assess resources are segmented appendages, including antennae, maxillary palps, and labial palps. There is no evidence to date that arthropods, including insects, use projections other than true segmental appendages to locate food. In this regard, it is noteworthy that some butterfly larvae possess a pair of long antenna-like projections on or near their heads. To date, the function of these projections has not been established. Larvae of pipevine swallowtail butterflies Battus philenor (Papilionidae) have a pair of long frontal fleshy projections that, like insect antennae generally, can be actively moved. In this study, we evaluated the possible function of this pair of long moveable frontal projections. In laboratory assays, both frontal projections and lateral ocelli were shown to increase the frequency with which search larvae found plants. The frontal projections increased finding of host and non-host plants equally, suggesting that frontal projections do not detect host-specific chemical cues. Detailed SEM study showed that putative mechanosensillae are distributed all around the frontal as well as other projections. Taken together, our findings suggest that the frontal projections and associated mechanosensillae act as vertical object detectors to obtain tactile information that, together with visual information from lateral ocelli and presumably chemical information from antennae and mouthparts, help larvae to find host plants. Field observations indicate that host plants are small and scattered in southern Arizona locations. Larvae must therefore find multiple host plants to complete development and face significant challenges in doing so. The frontal projections may thus be an adaptation for finding a scarce resource before starving to death. This is the first evidence that arthropods use projections other than true segmental appendages such as antennae, mouthparts and legs, to locate food resources.

Ectotherms increase in size dramatically during development, and this growth should have substantial effects on their body temperature and ability to thermoregulate. To better understand how this change in size affects temperature, we examined the direct effects of body size on body temperature in Battus philenor caterpillars, and also how body size affects both the expression and effectiveness of thermal refuge-seeking, a thermoregulatory behavior. Field studies of both live caterpillars and physical operative temperature models indicated that caterpillar body temperature increases with body size. The operative temperature models also showed that thermal refuges have a greater cooling effect for larger caterpillars, while a laboratory study found that larger caterpillars seek refuges at a lower temperature. Although the details may vary, similar connections between developmental growth, temperature, and thermoregulation should be common among ectotherms and greatly affect both their development and thermal ecology.

Plasticity in the development and expression of behavior may allow organisms to cope with novel and rapidly changing environments. However, plasticity itself may depend on the developmental experiences of an individual. For instance, individuals reared in complex, enriched environments develop enhanced cognitive abilities as a result of increased synaptic connections and neurogenesis. This suggests that costs associated with behavioral plasticity-in particular, increased investment in "self" at the expense of reproduction-may also be flexible. Using butterflies as a system, this work tests whether allocation of resources changes as a result of experiences in "difficult" environments that require more investment in learning. We contrast allocation of resources among butterflies with experience in environments that vary in the need for learning. Butterflies with experience searching for novel (i.e., red) hosts, or searching in complex non-host environments, allocate more resources (protein and carbohydrate reserves) to their own flight muscle. In addition, butterflies with experience in these more difficult environments allocate more resources per individual offspring (i.e., egg size and/or lipid reserves). This results in a mother's experience having significant effects on the growth of her offspring (i.e., dry mass and wing length). A separate study showed this re-allocation of resources comes at the expense of lifetime fecundity. These results suggest that investment in learning, and associated changes in life history, can be adjusted depending on an individual's current need, and their offspring's future needs, for learning.

The environment experienced during development can dramatically affect the brain, with possible implications for sensory processing, learning, and memory. Although the effects of single sensory modalities on brain development have been repeatedly explored, the additive or interactive effects of multiple modalities have been less thoroughly investigated. We asked how experience with multisensory stimuli affected brain development in the bumblebee Bombus impatiens. First, to establish the timeline of brain development during early adulthood, we estimated regional brain volumes across a range of ages. We discovered significant age-related volume changes in nearly every region of the brain. Next, to determine whether these changes were dependent upon certain environmental stimuli, we manipulated the visual and olfactory stimuli available to newly emerged bumblebee workers in a factorial manner. Newly emerged bumblebees were maintained in the presence or absence of supplemental visual and/or olfactory stimuli for 7 days, after which the volumes of several brain regions were estimated. We found that the volumes of the mushroom body lobes and calyces were larger in the absence of visual stimuli. Additionally, visual deprivation was associated with the expression of larger antennal lobes, the primary olfactory processing regions of the brain. In contrast, exposure to plant-derived olfactory stimuli did not have a significant effect on brain region volumes. This study is the first to explore the separate and interactive effects of visual and olfactory stimuli on bee brain development. Assessing the timing and sensitivity of brain development is a first step toward understanding how different rearing environments differentially affect regional brain volumes in this species. Our findings suggest that environmental factors experienced during the first week of adulthood can modify bumblebee brain development in many subtle ways., (© 2013 S. Karger AG, Basel.)

Floral displays are under selection to both attract pollinators and deter antagonists. Here we show that a common floral trait, a nectar guide pattern, alters the behavior of bees that can act opportunistically as both pollinators and as antagonists. Generally, bees access nectar via the floral limb, transporting pollen through contact with the plant's reproductive structures; however bees sometimes extract nectar from a hole in the side of the flower that they or other floral visitors create. This behavior is called "nectar robbing" because bees may acquire the nectar without transporting pollen. We asked whether the presence of a symmetric floral nectar guide pattern on artificial flowers affected bumble bees' (Bombus impatiens) propensity to rob or access nectar "legitimately." We discovered that nectar guides made legitimate visits more efficient for bees than robbing, and increased the relative frequency of legitimate visits, compared to flowers lacking nectar guides. This study is the first to show that beyond speeding nectar discovery, a nectar guide pattern can influence bees' flower handling in a way that could benefit the plant.

Some animal species increase resource acceptance rates in the presence of conspecifics. Such responses may be adaptive if the presence of conspecifics is a reliable indicator of resource quality. Similarly, these responses could represent an adaptive reduction in choosiness under high levels of scramble competition. Although high resource quality and high levels of scramble competition should both favor increased resource acceptance, the contexts in which the increase occurs should differ. In this paper, we tested the effect of social environment on egg-laying and aggressive behavior in the walnut fly, Rhagoletis juglandis, in multiple contexts to determine whether increased resource acceptance in the presence of conspecifics was better viewed as a response to increased host quality or increased competition. We found that grouped females oviposit more readily than isolated females when provided small (low-quality) artificial hosts but not when provided large (high-quality) artificial hosts, indicating that conspecific presence reduces choosiness. Increased resource acceptance was observed even when exposure to conspecifics was temporally or spatially separate from exposure to the resource. Finally, we found that individuals showed reduced aggression after being housed in groups, as expected under high levels of scramble competition. These results indicate that the pattern of resource acceptance in the presence of conspecifics may be better viewed as a response to increased scramble competition rather than as a response to public information about resource quality.

Plants often attract pollinators with floral displays composed of visual, olfactory, tactile and gustatory stimuli. Since pollinators' responses to each of these stimuli are usually studied independently, the question of why plants produce multi-component floral displays remains relatively unexplored. Here we used signal detection theory to test the hypothesis that complex displays reduce a pollinator's uncertainty about the floral signal. Specifically, we asked whether one component of the floral display, scent, improved a bee's certainty about the value of another component, color hue. We first trained two groups of bumble bees (Bombus impatiens Cresson) to discriminate between rewarding and unrewarding artificial flowers of slightly different hues in the presence vs absence of scent. In a test phase, we presented these bees with a gradient of floral hues and assessed their ability to identify the hue rewarded during training. We interpreted the extent to which bees' preferences were biased away from the unrewarding hue ('peak shift') as an indicator of uncertainty in color discrimination. Our data show that the presence of an olfactory signal reduces uncertainty regarding color: not only was color learning facilitated on scented flowers but also bees showed a lower amount of peak shift in the presence of scent. We explore potential mechanisms by which scent might reduce uncertainty about color, and discuss the broader significance of our results for our understanding of signal evolution.

Testes size often predicts the winner during episodes of sperm competition. However, little is known about the source of nutrients allocated to testes development, or testes plasticity under varying nutrient availability. Among many holometabolous insects, metabolic resources can derive from the larval or adult diet. Distinguishing the source of nutrients allocated to testes can shed light on life history factors (such as maternal influences) that shape the evolution of male reproductive strategies. Here we used an experimental approach to assess resource allocation to testes development in walnut flies (Rhagoletis juglandis) from differing nutritional backgrounds. We fed adult male walnut flies on sugar and yeast diets that contrasted with the larval diet in carbon and nitrogen stable isotope ratios. This design allowed us to assess the dietary source of testes carbon and nitrogen and its change over time. We found significant incorporation of adult dietary carbon into testes, implying that walnut flies are income breeders for carbon (relying more on adult resources). In contrast, we found little evidence that walnut flies incorporate adult dietary nitrogen into testes development. We discuss the implications of these allocation decisions for life history evolution in this species., (Copyright 2010 Elsevier Ltd. All rights reserved.)

Phenotypic plasticity is adaptive in variable environments but, given its costs, may be disfavored if only one environment is commonly encountered. Yet species in relatively constant environments often adjust phenotypes successfully in rare or novel environments. Developmental biases may reduce the costs of plasticity in common environments, favoring the maintenance of plasticity. We explored this proposition by studying the flexibility of visually guided host-selection behavior in cabbage white butterflies (Pieris rapae), wherein common and rare environments consisted of green and red host types, respectively. We demonstrated in greenhouse assays that adult females display an innate bias toward green color during host search but alter that bias through learning in red-host assemblages such that, after several hours of experience, red hosts are located as efficiently as green hosts. Full-sib analyses suggested there was genetic variation in host and color choice that was more pronounced in the red-host environment. We found no evidence of genetic correlations in behavior across host environments or of fitness costs of plasticity in color choice. Our results support the idea that learning may persist in less variable environments through the evolution of innate biases that reduce operating costs in common environments.

The identification of regulatory sequences in animal genomes remains a significant challenge. Comparative genomic methods that use patterns of evolutionary conservation to identify non-coding sequences with regulatory function have yielded many new vertebrate enhancers. However, these methods have not contributed significantly to the identification of regulatory sequences in sequenced invertebrate taxa. We demonstrate here that this differential success, which is often attributed to fundamental differences in the nature of vertebrate and invertebrate regulatory sequences, is instead primarily a product of the relatively small size of sequenced invertebrate genomes. We sequenced and compared loci involved in early embryonic patterning from four species of true fruit flies (family Tephritidae) that have genomes four to six times larger than those of Drosophila melanogaster. Unlike in Drosophila, where virtually all non-coding DNA is highly conserved, blocks of conserved non-coding sequence in tephritids are flanked by large stretches of poorly conserved sequence, similar to what is observed in vertebrate genomes. We tested the activities of nine conserved non-coding sequences flanking the even-skipped gene of the teprhitid Ceratis capitata in transgenic D. melanogaster embryos, six of which drove patterns that recapitulate those of known D. melanogaster enhancers. In contrast, none of the three non-conserved tephritid non-coding sequences that we tested drove expression in D. melanogaster embryos. Based on the landscape of non-coding conservation in tephritids, and our initial success in using conservation in tephritids to identify D. melanogaster regulatory sequences, we suggest that comparison of tephritid genomes may provide a systematic means to annotate the non-coding portion of the D. melanogaster genome. We also propose that large genomes be given more consideration in the selection of species for comparative genomics projects, to provide increased power to detect functional non-coding DNAs and to provide a less biased view of the evolution and function of animal genomes.

The role of brain size as a cost of learning remains enigmatic; the nature and timing of such costs is particularly uncertain. On one hand, comparative studies suggest that congenitally large brains promote better learning and memory. In that case, brain size exacts a global cost that accrues even if learning does not take place; on the other hand, some developmental studies suggest that brains grow with experience, indicating a cost that is induced when learning occurs. The issue of how costs are incurred is an important one, because global costs are expected to constrain the evolution of learning more than would induced costs. We tested whether brain size represented a global and/or an induced cost of learning in the cabbage white butterfly, Pieris rapae. We assayed the ability of full sibling families to learn to locate either green hosts, for which butterflies have an innate search bias, or red hosts, which are more difficult to learn to locate. Naïve butterflies were sacrificed at emergence and congenital brain volume estimated as a measure of global costs; experienced butterflies were sacrificed after learning and change in brain volume estimated as a measure of induced costs. Only for the mushroom body, a brain region involved in learning and memory in other insects, was volume at emergence related to learning or host-finding. Butterfly families that emerged with relatively larger mushroom bodies showed a greater tendency to improve their ability to find red hosts across the two days of host-search. The volume of most brain regions increased with time in a manner suggesting host experience itself was important: first, total number of landings during host-search was positively related to mushroom body calyx volume, and, second, experience with the red host was positively related to mushroom body lobe volume. At the family level, the relative volume of the mushroom body calyx and antennal lobes following learning was positively related to overall success in finding red hosts. Overall, our results suggest that within species, brain size might act as a small global cost of learning, but that environment-specific changes in brain size might reduce the overall costs of neural tissue in the evolution of learning., ((c) 2009 S. Karger AG, Basel.)

Multimodal signals are common in nature and have recently attracted considerable attention. Despite this interest, their function is not well understood. We test the hypothesis that multimodal signals improve decision making in receivers by influencing the speed and the accuracy of their decisions. We trained bumble-bees (Bombus impatiens) to discriminate between artificial flowers that differed either in one modality, visual (specifically, shape) or olfactory, or in two modalities, visual plus olfactory. Bees trained on multimodal flowers learned the rewarding flowers faster than those trained on flowers that differed only in the visual modality and, in extinction trials, visited the previously rewarded flowers at a higher rate than bees trained on unimodal flowers. Overall, bees showed a speed-accuracy trade-off; bees that made slower decisions achieved higher accuracy levels. Foraging on multimodal flowers did not affect the slope of the speed-accuracy relationship, but resulted in a higher intercept, indicating that multimodal signals were associated with consistently higher accuracy across range of decision speeds. Our results suggest that bees make more effective decisions when flowers signal in more than one modality, and confirm the importance of studying signal components together rather than separately.

Prospects of global increases in extreme weather change provide incentive to examine how such change influences animal behavior, for example, behavior associated with resource use. In this study, we examined how oviposition behavior in a southern Arizona population of pipevine swallowtails (Battus philenor L.) responded to changes in their Aristolochia host resource and vegetative background caused by the North American monsoon system. Summer monsoon rains resulted in a flush of non-host vegetation and a more than doubling in rate of landings by host-searching females on non-host vegetation. Rates of discovery of the host species A. watsoni Woot. Standl. decreased by 50% after monsoon rains. Rains did not alter host density appreciably, but resulted in significant increases in host plant size and new growth, two indicators of host suitability for B. philenor larvae. After the rains, mean clutch size on individual host plants increased by a factor of 2.5; the mean proportion of host plants encountered on which a female laid eggs also increased significantly. Females were discriminating about the host plants on which they laid eggs after alightment; plants accepted for oviposition were larger, bore more new growth, and bore fewer larvae than rejected plants. Contrary to predictions from foraging theory, degree of discrimination did not change seasonally. Finally, the rate at which eggs were laid increased seasonally, suggesting that oviposition rates were limited more before monsoon rains by the relatively low quality of hosts than they were after the rains by the relatively low rate at which hosts were found. This latter result suggests that, while butterflies possess behavioral flexibility to respond to extreme weather change, such flexibility may have limits. In particular, expected increases in the severity and frequency of droughts may result in reduced oviposition rates, reductions that could have adverse demographic consequences.

We tested a hypothesis originating with Darwin that bees outside the nest exhibit social learning in flower choices. Naive bumblebees, Bombus impatiens, were allowed to observe trained bees or artificial bees forage from orange or green flowers. Subsequently, observers of bees on green flowers landed more often on green flowers than non-observing controls or observers of models on orange flowers. These results demonstrate that bumblebees can change flower choice by observations of non-nest mates, a novel form of social learning in insects that could provide unique benefits to the colony.

Aposematic, or warning, signals are generally interspecific in form: one species advertises noxiousness to a predator or parasite species. In a study of the pipevine swallowtail butterfly (Battus philenor), we show that a pattern of colouration in the caterpillars that is considered to be aposematic in the context of attack by natural enemies also deters oviposition by conspecific females. In field and laboratory assays, females avoided oviposition on plants bearing live conspecific larvae. Females avoided oviposition on plants bearing artificially constructed models identical to larvae in shape, size and colour pattern. Finally, oviposition on plants harbouring a model bearing the larval colour pattern was reduced relative to plants bearing a leaf-green model, suggesting that the larval colour pattern was essential for avoidance. We discuss how intraspecific and interspecific processes might interact in the evolution of an aposematic signal.

"Peak shift" is a behavioral response bias arising from discrimination learning in which animals display a directional, but limited, preference for or avoidance of unusual stimuli. Its hypothesized evolutionary relevance has been primarily in the realm of aposematic coloration and limited sexual dimorphism. Here, we develop a novel functional approach to peak shift, based on signal detection theory, which characterizes the response bias as arising from uncertainty about stimulus appearance, frequency, and quality. This approach allows the influence of peak shift to be generalized to the evolution of signals in a variety of domains and sensory modalities. The approach is illustrated with a bumblebee (Bombus impatiens) discrimination learning experiment. Bees exhibited peak shift while foraging in an artificial Batesian mimicry system. Changes in flower abundance, color distribution, and visitation reward induced bees to preferentially visit novel flower colors that reduced the risk of flower-type misidentification. Under conditions of signal uncertainty, peak shift results in visitation to rarer, but more easily distinguished, morphological variants of rewarding species in preference to their average morphology. Peak shift is a common and taxonomically widespread phenomenon. This example of the possible role of peak shift in signal evolution can be generalized to other systems in which a signal receiver learns to make choices in situations in which signal variation is linked to the sender's reproductive success.

We examined the effect of learning a new task on the performance of a previously learned task with the same set of visual cues in bumblebees, Bombus impatiens. Previous studies have shown that given a binary choice at each task, bumblebees do not show retroactive interference, or mistakes in the first task, if the two tasks are in different contexts, feeding and nest location. Here we tested whether adding a third, unrewarded choice to each task affects the performance of bees learning in two contexts. In addition, we examined whether workers differ in their expression of interference and learning ability based on size. Performance of workers at a feeder task was degraded by the introduction of training to a second task at the nest entrance. Mistakes at the feeder were biased toward the color cue that was not rewarding in both tasks; suggesting that irrelevant or background stimuli are more prone to decay or forgetting during interference. With respect to interference, we did not find an effect of body size on the amount of interference; however, size was related to how quickly interference occurred. Among individuals showing retroactive interference, larger bees showed interference earlier in phase 2 than did smaller bees. Overall, larger workers learned each task more rapidly than smaller workers. We conclude that the timing of interference is a tradeoff between acquisition of the new task and performance at a previously learned task. Given that foragers in nature tend to be larger than nest workers, we suggest that size-related learning differences be considered as a factor in division of labor between large and small bumblebees.

The oviposition-preference-offspring-performance hypothesis predicts that female insects should prefer to deposit clutches on or in hosts that maximize offspring performance. An important assumption behind this prediction is that female fitness is tightly correlated with the fitness of any one offspring. In this study, we evaluate offspring performance in the walnut fly, Rhagoletis juglandis Cresson (Diptera: Tephritidae), in relation to a previously described oviposition preference for previously exploited host fruit. In particular, we examined how superparasitism of walnut hosts influences offspring survival and weight at pupation under field conditions. We found that superparasitism was common and that increases in larval densities within fruit were associated with reduced larval survival and weight at pupation. In a laboratory experiment, female size was correlated with lifetime fecundity. In this system, oviposition preference is therefore negatively, not positively, correlated with offspring performance. We argue that patterns of female preference in this system reflect direct benefits to females that are traded off against costs in terms of offspring fitness. Because female fitness is a product not only of offspring quality but also of the total number of offspring produced, female walnut flies may be optimizing their fitness by producing many less fecund offspring. Studies examining the preference-performance hypothesis should consider the reproductive conflicts between parents and offspring as potential factors that influence the congruence between parental preference and offspring performance.

The invertebrate neuromodulator octopamine is known to be involved in bees' associative learning, enhancing the responsiveness of a bee to a conditioned stimulus. In this study, we tested the effect of octopamine on the choice behavior of free-flying bumblebees using a two-phase experiment in an array of artificial flowers. During the first phase of the experiment, the bee was allowed to collect octopamine-laden sugar water from two types of equally rewarding flowers (yellow versus blue). In the second phase, one type of flower was set to be unrewarding. The behavior of the bee (proportion of visits to the unrewarding flowers) over the two phases was fitted to a sigmoid regression model. Our results show that octopamine had no significant effect on the bees' equilibrium choice or on the overall rate of the behavioral change in response to the change in reward. Rather, octopamine significantly affected the time interval between the change in reward status and the initiation of behavioral change in the bee.

Oocyte counts, used as a measure of egg load, were compared among three different age groups (15, 30 and 45 days) of two polyphagous species of tephritid fruit flies, Anastrepha ludens and Anastrepha obliqua, which were exposed to varying conditions of diet (sucrose vs sucrose and protein), availability of oviposition substrate (present vs absent), adult female density (1, 2 and 4 females/cage), and semiochemical context (presence vs absence of male pheromones and fruit volatiles). In both species, oocyte counts were higher in older females and for females fed sucrose and protein than for females fed sucrose only. The presence of artificial oviposition substrates influenced oocyte counts in A. obliqua, but not in A. ludens. Female density influenced oocyte counts in both species. Females maintained in groups had higher egg loads than isolated females. Finally, preliminary evidence suggests that semiochemical context influenced oocyte counts. Counts were highest for females in a room containing both fruit volatiles and male pheromone, lowest for females in a room containing neither volatiles nor pheromone, and intermediate for females in rooms containing either volatiles or pheromone but not both. Our results suggest that egg load is influenced by environmental factors in different ways in these two species. Egg load in A. obliqua, a species whose host fruits are highly ephemeral, is responsive to access to the host resource. By contrast, in A. ludens, a species infesting less ephemeral fruit, female density and age played a more important role than host stimuli. The role of ovarian maturation and oviposition in mediating these effects, as well as implications for mass rearing and pest management, are discussed.

Oviposition behavior in herbivorous and frugivorous insects and parasitoids is dynamic at the level of the individual, responding to variation in host quality and availability. Patterns of variation in egg load in response to host presence and quality suggest that ovarian development also responds to variation in the host environment. Ovarian dynamics are mediated by feedback from oviposition, by host feeding, and by sensory input from the host. The last of these mechanisms, host sensory cuing, is known to occur in three major orders and provides strong evidence that ovarian dynamics are adaptive by design. Conditions favoring host effects on ovarian development include trade-offs between egg production and either survival or dispersal, uncertainty in the host environment, and a correlation in host conditions between the time that oogenesis is initiated and the time that eggs are laid. Some host defenses block ovarian development, suggesting that ovarian dynamics in host-specific insects should be viewed from a coevolutionary perspective.

The effects of the interaction between the operational sex ratio (OSR) and a resource (i.e. oviposition site) on mating dynamics have rarely been considered. We examined the effect of the resource presence and its interaction, with the effect of OSR on copulation duration in Rhagoletis juglandis, a tephritid fly species characterized by a resource-defence mating system in which males defend territories on walnut fruit. In this species, copulation duration varies from 30 s to over 1 h and was shown previously to respond strongly to changes in OSR. In the field, short copulations tended to begin and end on fruit, whereas most long copulations generally began on fruit but ended in the foliage, suggesting a possible effect of resource presence on the copulation duration. In laboratory assays of isolated pairs, copulations were significantly shorter in the presence of a surrogate fruit, confirming the effect of resource presence. In another laboratory assay, in which we manipulated OSR independently of resource presence, resource presence and OSR were additive in their effects. Results are discussed in the context of sperm competition theory. Copyright 1999 The Association for the Study of Animal Behaviour.

We studied the function of learning in the parasitoid Leptopilina heterotoma by looking at discrimination of odour stimuli used in foraging for a host. To optimize the rate of encounters with hosts, these parasitoids are expected to assess the extent to which variation in host-substrate odours is reliably associated with variation in the presence of hosts, that is, substrate profitability. Where the association is reliable, parasitoids should attend to variation in odours and discriminate between them; where it is not, they should ignore it. We hypothesized that foraging decisions are based on the completeness of information the animal has about differences in substrate profitabilities. Our laboratory studies showed that discrimination and non-discrimination of odour stimuli are dynamic behavioural decisions that can be related to the degree of substrate variation and to an animal's informational state. In wind-tunnel studies, females learned to discriminate between odours from substrates that were qualitatively different, for example, between odours from apple and pear substrates or between yeast substrates with different C6 compounds added. They did not discriminate when differences were small (e.g. between odours from two apple varieties or between yeast patches with different concentrations of ethyl acetate), unless unrewarding experiences provided evidence of the absence of hosts in one of the substrates. Hence, we suggest that non-discrimination between odour stimuli in L. heterotoma is not a lack of ability to discriminate but a functional decision by the parasitoid. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

This study evaluated a possible fitness advantage, specifically time savings, that might account for an unusual propensity in walnut flies (Rhagoletis spp.) to superparasitize their walnut hosts and to place eggs into existing egg-laying cavities. The first part of this study demonstrated that, in laboratory assays, females of two walnut fly species, R. boycei and R. juglandis, save time when cavities are reused and that in R. juglandis, where it was examined in detail with in vivo staining of eggs, time saving was not an artifact of differences in the size of clutches deposited at new versus existing sites. We further demonstrated that time savings reflected a reduction in the time required to generate the cavity itself. In the second part of the study, we evaluated the possibility that, in the field, time saved by reusing existing cavities is nullified by extra time spent mating associated with a previously described tendency for males to guard these cavities. Field observations of R. juglandis indicated that use of existing sites was, as expected, associated with increased mating. Yet, despite the added time spent mating, in observations of similar length females attempting to lay eggs at existing sites deposited clutches more often than females attempting to lay eggs at new sites. We discuss these results in the context of the more common pattern of superparasitism avoidance observed in host-specific insects.

Oviposition by females of the pipevine swallowtail butterfly,Battus philenor, was stimulated by contact with alcoholic extracts of host foliage.D-(+)-Pinitol was isolated and identified from leaf material of one host species,Aristolochia macrophylla (Aristolochiaceae). In combination with chloroform-soluble components of host leaf material, this compound was comparable to the parent extract in stimulating oviposition.

Systematic quantitative observations of the location and diel pattern of adult Mediterranean fruit fly, Ceratitis capitata (Wiedemann), activities were carried out in an orange grove and surroundings on the island of Chios in Greece. Natural fly foods were assessed for their contribution to fly longevity, fecundity and fertility. There were diel shifts in male and female location. Females required a substantial and varied diet to realize peak fecundity. This diet was acquired away from the primary host, orange. Foraging for food throughout most of the day on fig and non-host foliage (including feeding on bird droppings) as well as on fig fruit and grapes, females dispersed and fed more than males. A diet of grapes alone did not support any fecundity, contributing only to longevity. A diet of figs alone, on the other hand, sustained both longevity and egg production. Bird feces alone supported neither egg production nor longevity. However, when added to a diet of figs, bird feces significantly increased fly fecundity. Throughout most of the day, males aggregated in leks within the inner canopy of the primary host, orange. The arrival here during the warmest hours of the day of receptive females, followed by pair formation, reinforced the lek mating system on host foliage. In the afternoon, females shifted to orange fruit where they suffered from high predation mortality while ovipositing. Soon after, males also shifted to orange fruit, where they attempted matings with non-receptive ovipositing females. Male feeding on fig fruit occurred late in the day, a time when they were least likely to find a mate. Male survival did not differ between the natural diets. Tradeoffs between food consumption, mating success and predator evasion are discussed for each sex and related to fruit fly mating systems.

A brief 2-hr experience with hostDrosophila larvae in artificial apple-yeast or mushroom microhabitats had three effects on the foraging behavior of femaleLeptopilina heterotoma (Hymenoptera: Eucoilidae) parasitoids under field conditions. First, experienced females released at the center of circular arrays of apple-yeast and mushroom baits were more likely to find a microhabitat over the course of a daily census than naive ones. Second, for those females that found a microhabitat, experienced ones found it faster than naive ones (i.e., experience reduced travel times). Third, females experienced with a particular microhabitat were more likely to find that micro-habitat than an alternative one. Learned preferences were retained for at least one day and possibly as many as seven. Results generally did not depend on the host species (D. melanogaster orD. simulans) with which females were given experience. Females tended to arrive at baits upwind of the point of release, suggesting that odor is involved in finding host microhabitats and, in particular, in learning to find them more effectively. The implications of these results for the application of semiochemicals in biological control are discussed briefly.

Numerous authors have suggested that genetic subdivision within a population in a heterogeneous environment is more likely if individuals tend, through prior experience, to breed in the same habitat in which they developed. Under semi-field conditions we demonstrate that prior adult experience alters habitat preference in the apple maggot fly, Rhagoletis pomonella (Tephritidae), a frugivorous parasitic insect thought to have undergone sympatric divergence in host use in historical times. Females exposed to a particular host fruit species - apple (Malus pumila) or hawthorn (Crataegus mollis) - in a field cage oviposited at a higher rate in test fruit of that species than did inexperienced females or females exposed to the other species. Females exposed to a particular host fruit species also tended to remain longer in test trees harboring fruit of that species than did inexperienced females or females exposed to the other species. Prior adult experience thus alters two components of habitat preference in the apple maggot fly: oviposition preference and habitat fidelity. We discuss how these effects of experience on habitat preference should increase the likelihood that individuals mate assortatively and may further increase the likelihood that apple maggot populations become genetically subdivided.

The host-discrimination behavior of the adult female pipevine swallowtail butterfly (Battus philenor) was investigated for two populations, an east Texas population that uses two host species with different leaf shapes and a Virginia montane population that uses one host species with a single leaf shape. While Texas and Virginia females exhibited similar chemotactile responses after landing on various host species, butterflies from each population landed more frequently on certain host species used by that particular population. Despite this difference in searching behavior, Texas and Virginia populations were equally capable of learning to search for the leaf shape of a particular host species in artificial enclosure arrays. In addition, learning of leaf-shape preference was confounded equally when each population was introduced into arrays containing all the host species found in both populations. The lack of differentiation in learning of leaf-shape preference between populations that differ in host breadth and host preference argues against the specific hypothesis that learning of leaf-shape preference represents a local adaptation that permits foraging butterflies in the east Texas population to respond to seasonal changes in the relative quality or abundance of host species that differ in leaf shape. Three hypotheses are proposed to explain the apparent absence of interpopulation genetic variation in learning traits. One hypothesis supposes that learning of leaf-shape discrimination is mediated by the same physiological mechanisms that permit females to learn to discriminate among suitable and unsuitable conspecific plants. Selection for restriction of learning of leaf-shape preference in the Virginia montane population may therefore be constrained by selection for learning of other types of discrimination behavior., (© 1986 The Society for the Study of Evolution.)