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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.

Phenotypic plasticity in heterogeneous environments can provide tight environment-phenotype matching. However, the prerequisite is a reliable environmental cue(s) that enables organisms to use current environmental information to induce the development of a phenotype with high fitness in a forthcoming environment. Here, we quantify predictability in the timing of precipitation and temperature change to examine how this is associated with seasonal polyphenism in tropical Mycalesina butterflies. Seasonal precipitation in the tropics typically results in distinct selective environments, the wet and dry seasons, and changes in temperature can be a major environmental cue. We sampled communities of Mycalesina butterflies from two seasonal locations and one aseasonal location. Quantifying environmental predictability using wavelet analysis and Colwell's indices confirmed a strong periodicity of precipitation over a 12-month period at both seasonal locations compared to the aseasonal one. However, temperature seasonality and periodicity differed between the two seasonal locations. We further show that: (a) most females from both seasonal locations synchronize their reproduction with the seasons by breeding in the wet season but arresting reproduction in the dry season. In contrast, all species breed throughout the year in the aseasonal location and (b) species from the seasonal locations, but not those from the aseasonal location, exhibited polyphenism in wing pattern traits (eyespot size). We conclude that seasonal precipitation and its predictability are primary factors shaping the evolution of polyphenism in Mycalesina butterflies, and populations or species secondarily evolve local adaptations for cue use that depend on the local variation in the environment.

International audience; Resource use by consumers across patches is often proportional to the quantity or quality of the resource within these patches. In folivores, such proportional use of resources is likely to be more efficient when plants are spatially proximate, such as trees forming a forest canopy. However, resources provided by forest-trees are often not used proportionally. We hypothesised that proportional use of resources is reduced when host trees are isolated among phylogenetically distant neighbours that mask olfactory and visual search cues, and reduce folivore movement between trees. Such phylogenetically distant neighbourhoods might sort out species that are specialists, poor dispersers, or have poor access to information about leaf quality. We studied individual oaks, their leaf size and quality, their folivory and abundance of folivores (mostly Lepidopteran ectophages, gallers and miners), and parasitism of folivores. We found that leaf consumption by ectophages hardly increased with increasing leaf size when host trees were phylogenetically isolated. We found a similar effect on host use by parasitoids in 1 year. In contrast, we found no consistent effects in other folivore guilds. Relative abundances of specialists and species with wingless females declined with phylogenetic isolation. However, resource use within each of these groups was inconsistently affected by phylogenetic isolation. We suggest that phylogenetic isolation prevents ectophages from effectively choosing trees with abundant resources, and also sorts out species likely to recruit in situ on their host tree. Trees in phylogenetically distant neighbourhoods may be selected for larger leaves and greater reliance on induced defences.

If the evolutionary determinants of longevity were mainly extrinsic, related species from different habitats should systematically differ in individual lifespans. Comparative studies of insects can significantly contribute to understanding the evolution of lifespan, as the trait can feasibly be measured in a high number of species. We recorded adult longevities for 110 species of geometrid moths from a tropical community. Comparative analyses based on an original phylogenetic reconstruction were applied to reveal the correlates of species-specific values of lifespan. Larger moth species were found to live longer, and females tended to have shorter lifespans than males. Average adult lifespans in tropical geometrids, and the relationships of lifespan with other variables, were found to be highly similar to those reported for their temperate region relatives. The among-region similarity leads to the conclusion of the dominance of intrinsic (physiological) determinants of longevity over the extrinsic (ecological) ones: the contrasting environments of tropical and temperate forests have not produced detectable differences in moth longevities.

1. Comparative studies on insect life histories are facilitated by the increasing availability of reliable phylogenies but are hampered by the scarcity of comparable data. Fortunately, morphological proxies of some life‐history traits can be measured on preserved specimens. 2. This study compared values of size‐related life‐history traits among a tropical (Ugandan) and a temperate (Estonian) assemblage of geometrid moths. 3. A comparative analysis based on an originally derived phylogeny revealed that tropical moths were, on average, larger than temperate ones. Tropical moths also had somewhat lower relative abdomen masses than temperate ones. This indicates that the tropical rather than the temperate moths tend to use an income (rather than capital) breeding strategy. Nevertheless, no difference was found in a related index of pro‐ovigeny. When body size was accounted for, tropical moths were found to lay smaller eggs than temperate ones. 4. The differences between the two compared areas are consistent with selection on higher mobility of the moths imposed by the more diverse tropical vegetation. Relatively larger eggs of temperate moths may constitute an adaptation to overcome the presumably stronger quantitative defences of their host plants. 5. Overall, however, we conclude that the differences in ecologically relevant size‐related traits are relatively low among moth assemblages of a tropical and a temperate forest region, indicating that these environments may not impose radically different selective pressures on insect life histories.

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.

Sexual traits are often the most divergent characters among closely related species, suggesting an important role of sexual traits in speciation. However, to prove this, we need to show that sexual trait differences accumulate before or during the speciation process, rather than being a consequence of it. Here, we contrast patterns of divergence among putative male sex pheromone (pMSP) composition and the genetic structure inferred from variation in the mitochondrial cytochrome oxidase 1 and nuclear CAD loci in the African butterfly Bicyclus anynana (Butler, 1879) to determine whether the evolution of “pheromonal dialects” occurs before or after the differentiation process. We observed differences in abundance of some shared pMSP components as well as differences in the composition of the pMSP among B. anynana populations. In addition, B. anynana individuals from Kenya displayed differences in the pMSP composition within a single population that appeared not associated with genetic differences. These differences in pMSP composition both between and within B. anynana populations were as large as those found between different Bicyclus species. Our results suggest that “pheromonal dialects” evolved within and among populations of B. anynana and may therefore act as precursors of an ongoing speciation process.

International audience; Aim Under spatial isolation on oceanic islands, species tend to show extreme body sizes. From the point of view of many colonizers, individual hosts surrounded by phylogenetically distant neighbours are phylogenetically isolated. This study addresses for the first time how phylogenetic isolation of individual hosts affects body size of colonizers, and whether effects on body size reflect selection among colonizers established on host individuals rather than selection among colonizers dispersing toward trees or phenotypic plasticity of colonizers. Location Rennes National Forest, Western France. Methods We sampled click beetles (Elateridae) on individual oak trees varying in phylogenetic isolation from their neighbours and in age. We measured body size and fluctuating asymmetry (which we found to correlate to reduced body size) and related both to phylogenetic isolation and age of trees. We compared these relationships among species of different larval trophic position and adult body size, using meta-analytical approaches. Results Within species, body size changes with phylogenetic isolation of individual host trees: root feeders tend to become larger, predators smaller. Effects were independent of mean body-size, disappeared with tree age, and were inconsistent with patterns of fluctuating asymmetry. Main conclusions Our results are consistent with body-size selection among colonizers established on individual trees, rather than selection among colonizers dispersing toward trees or phenotypic plasticity. Overall, phenotype patterns of animals across islands in the ocean may resemble those across host individuals in a phylogenetically distant neighbourhood, suggesting micro-evolution of colonizers in response to the macro-evolutionary structure of the host community.

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.

International audience; In communities of tropical insects, adult abundance tends to fluctuate widely, perhaps in part owing to predator–prey dynamics. Yet, temporal patterns of attack rates in tropical forest habitats have not been studied systematically; the identity of predators of insects in tropical forests is poorly known; and their responses to temporal variation in prey abundance have rarely been explored. We recorded incidence and shape of marks of attacks on dummy caterpillars (proxy of predation rate) in a sub-montane tropical forest in Uganda during a yearlong experiment, and explored correlations with inferred caterpillar abundance. Applying the highest and lowest observed daily attack rates on clay dummies over a realistic duration of the larval stage of butterflies, indicates that the temporal variation in attack rate could cause more than 10-fold temporal variation in caterpillar survival. Inferred predators were almost exclusively invertebrates, and beak marks of birds were very scarce. Attack rates by wasps varied more over time than those of ants. Attack rates on dummies peaked during the two wet seasons, and appeared congruent with inferred peaks in caterpillar density. This suggests (1) a functional response (predators shifting to more abundant resource) or adaptive timed phenology (predators timing activity or breeding to coincide with seasonal peaks in prey abundance) of predators, rather than a numerical response (predator populations increasing following peaks in prey abundance); and (2) that predation would dampen abundance fluctuations of tropical Lepidoptera communities

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.

Although the general concept of capital vs. income breeding has become widely used in insect ecology, finding easy-to-measure indices for quantifying the role of larval- vs. adult-derived nutrients in egg production has remained a challenge. When searching for possible candidates for this task, we evaluated the applicability of three morphometric ratios to be measured on freshly eclosed adults: (1) relative size of female abdomen, (2) sexual size dimorphism, and (3) proportion of volume of mature eggs relative to total volume of female abdomen. We report the values of these indices in five species of geometrid moths [Hypomecis punctinalis Scopoli, Ematurga atomaria L., Selenia tetralunaria Hufnagel, Semiothisa clathrata L., and Epirrhoe alternata Muller (all Lepidoptera: Geometridae)] and compare them to the degrees of income breeding measured directly by oviposition experiments. As a comparison, we also recorded the ovigeny index and the effect of income on lifespan. All morphometric indices varied considerably between the species studied, and, as predicted, the values of all three proposed indices were higher in the more capital-breeding species. The only exception to this pattern was a low proportion of mature eggs in abdomens in one primarily capital-breeding species, S. tetralunaria. Unlike other capital breeders, this species also proved to be largely synovigenic, indicating that capital breeding and pro-ovigeny are not strictly linked. The reproductive traits measured concord with ecological niches occupied by the species studied: the oligophagous meadow dwellers were largely income breeding and synovigenic, whereas the polyphagous forest dwellers were capital breeders. The high and predictable among-species variation in egg-production strategies suggests that geometrid moths form a promising target group for comparative studies on respective traits. Simple anatomical indices, in turn, appear to be applicable as easy-to-record proxies of egg production strategies.

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.

In polygynous insect species, male reproductive success is directly related to lifetime mating success. However, the costs for males of sexual activities such as courting, signaling, and mating are largely unknown. We studied the cost of sexual activities in male Mediterranean fruit flies, Ceratitis capitata (Tephritidae), a polygynous lekking species, by keeping cohorts of individual male flies under relaxed crowding conditions in the laboratory. We used 5 cohorts among which individuals differed in their opportunities to interact with con-specifics and recorded life span, and in one treatment, mating rate. We found that males kept singly lived more than twice as long as males that interacted intensively with mature virgin females, while male–male interactions caused a smaller reduction in longevity. Because longevity of males that could court but not mate was not significantly different from those that could court and mate, we conclude that courting (not mating) was responsible for the observed longevity reduction. Moreover, we detected high variability in male mating success, when 5 virgin females were offered daily. In contrast to the cohort level, individual males that mated at a high rate lived relatively long, thus indicating heterogeneity in quality or sexual strategy among males.

The term 'puddling' includes feeding on (dried) mud and various excrements and secretions of vertebrates, and carrion. It is thought to be a form of supplementary feeding, not targeted at obtaining energy. Although the natural history of the puddling phenomenon in herbivorous arthropods becomes better known, it is still largely unclear how puddling (in particular for sodium) affects fitness despite the growing knowledge of insect physiology at the cellular level. If we follow the definition used for puddling in Lepidoptera, representatives of a wide range of herbivorous and detrivorous terrestrial arthropods (Lepidoptera, Orthoptera, Blattodea, Hymenoptera, Hemiptera, Diptera, and Diplopoda) have been observed to puddle. It appears that those species with diets low in sodium (e.g., folivorous larvae) puddle for sodium whereas those with diets low in nitrogen (e.g., detritivores) puddle for nitrogen. Sex differentials in puddling behavior can usually be explained by transfers of nutrients from males to females during mating. Puddling is rare or absent in immature stages and there is some evidence that nutrients from puddles increase female reproductive success. Strong evidence for the widely cited hypothesis that sodium from puddles is used to enhance neuromuscular activity is still lacking. High mobility and long life spans could be associated with puddling behavior, whereas insects that are concealed or well defended are less likely to puddle (e.g., beetles). The role that risks of pathogen and parasite infection as well as predation at puddling substrates may play in the evolution of puddling remains virtually unexplored.

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.

This study was concerned with the impact of different levels of artificial impairment (leg amputations) on male and female survival and female reproduction in the Mexican fruit fly, Anastrepha ludens . We monitored the demographic responses in a total of 100 flies of each sex that were maintained individually in 4 × 4 × 10 cm cages and subject to 1-of-11 different leg amputations (plus intact control) including cohorts in which either one front, one middle or one rear leg was severed (3 cohorts total), in which two legs were severed in different front–middle–rear combinations (6 cohorts total), or in which the two middle and one additional leg were severed (2 cohorts total). The two main findings were that: (i) although the effects on mortality of impairments were sex-specific, no universal patterns emerged that applied to either sex; and (ii) reproduction occurred in all cohorts of impaired females. Moderately-impaired flies (e.g. amputation of a single middle leg) laid nearly as many eggs in their lifetime as did intact controls. However, severely impaired flies (i.e. 3 legs amputated) laid significantly fewer eggs.

Fruit-feeding butterflies can experience a more nutrient rich adult diet than nectar-feeding species, and can be expected to use these nutrients for egg production. Here we compare life span, and reproduction parameters of wild-caught females of large and long-lived species on either a sucrose or a mashed banana diet. With small sample sizes per species, but rich longitudinal data for each individual, we examined the longitudinal reproduction pattern, egg size and hatchability of these butterflies in captivity. Diet significantly affected mortality in captivity in a time-dependent manner. On average, we found that butterflies fed mashed banana laid 1.855 times more eggs than those fed sugar. They laid significantly more eggs when they laid and conserved egg size with age while butterflies fed sucrose showed significantly declining egg sizes. Egg hatchability was not significantly affected by diet. Long pre-oviposition periods, significantly smaller first eggs, and absence of age at capture effects on intensity of reproduction indicate low reproduction rates in the field that are due to low food availability. With our small sample sizes, we did not detect significant differences between the species in their response to the diet treatments.

Animals in the wild often have physical impairments that can affect their fitness. The aim of this study was to compare injuries and impairments of four different primate species (black-and-white colobus, red colobus, red-tailed monkeys, and grey-cheeked mangabeys) living in Kibale National Park (Uganda), and estimate the proportion of injured and impaired individuals among the sexes and age classes. The species differed in the proportion of individuals with injuries and impairments, with 16.7% in black-and-white colobus, 23.1% in red colobus, 16.2% in red-tailed monkey and 30.2% in grey-cheeked mangabeys. Species also differed in the types and location on the body of injuries. Adult animals had more injuries than immatures. Males had more injuries than females, in all but red-tailed monkeys. The results are discussed in relation to the literature on aggressive behavior, locomotion, and predation in these species.

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.

Reproductive tactics of males can change with individual quality, relat-edness and social structure. Here we describe the behaviours of malegrey-cheeked mangabeys towards other males, and females and theiroffspring (Lophocebus albigena) in relation to male status (high-rank-ing⁄low-ranking⁄transient) and group composition in Kibale NationalPark, Uganda. High-ranking males had the highest mating success, fre-quency of loud calls, mate guarding and aggression towards females andmales. Only transient males were often observed to be aggressivetowards juveniles, while some high-ranking males provided infant care.Mating tactics of high-ranking males varied greatly among the five stud-ied groups, probably as a function of the intensity of male–male compe-tition. These results are discussed with regard to the role of male–malecompetition and behaviours that could affect female mate choice as tac-tics to obtain reproductive success. Ethology Ethology 114 (2008) 851–862 a 2008 The AuthorsJournal compilation a 2008 Blackwell Verlag, Berlin

Female sexual strategies affect male strategies and can play an important role in shaping mating systems. We investigated female sexual behaviour within five groups of grey-cheeked mangabeys in Kibale National Park, Uganda, and tested the hypothesis that females exhibit mate choice using as indications the prevalence of (1) females soliciting matings by presenting to males and (2) females refusing to mate with approaching males. In addition, we describe how these behaviours as well as grooming and copulation calls are distributed over high-ranking, low-ranking and migrating males and discuss these patterns with regard to trade-offs that could play a roll in female mate choice in multi-male groups. Females were promiscuous and initiated almost half of the matings, with both resident and migrating males. More than half of male mating approaches were refused by peak females. Female mate choice in this species may depend on individual female preferences, oestrus phase and male tactic.

Crop-raiding is a significant problem in small-scale farming, both for the people who depend on their crops and the success of conservation projects. Here, we present the results of a survey on subsistence agriculture, describing crop damage and methods to prevent them, in three communities (35 households) at the northern periphery of the Dja Faunal Reserve, in Cameroon. A total of 96.7% of fields experienced crop damage, 13 species were responsible for this damage, and small mammals, such as cane rat, were the most severe (73% fields affected). None of the methods to prevent damage to crops was negatively correlated with damage. Cumulative damage was negatively correlated with the distance from the forest only on the peanut fields. Distance of a field to the forest was negatively correlated with crop-raiding by only one of the 13 species: the agile mangabeys. Species that were the subject of conservation efforts were responsible for a small proportion of the damage.

Information on the life span of organisms in the field is essential for elucidating the evolution of life span and aging. We present mark-recapture data (>30 000 marked individuals, >4000 recaptured at least once) on forty-seven species of fruit-feeding butterflies in a tropical forest in Uganda. The data reveal adult life spans in the field for several species that are significantly longer than previously recorded in Lepidoptera (butterflies and moths). Longevity records for species of which more than 100 individuals were recaptured ranged from 67 (Bicyclus auricruda) to 293 days (Euphaedra medon). In contrast to the majority of Lepidoptera which are short-lived, these all show exceptionally long life spans, and may thus help to better identify factors that affect aging, particularly when combined with information on temporal patterns in reproduction, strategies to avoid predation, and nutritional ecology. These key traits are readily measurable in butterflies and thus studies on fruit-feeding butterflies have much potential for gaining insight into the evolution of life span and aging, especially given the tradition of field-research on butterflies.

International audience; Predation can be one of the key factors that determine abundance in insect herbivore communities, and drive evolution of body size, and anti-predator traits, including crypsis. Population dynamics and selection pressures will depend on the identity of dominant predators in the system, and these may vary substantially among habitats. Arthropods emerge as chief predators on caterpillars in the understorey of non-montane tropical forest, whereas birds dominate elsewhere. In a tropical forest in Uganda, Africa, we evaluated marks on dummy caterpillars that differed in size, material (clay vs. dough), colourant, and plant species on which dummy caterpillars were exposed. We included live caterpillars to estimate the extent to which studies using artificial caterpillars reflect actual levels of predation. Ants and wasps were the most important damagers of dummy caterpillars, whereas bug and beetle damage was very rare, and no bird or small mammal damage was observed. Daily attack rates did not differ significantly from apparent mortality of live caterpillars (daily mortality = 12.1%), but dummy caterpillars made from dough were attacked more frequently (daily attack rate = 18.4%) than those from clay (daily attack rate = 6.9%). Caterpillars of different colour and size, and caterpillars exposed on different plant species had the same chances to be predated. This is in contrast to results from temperate area studies where birds dominate and are not affected by dummy caterpillar material, but prefer larger caterpillars. Our results are consistent with dominant predators on tropical forest caterpillars being invertebrates that are more chemically than visually oriented, so that: (1) material used for dummy caterpillars is important, (2) background matching is relatively unimportant, and (3) being large may have less of a cost. These patterns in predation might facilitate polyphagy and evolution of large body size in tropical Lepidoptera

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.

International audience; Identifying the causes of differential reproductive success is key to understanding natural selection and the forces of selection operating on animals. Here we present results from a 9-yr (2004–2012) study of female reproductive performance in relation to mother’s age and rank, presence of immigrant males, rainfall, and fig fruit abundance in four groups of gray-cheeked mangabeys (Lophocebus albigena) in Kibale National Park, Uganda. We found that females had a rank- and age-specific reproductive pattern, with high-ranking females maturing earlier, having their first births earlier, and exhibiting significantly slower reproductive aging than low-ranking females. We also found that both immigrant and resident males were associated with higher birth rates. Finally, we found that reproduction was aseasonal and did not correlate with rainfall, but that births correlated positively with the abundance of fruits of Ficus spp. Our results show broad similarities between arboreal, forest-dwelling gray-cheeked mangabeys and their more terrestrial, open habitat-dwelling papionin relatives in the importance of dominance rank in estrous cycle initiation, first reproduction, and reproductive aging.

Many butterflies in tropical forests feed on fruits that have fallen to the forest floor. This substrate differs in many ways from floral nectar, and therefore fruit-feeding butterflies are expected to possess adaptations for efficient foraging, choice, and ingestion of their food. Differences in food quality and in spatial and temporal availability are also likely to have led to life history evolution. Here we describe the sugar and nitrogen content of fruits that butterflies feed on in at ropical forest in Uganda, and measure the attractiveness of these fruits to the local butterfly fauna together with the role that decay plays in the attraction. These data are supplemented with feeding observations at fruit falls in the forest. Our results show that (1) fruits contain significant and variable concentrations of sugar and nitrogen, and constitute a nutritious food source for butterflies in tropical forests; (2) fruit-feeding butterflies use cues from the fruits and fermentation products to locate their food; (3) different classes of fruit-feeding butterflies may vary in their preferences for certain fruits, and differ in their ability to find preferred food; an d( 4) fruit choice is not strongly correlated with attractiveness or nutrient content. The results are discussed in the light of the evolution of food searching and life history strategies.

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; Differences among females in infant survival can contribute substantially to variance in fitness. Infant survival is a product of external risk factors and investment by kin, especially the mother, and is thus closely tied with the evolution of behavior and life history. Here we present a 9-yr study (2004–2012) of infant survival and sex ratio relative to age and dominance ranks of mothers and the presence of immigrant males in a free-ranging population of gray-cheeked mangabeys (Lophocebus albigena) in Kibale National Park, Uganda.We consider immigrant males because they are known to increase infant mortality in several other species.We found that infants of older mothers had higher survival than those of younger mothers but that high rank did not confer a significant benefit on infant survival. Female infants had higher survival than male infants. Young, low-ranking females had more male infants than young, high-ranking females, which had slightly more daughters, but this difference declined as females aged because low-ranking females had more daughters as they aged. With limited data, we found a significantrelationship between the presence of male immigrants and infant mortality (falls and unexplained disappearances) to 18 mo. Our results suggest that infant survival in graycheeked mangabeys is most precarious when mothers must allocate energy to their own growth as well as to their infants, that sons of young mothers are at greatest risk, and that immigrant males can negatively affect infant survival.

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.

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.

Because tropical forests face serious threats and are usually situated in developing countries, cheap and easy Rapid Biodiversity Assessment (RBA) seems an ideal strategy for documenting changes in habitat quality and setting conservation priorities. Fruit-feeding butterflies are often used as focal taxa to document biodiversity trends in tropical forests. However, the choice of inappropriate sampling strategies may nullify its value. More extensive data sets that sample multiple months and seasons and incorporate the vertical dimension are needed as a baseline for designing reliable sampling regimes. Therefore, we performed a butterfly trapping study in Ghana, using both understorey and canopy traps, and sampling seven periods covering dry and wet seasons. Both individual numbers and species richness were on average three to four times higher in the understorey than in the canopy traps with strikingly different species composition (only 11% overlap in species). The number of species and individual abundance recorded in a month varied extensively. This study underlined the importance of taking into account temporal variation and vertical stratification when designing RBAs of fruit-feeding butterflies in West African forests. We recommend the use of both understorey and canopy traps and inclusion of both ‘wet’ and ‘dry’ seasons into RBA sampling regimes. Resume Comme les forets tropicales font face a des menaces graves et qu'elles sont en general situees dans des pays en developpement, une evaluation rapide de la biodiversite (RBA) semble une strategie ideale pour documenter les changements de qualite de l'habitat et etablir des priorites de conservation. On se sert souvent des papillons frugivores comme taxons focaux pour documenter les tendances de la biodiversite dans des forets tropicales. Cependant, le choix de strategies d’echantillonnage inappropriees risque d'annuler leur valeur. Il faut des ensembles de donnees plus complets, qui echantillonnent sur des mois et des saisons et qui integrent une dimension verticale pour etablir la base de reference permettant de concevoir des regimes d’echantillonnage fiables. C'est pourquoi nous avons fait une etude de piegeage des papillons au Ghana, en utilisant des pieges en sous-bois et dans la canopee, et en couvrant sept periodes comptant des saisons seches et des saisons des pluies. Aussi bien les chiffres individuels que la richesse en especes etaient en moyenne 3 a 4 fois superieurs dans les pieges places en sous-bois que dans la canopee, avec une composition des especes remarquablement differente (seulement 11% de recouvrement des especes). Le nombre d'especes et l'abondance individuelle enregistres en un mois variaient enormement. Cette etude souligne qu'il est important de prendre en compte la variation temporelle et la stratification verticale lorsque l'on prepare une RBA de papillons frugivores dans des forets d'Afrique de l'Ouest. Nous recommandons l'emploi de pieges en sous-bois et dans la canopee et l'inclusion de saisons « seches » et « humides » dans les regimes d’echantillonnages d'une RBA.

The broad objective of this paper is to present an overview and synthesis of selected studies on reproduction and aging in two model tephritid fruit fly species including the Mediterranean fruit fly, Ceratitis capitata, and the Mexican fruit fly, Anastrepha ludens. We summarize the research findings from empirical studies and modeling investigations involving reproduction in the two tephritid species. At the end we identify and discuss four general principles regarding reproductive aging in tephritids including reciprocity of reproductive and aging costs, qualitative tradeoffs, plasticity of lifespan and reproduction, and life history constraints and determinacy.

Concentration of the hormone cortisol is often used as an indicator of stress, and chronically high cortisol levels are often associated with poor health. Among group living animals that compete for resources, agonistic social interactions can be expected to contribute to variation in cortisol levels within and among individuals over time. Reproductive tactics of males can change with individual quality, relatedness, and social structure, and affect cortisol levels. In gray-cheeked mangabey (Lophocebus albigena) groups, male rank is an important factor in social interactions, and males also move between groups while actively competing for females or sneaking copulations. During a 20-month study we observed the social behavior and collected 461 fecal samples from 24 adult male gray-cheeked mangabeys from five groups in Kibale National Park, Uganda. Aggressive interactions and the presence of females at the peak of sexual swelling were associated with elevated cortisol concentrations in all males. Independently, dominant (i.e., highest-ranking) males within groups had higher cortisol concentrations than subordinate males, and immigrant males had higher cortisol concentrations than dominant males.

The results of life table experiments to determine the effects of artificial impairment (leg amputation) in 7500 Drosophila melanogaster adults revealed that the extent to which life expectancy was reduced in impaired individuals was conditional on: (1) leg location and number amputated – front leg had greatest impact and the number of legs amputated directly correlated with mortality impact; (2) age of amputation – the greatest relative reduction in remaining life expectancy occurred when young flies were impaired; (3) vial orientation – mortality in impaired flies was the least when vials held upside-down (most friendly environment) and the greatest when they were right-side up (least friendly environment); and (4) sex – male mortality was increased more than female mortality in nearly all impairment treatments. These results were used to formulate a set of general principles of disability that would apply not only to humans but to all organisms.