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Phytophagous insects may be expected to prefer host-plant species on which their larvae perform best, but this has rarely been explored in grass-feeding butterflies. We explored links between oviposition preferences, larval food preferences, and performance (larval survival and development time, pupal mass, and adult longevity) on 18 species of grass in the common evening brown, Melanitis leda L. (Nymphalidae: Melanitini), a tropical butterfly that has been recorded from a large number of species of grass. Melanitis leda oviposited on all of the grass species offered in choice experiments, and larvae were able to develop to the adult stage on almost all of them. Oviposition preferences were for species of plants on which larval survival was higher, but were not correlated with larval development. The mother's choice of host plant appeared to affect performance of the resulting adults as large butterflies had longer life spans. Ovipositing females appeared to adjust clutch size and position of their eggs on a plant depending on plant traits. Larval survival was only marginally correlated with larval growth rate on the different species of plants. Larvae showed host-plant preferences that were not correlated with adult oviposition preferences, larval survival, or larval growth. Overall, while M. leda can utilize a large number of species of grass, it shows marked oviposition preferences. Apparent fine-tuning of clutch size and positioning of eggs on plants with different traits might be another adaptation associated with using a large number of species of plants. We conclude that this grass feeding butterfly selects oviposition sites that maximize the chances of their larvae surviving.

International audience; Evading predators is a fundamental aspect of the ecology and evolution of all prey animals. In studying the influence of prey traits on predation risk, previous researchers have shown that crypsis reduces attack rates on resting prey, predation risk increases with increased prey activity, and rapid locomotion reduces attack rates and increases chances of surviving predator attacks. However, evidence for these conclusions is nearly always based on observations of selected species under artificial conditions. In nature, it remains unclear how defensive traits such as crypsis, activity levels, and speed influence realized predation risk across species in a community. Whereas direct observations of predator-prey interactions in nature are rare, insight can be gained by quantifying bodily damage caused by failed predator attacks. 2). We quantified how butterfly species traits affect predation risk in nature by determining how defensive traits correlate with wing damage caused by failed predation attempts, thereby providing the first robust multi-species comparative analysis of predator-induced bodily damage in wild animals. 3). For 34 species of fruit-feeding butterflies in an African forest, we recorded wing damage and quantified crypsis, activity levels, and flight speed. We then tested for correlations between damage parameters and species traits using comparative methods that account for measurement error. 4). We detected considerable differences in the extent, location, and symmetry of wing surface loss among species, with smaller differences between sexes. We found that males (but not females) of species that flew faster had substantially less wing surface loss. However, we found no correlation between cryptic colouration and symmetrical wing surface loss across species. In species in which males appeared to be more active than females, males had a lower proportion of symmetrical wing surface loss than females. 5). Our results provide evidence that activity greatly influences the probability of attacks and that flying rapidly is effective for escaping pursuing predators in the wild, but we did not find evidence that cryptic species are less likely to be attacked while at rest.

Information on the mating system of an insect species is necessary to gain insight into sexual selection and population structure. Male territoriality of the common evening brown butterflyMelanitis ledahas been studied in the wild, but other aspects of its mating system remain largely unknown. For a population ofM. ledain South India, we observed male-male and male-female interactions in captivity, measured mating duration and spermatophore mass, and also determined the degree of polyandry in the wild. We found that mating behavior takes place for short periods of time around dawn and dusk. Our observations corroborate that males compete in aerial combats (twirling) and interfere with mating pairs. In the morning, they may use shivering to warm up. Females can twirl with males and refuse mating by pointing their abdomens upwards or by flying away. Males court females by fluttering their wings while perched behind females, and then initiate copulation by curling their abdomens ca. 180 degrees sideways to make genital contact. While in the morning, matings lasted on average one hour and twenty-three minutes and never exceeded three hours, in the evening, matings could be of similar duration, but 42% of butterflies only separated when dawn was approaching. However, such long nocturnal matings did not result in heavier spermatophores. The first spermatophore of a male tended to be larger than subsequent spermatophores. Together with previous studies on this species, our findings suggest that males compete mainly through territorial defense (as reported before), courtship performance, and interference, and to a lesser extent by providing spermatophores, while females exert some control over the mating system by the timing of their receptivity and mate choice.

Specificity is one of the fundamental concepts in ecology. Host specificity of phytophagous insects has been of particular interest because of its crucial role in diversification and life-history evolution. However, the majority of tropical insects remain insufficiently explored with respect to their host-plant relations. A lack of respective data is also hindering the debate over whether higher levels of host-plant specificity prevail in tropical insects compared to temperate ones. We investigated host-plant specificity of forest geometrid moths (Lepidoptera: Geometridae) in equatorial Africa using host-plant acceptability trials with neonate larvae, with the addition of field observations. We compare our experimental data to the (well-known) host-specificity patterns of closely related temperate (hemiboreal) species. Similarly to the temperate region, there were broadly polyphagous tropical species in several clades of Geometridae utilizing hosts belonging to different plant families. Phylogenetic comparative analysis returned no significant differences in host specificity between the two regions. Our study contributes to the evidence that host-plant specificity of herbivores is not necessarily substantially higher in tropical than temperate regions.

In warm climates, lizards may be important as predators of butterflies and thus exert selection pressure on butterfly colour patterns, including eyespots. However, this has received little attention. Two studies reported no evidence that butterfly eyespots deflect lizard attacks, while possible intimidating effects of butterfly eyespots on lizards have not been investigated. We exposed common evening brown butterflies (Melanitis leda) with a wide range of eyespot sizes (dry season forms with very small and faint eyespots, intermediate phenotypes with small eyespots, and wet season forms with large or very large eyespots) to house geckos, and recorded the location and shape of wing surface loss. We supplemented these data with direct observations of attacks of house geckos (Hemidactylus frenatus) and bronze skinks (Eutropis macularia) on common evening brown butterflies. We found that about three quarters of symmetrical wing surface loss, and about half of the non-symmetrical wing surface loss was rounded like the mouth of a gecko. Symmetrical as well as non-symmetrical wing surface loss was significantly more common in hindwings. We surmise that rounded wing surface loss and a bias in symmetrical wing surface loss to hindwings can be taken as evidence for failed lizard attacks. Rounded wing surface loss was more common in butterflies with small eyespots than in those with very small eyespots or with large eyespots. This suggests that small eyespots can deflect lizard attacks away from the head towards the wing margins, and that large eyespots can thwart lizard attacks. In four cases, skinks appeared to explore rather than attack butterfly eyespots. In contrast to earlier studies, our data provide evidence that butterfly eyespots can be an effective defence against lizards, and thus that predation by lizards can select for eyespots in butterflies.

Reproductive data of individual insects are extremely hard to collect under natural conditions, thus the study of research questions related to oviposition has not advanced. Patterns of oviposition are often inferred only indirectly, through monitoring of host infestation, whereas the influence of age structure and several other factors on oviposition remains unknown. Using a new approach, in this article, we live-trapped wild Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) females on the Greek island of Chios during two field seasons. For their remaining lifetime, these females were placed individually in small cages and their daily oviposition was monitored. Reproduction rates between cohorts from different collection dates were then compared. The results showed that in the different captive cohorts the average remaining lifetime and reproduction were highly variable within and between seasons. Multivariate regression analysis showed that the month of capture had a significant effect on captive life span, average daily reproduction, and patterns of egg laying. The effect of year was significant on reproduction, but not on captive life span. These differences between sampling periods probably reflect differences in the availability of hosts and other factors that vary during the season and affect age structure and reproduction. Using a non-parametric generalized additive model, we found a statistically significant correlation between the captive life span and the average daily reproduction. These findings and the experimental approach have several important implications.

Testosterone can be expected to play a significant role in mediating behavior and life history in social animals, but the number of species with data from the wild is still too small to make generalizations. We investigated the influence of social factors (social status, aggression, and reproductive females) and environmental variation (rainfall and temperature) on fecal testosterone concentrations in wild male gray-cheeked mangabeys (Lophocebus albigena) living in five groups in Kibale National Park, Uganda. This species is phylogenetically closely related to baboons, but is arboreal, with multi-male, multi-female groups rarely exceeding 20 individuals. We analyzed 358 fecal samples from 21 adult males. We found that the highest-ranking males had the highest testosterone concentrations while immigrant males had the lowest testosterone concentrations. Aggression was not correlated with testosterone levels. The presence of females with sexual swellings at their most tumescent stage increased testosterone concentrations in all males. Finally, individuals tended to have lower testosterone when the temperature was higher.

We use unique experimental data on daily reproduction and survival of individual fruit flies from eight cohorts eclosed at different dates in 2004 and 2005 who were treated with varying proportions of sugar and yeast and subject to different caloric restrictions (CR). We investigate the relationship between eclosion date and longevity across diets and reproduction in Anastrepha ludens. We show that eclosion date can be associated with uncontrolled external or internal factor(s) which can modulate longevity of males and females independently of diet and reproduction to the extent similar to the effect of diet on longevity. The effect of diet manipulation on longevity is sensitive to date of eclosion with the role of CR in life extension ranging from beneficial to harmful. Interaction of date of eclosion with compositional changes of sugar and yeast but not with CR is responsible for life extension. Highly protein-enriched diets reliably maximize reproduction but not life span. Decreased longevity of flies treated with high-protein diets may be associated with harmful consequences of protein ingestion but is unlikely a result of high reproduction rates. We present evidence for the presence of two frailty-sensitive weakly interacting mechanisms of longevity in female flies associated with differences in predisposed fitness.

Male life history and resource allocation is not frequently studied in aging and life span research. Here, we verify that males of long-lived fruit-feeding butterfly species have reduced longevity on restricted diets [Beck, J., 2007. The importance of amino acids in the adult diet of male tropical rainforest butterflies. Oecologia 151, 741-747], in contrast to the common finding of longevity extension in dietary restriction experiments in Drosophila and some other organisms. Males of some of the most long-lived species of fruit-feeding butterflies were collected from Kibale Forest, Uganda, and kept on diets of either sugar or mashed banana. Seven out of eight species had non-significantly longer life spans on mashed banana diets. Data analysis using a time-varying Cox-model with species as covariate showed that males had reduced survival on the sugar diet during the first 35 days of captive life, but the effect was absent or reversed at more advanced ages. These results challenge the generality of dietary restriction as a way to extend life span in animals. We argue that such studies on males are promising tools for better understanding life history evolution and aging because males display a wider variety of tactics for obtaining reproductive success than females.

Fruit-feeding butterflies are among the longest lived Lepidoptera. While the use of pollen-derived amino acids by Heliconius butterflies has been interpreted as important for the evolution of extended lifespans, very little is known about the life-history consequences of frugivory. This issue is addressed by investigating effects of four adult diets (sugar, sugar with amino acids, banana, and moistened banana) on lifespan and reproduction in the fruit-feeding butterfly Charaxes fulvescens Aurivillius (Lepidoptera: Nymphalidae). Female butterflies were collected from Kibale National Park, Uganda, and kept individually in cages near their natural habitat and data were collected on lifespan, oviposition, and hatching of eggs. Lifespan in captivity was longer for the sugar and the amino acid cohort, than for the banana cohorts. The longitudinal pattern of oviposition was erratic, with many days without oviposition and few periods with high numbers of eggs laid. Butterflies typically did not lay eggs during their 1st week in captivity and the length of the period between capture and first reproduction was significantly shorter for butterflies fed moistened banana. The length of the reproduction period (first reproduction–last reproduction in captivity) and the reproduction rate (total number of eggs/length of the reproduction period) did not differ significantly between the diet treatments. Those fed with amino acid and moistened banana had significantly higher egg hatchability than those fed with sugar and banana. We found no evidence for a lifespan cost of reproduction. Our results show that (1) female C. fulvescens can use amino acids in their diet for laying fertile eggs, (2) more wing-wear does correlate with lower survival in captivity (indicating aging in the wild), but not with intensity of reproduction (providing no evidence for reproductive aging), and (3) fruit-feeding butterflies may be dietary restricted in the field.

International audience; Intestinal parasites may constitute an important evolutionary and ecological force. We aimed to identify social, physiological, and environmental factors that correlate with intestinal parasite infections in adult male gray-cheeked mangabeys (Lophocebus albigena). We analyzed 102 fecal samples collected from 18 adult males over 19 mo for the incidence (proportions of samples with parasites) and intensity (total number of parasites per gram of feces) of infection of nematodes relative to social status, fecal glucocorticoid and testosterone metabolites, group size, and rainfall, all of which are factors that earlier studies suggested can be important mediators of parasite load. Parasite incidence was greater in immigrant males compared to low- and high-ranking males whereas parasite intensity was greater in immigrant males and low-ranking males compared to high-ranking males. Fecal samples with more parasites had higher concentrations of fecal glucocorticoid and testosterone metabolites than those with fewer parasites. As immigrant males had a greater incidence of parasites and higher concentrations of both metabolites than resident males, this profile appears to fit immigrant males best. We also found higher nematode intensities in mid-sized groups and during periods with more rainfall. Our results suggest that it will be fruitful for future studies to explore the role of immigrant males as spreaders of intestinal parasites.

Adult butterflies feed from a variety of substrates and have appropriate adaptations. We examined proboscis morphology in a community of fruit-feeding butterflies (Nymphalidae) in a tropical forest in Uganda. These data were supplemented with behavioural observations and measurements of intake rate on natural and artificial substrates. We found no sexual dimorphism in proboscis morphology even though puddling behaviour is usually performed by males. Two main feeding techniques could be distinguished on the basis of behaviour and morphology: the piercing technique, typically found in Charaxinae, and the sweeping technique employed by both Nymphalinae and Satyrinae. These techniques, distinguished in previous studies, are described in more detail and their relative efficiencies are discussed in the context of sexual dimorphism, food-choice and life history evolution. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86, 333–343.

International audience; An elevated concentration of glucocorticoids is an indicator of stress, and chronically high glucocorticoid levels are often associated with poor health and reduced fertility.We explored conditions that might be stressors in the lives of adult female gray-cheeked mangabeys (Lophocebus albigena) by measuring fecal glucocorticoid metabolites (fGCM) concentrations. During a six-month study we collected 109 fecal samples from 28 adult females from five groups in Kibale National Park, Uganda. We examined fecal fGCM levels of individual females relative to their own reproductive status (cycling or not cycling) and that of other females and to the presence of newcomer (immigrant) males. We found elevated fGCM concentrations in females when other females in their groups were at the peak of sexual swelling, and when immigrant males joined the group.

The males of butterflies transfer a spermatophore to the female during mating that can contain nutrients enhancing the reproductive potential of their partners. The nutrients transferred by males can be derived from both larval and adult feeding. These nutrients may be depleted by multiple matings. An apparent difference in adult feeding behaviour between the sexes is puddling on mud, dung and carrion, which in most butterfly species is exclusively a male behaviour. A possible explanation for this division in feeding behaviour is that nutrients derived from puddling by males are transferred to the female during mating. Here, we test this hypothesis in the African fruit-feeding butterfly Bicyclus anynana. We varied the male nuptial gift by (1) feeding males either a diet with or without sodium, and (2) varying the number of previous successful copulations by remating males up to five times on consecutive days. The results show both a strong effect of order of mating on the mating duration, and an individual effect with some males typically copulating for a shorter time than others. The effects on female reproduction were, however, minimal. The total number of eggs per female and the sodium content of the eggs did not differ significantly between diets, nor were they affected by the mating histories of the males. Eggs showed a non-significant lower hatching for females partnered by a male who had already mated several times. There was an indication of an interaction with male diet: the sodium treatment showing a decline in egg hatchability with order number of male mating, whilst the control treatment showed a constant hatchability. The results are discussed in relation to determinants of male gift-giving strategy and to other potential explanations for a restriction of puddling to males in butterflies.