Supplementary Tree 1.nex Nymphalidae backbone tree inferred with RAxML and time-calibrated using BEAST, with mean posterior node ages and 95% credibility intervals summarized. More information can be found in Chazot et al. (2021). Conserved ancestral tropical niche but different continental histories explain the latitudinal diversity gradient in brush-footed butterflies. Nature Communications.Supplementary Tree 2.nex Maximum clade credibility tree of all Nymphalidae included in "Chazot et al. (2021). Conserved ancestral tropical niche but different continental histories explain the latitudinal diversity gradient in brush-footed butterflies. Nature Communications.", with mean posterior node ages and 95% credibility intervals estimated from the posterior distribution of 1000 grafted trees (1000 subclades posterior trees combined with 1000 backbone posterior trees). More information can be found in Chazot et al. (2021). Conserved ancestral tropical niche but different continental histories explain the latitudinal diversity gradient in brush-footed butterflies. Nature Communications.
Abstract The origins, evolutionary history and diversification of the Australian butterfly fauna are poorly known and uncertain. Two competing hypotheses have been proposed to explain the occurrence of butterflies on this isolated continental landmass. The common view is that all Australian butterflies entered the continent relatively recently from the northern hemisphere via Southeast Asia and/or mainland New Guinea (i.e. northern dispersal origin hypothesis). The alternative view is that part or all of the Australian butterfly fauna ultimately evolved in remnant or Southern Gondwana when Australia was connected to South America through Antarctica (i.e. Southern Gondwanan origin hypothesis). However, robust phylogenies with strong support for monophyly are lacking for the majority of Australian endemic butterfly lineages, thereby precluding determination of their systematic relationships and hence their geographic origins. Here, we use molecular data to reconstruct phylogenetic relationships of the globally distributed butterfly subtribe Coenonymphina (Satyrinae: Satyrini). This group represents a major component of the butterfly fauna of the wider Australasian region, with 19 genera and 71 species endemic to the region. Dating estimates extrapolated from secondary calibration sources indicate that the subtribe arose c . 48 Ma (95% credibility interval, 52–42 Ma), and the crown group first diverged in the Eocene ( c . 44 Ma, 95% credibility interval 51–37 Ma). Rapid speciation events subsequently followed around the Eocence–Oligocene boundary, resulting in a near‐hard polytomy comprising short basal branches with nodes that are difficult to resolve. Based on strongly supported phylogenetic relationships and estimates of divergence times, we conclude that the group probably had its origin in the fragment of Southern Gondwana consisting of Australia, Antarctica and South America. However, we are unable to rule out the northern dispersal scenario, particularly as Coenonymphina are closely related to a set of predominantly Asian lineages. Dispersal and extinction events following the final break‐up of Gondwana have played a pivotal role in shaping the extant distributions of the group.
Abstract 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 butterfly Melanitis leda has been studied in the wild, but other aspects of its mating system remain largely unknown. For a population of M. leda in 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.
Abstract Avoiding detection is vital for the survival of many animals. Factors extrinsic to animals, such as the visual complexity of the background, have been shown to impede the detection of animals. Studies using artificial and natural backgrounds have attributed background complexity to various visual features of the background. One feature that has received less attention is the diversity of color (hue) in the background. We used chickens and artificial backgrounds containing perceptually distinct elements in experiments to test whether color and luminance diversity affect detection time. We found that color diversity in the background impeded detection, while color diversity in prey and luminance diversity in the background did not impede detection. We also did not find an effect of luminance contrast on detection time. Our study suggests a prey animal can benefit in terms of increased detection times by predators when resting on backgrounds with enhanced color diversity.
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. 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. 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. 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 coloration 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. 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.
Butterflies of the subtribe Mycalesina (Nymphalidae: Satyrinae) are important model organisms in ecology and evolution. This group has radiated spectacularly in the Old World tropics and presents an exciting opportunity to better understand processes of invertebrate rapid radiations. However, the generic-level taxonomy of the subtribe has been in a constant state of flux, and relationships among genera are unknown. There are six currently recognized genera in the group. Mycalesis, Lohora and Nirvanopsis are found in the Oriental region, the first of which is the most speciose genus among mycalesines, and extends into the Australasian region. Hallelesis and Bicyclus are found in mainland Africa, while Heteropsis is primarily Madagascan, with a few species in Africa. We infer the phylogeny of the group with data from three genes (total of 3139 bp) and use these data to reconstruct events in the biogeographic history of the group. The results indicate that the group Mycalesina radiated rapidly around the Oligocene-Miocene boundary. Basal relationships are unresolved, but we recover six well-supported clades. Some species of Mycalesis are nested within a primarily Madagascan clade of Heteropsis, while Nirvanopsis is nested within Lohora. The phylogeny suggests that the group had its origin either in Asia or Africa, and diversified through dispersals between the two regions, during the late Oligocene and early Miocene. The current dataset tentatively suggests that the Madagascan fauna comprises two independent radiations. The Australasian radiation shares a common ancestor derived from Asia. We discuss factors that are likely to have played a key role in the diversification of the group. We propose a significantly revised classification scheme for Mycalesina. We conclude that the group originated and radiated from an ancestor that was found either in Asia or Africa, with dispersals between the two regions and to Australasia. Our phylogeny paves the way for further comparative studies on this group that will help us understand the processes underlying diversification in rapid radiations of invertebrates.
Abstract Objectives In primates, allogrooming and other affiliative behaviors confer many benefits and may be influenced by many socioecological factors. Of these, the impact of anthropogenic factors remain relatively understudied. Here we ask whether interactions with humans decreased macaques' affiliative behaviors by imposing time‐constraints, or increased these behaviors on account of more free‐/available‐time due to macaques' consumption of high‐energy human foods. Materials and methods In Southern India, we collected data on human–macaque and macaque–macaque interactions using focal‐animal sampling on two groups of semi‐urban bonnet macaques for 11 months. For each macaque within each climatic season, we calculated frequencies of human–macaque interactions, rates of monitoring human activity and foraging on anthropogenic food, dominance ranks, grooming duration, number of unique grooming partners, and frequencies of other affiliative interactions. Results We found strong evidence for time‐constraints on grooming. Macaques that monitored humans more groomed for shorter durations and groomed fewer partners, independent of their group membership, sex, dominance rank, and season. However, monitoring humans had no impact on other affiliative interactions. We found no evidence for the free‐time hypothesis: foraging on anthropogenic food was unrelated to grooming and other affiliation. Discussion Our results are consistent with recent findings on other urban‐dwelling species/populations. Macaques in such environments may be especially reliant on other forms of affiliation that are of short duration (e.g., coalitionary support, lip‐smacking) and unaffected by time‐constraints. We stress on the importance of evaluating human impact on inter‐individual differences in primate/wildlife behavior for conservation efforts.
Many butterflies possess striking structures called eyespots on their wings, and several studies have sought to understand the selective forces that have shaped their evolution. Work over the last decade has shown that a major function of eyespots is their ability to reduce predation by being intimidating to attacking predators. Two competing hypotheses seek to explain the cause of intimidation, one suggesting 'eye-mimicry' and the other their 'conspicuousness' as the reason. There is an on-going debate about which of these better explains the effectiveness of eyespots against predation. We undertook a series of indoor experiments to understand the relative importance of conspicuousness and eye-mimicry, and therefore how predator perception may have influenced the evolution of eyespots. We conducted choice tests where artificial paper models mimicking Junonia almana butterflies were presented to chickens and their preference of attack recorded.We first established that birds avoided models with a pair of eyespots. However, contrary to previous, outdoor experiments, we found that the total area of eyespots did not affect their effectiveness. Non-eye-like, fan shaped patterns derived from eyespots were found to be just as effective as eye-like circular patterns. Furthermore, we did not find a significant effect of symmetry of patterns, again in discordance with previous work. However, across all experiments, models with a pair of patterns, symmetric or asymmetric, eyelike or non-eye-like, suffered from fewer attacks compared with other models.The study highlights the importance of pairedness of eyespots, and supports the hypothesis that two is a biologically significant number that is important in prey-predator signalling. We discuss the implications of our results for the understanding of eyespot evolution.
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