Biology subjects

Willmott, K. R.

Publications and source records attributed to Willmott, K. R..

10 recordsLinked to original sources

bioRxiv2026

Decoupled wing evolution reveals trait-specific links between morphology, geography, and diversification in Ithomiini butterflies.

Understanding when species diversification and phenotypic evolution are coupled remains a central challenge in macroevolution. Such coupling may depend on the traits considered, phenotypic integration, or the geographic context in which diversification occurs. Here, we investigate these relationships in Ithomiini, a diverse Neotropical butterfly tribe whose diversification is strongly associated with the Andes-Amazonia system. We quantified forewing and hindwing size and shape for 1,618 individuals representing 291 species using geometric morphometrics, and combined these data with phylogenetic comparative methods, diversification-rate estimates, and spatial analyses of species distributions. Evolutionary rate shifts were largely trait- and wing-specific, indicating limited coupling between forewing and hindwing evolution. Across the phylogeny, wing size evolution and extant wing size were not associated with species diversification, nor was hindwing shape evolution. By contrast, forewing shape evolutionary rates showed a positive association with diversification. Spatial analyses revealed that this coupling was geographically concentrated in the Andes, where high diversification rates coincided with elevated forewing size and shape diversity and increased forewing evolutionary rates. Hindwing diversity, by contrast, more closely tracked phylogenetic diversity and showed weaker or opposing relationships with diversification. Our results show that diversification and phenotypic evolution in Ithomiini are neither generally coupled nor fully independent, but linked in trait-specific and geographically structured ways. More broadly, they highlight the need to decompose complex phenotypes and integrate spatial context when testing macroevolutionary coupling.

ecology↗

bioRxiv2025

Aposematic color patterns are the dominant axis of phenotypic diversification in Nymphalid butterflies

Butterfly wing patterns serve diverse roles in visual communication, from aposematic signaling and mimicry to mate attraction and camouflage. In brush-footed butterflies (Nymphalidae), this diversity can be traced to the wing pattern "ground plan" that generates phenotypes deterministic in origin yet highly multidimensional in form. Quantifying such complexity at scale has long been a challenge, limiting our understanding of how visual signals interact, constrain one another, and evolve. Here, we used computer vision to extract high-dimensional traits from standardized museum specimens and assembled the largest comparative dataset of wing color patterns to date, spanning over one third of all known Nymphalid species. We first tested whether chemically defended species occupy a distinct region in morphospace and then derived a quantitative score for aposematism from the principal color patterns associated with defense. Using this score, we examined whether aposematic signals are expressed consistently across wing surfaces and sexes, and whether their origins are linked to shifts in evolutionary rate. We found that the dominant axis of morphospace is defined by chromatic and achromatic contrast, along which defended and undefended species cluster. Validation with an expert-labeled moth dataset confirmed that this axis separates aposematic from non-aposematic phenotypes across Lepidoptera. Consistent with theory, strongly aposematic species showed greater visual similarity between dorsal and ventral surfaces, between sexes, and among individuals. Rate analyses further indicated that aposematic patterns evolved repeatedly and were associated with non-linear shifts in evolutionary tempo. Together, these results identify aposematism as the dominant organizing axis of wing color pattern evolution in Nymphalidae. Significance statementButterflies are renowned for their striking diversity of wing patterns, including the warning colors that signal chemical defense to predators. Yet whether such warning patterns share common features across lineages has remained unclear. Here, we applied a metric computer vision model to more than 16,000 museum specimens spanning one third of all Nymphalid species, encoding their wing patterns into a common morphospace. Within this space, high-contrast aposematic patterns emerged as the dominant axis of diversification, explaining up to 20% of phenotypic variance. These signals were expressed consistently across wing surfaces, sexes, and individuals, and evolved repeatedly across the family. Our approach demonstrates how computer vision enables meaningful comparative analyses of complex patterns, revealing general principles of butterfly diversification.

evolutionary biology↗

bioRxiv2025

Phylogeny, systematics and evolution of mimicry patterns in Neotropical limenitidine butterflies

The Neotropical butterfly genus Adelpha Hubner exhibits remarkable species diversity and striking convergence in wing colour patterns potentially explained by mimicry, making it an exceptional model for exploring trait evolution and its relationship with speciation. To date, unresolved phylogenetic relationships hinder a comprehensive understanding of the evolutionary biology of the genus. Using a novel multi-marker dataset combining one mitochondrial and 15 nuclear gene fragments, we generate the most comprehensive phylogeny of the genus Adelpha to revisit its systematics and investigate the evolution of mimicry colour patterns. Our data set encompasses 83 of the 87 known extant species and six Limenitis species that were recently excluded from Adelpha (134 of c. 160 subspecies in total), collectively displaying 14 distinct mimicry patterns. We provide conclusive evidence that corroborates previous work on the polyphyly of Adelpha as historically conceived, and describe the genus Adelphina Paez & Willmott n. gen. to stabilize the nomenclature, both genera representing Neotropical limenitidines. The comprehensive phylogeny provided in this study lays a solid foundation for future research into the processes driving diversification within these species interacting through mimicry. Ancestral character state reconstruction reveals gradual evolution of mimicry pattern. The more common mimicry pattern IPHICLUS (forewing with orange subapical spot and white band) is inferred as ancestral, but repeated convergent evolution is also recovered. Evolutionary convergence is also observed for the second most abundant mimicry pattern, COCALA (orange-white banded). Increased rates of mimicry pattern evolution are also found toward the equator. These results underscore the complexity of mimicry evolution in the Neotropical limenitidines i.e., Adelpha and Adelphina, emphasizing the need to explore its interplay with other biotic and abiotic factors.

evolutionary biology↗

bioRxiv2024

Genomics of Neotropical biodiversity indicators: two butterfly radiations with rampant chromosomal rearrangements and hybridisation

A major question in evolutionary biology is what drives the diversification of lineages. Rapid, recent radiations are ideal systems for addressing how new species arise because they still show key morphological and ecological adaptations associated with speciation. While most studied recent radiations have evolved in an insular environment, less research has been carried out on continental radiations with complex species interactions. Melinaea and Mechanitis butterflies (Nymphalidae: Ithomiini) have rapidly radiated in the Neotropics. They are classical models for Amazonian biogeography and colour pattern mimicry and have been proposed as biodiversity indicators. We generated reference genomes for five species of each genus, and whole-genome resequencing data of most species and subspecies covering a wide geographic range to assess phylogeographic relationships, patterns of hybridisation and chromosomal rearrangements. Our data help resolve the classification of these taxonomically challenging butterflies and reveal very high diversification rates. We find rampant evidence of historical hybridisation and putative hybrid species in both radiations, which may have facilitated their rapid diversification. Moreover, dozens of chromosomal fusions and fissions were identified between congeneric species, and even some within species. We conclude that interactions between geography, hybridisation and chromosomal rearrangements have contributed to these two rapid radiations in the highly diverse Neotropical region. We suggest that rapid radiations may be spurred by repeated periods of geographic isolation during Pleistocene climate oscillations, combined with lineage-specific rapid accumulation of incompatibilities during allopatric phases, followed by secondary contact with some gene exchange. Significance StatementUnderstanding factors contributing to rapid speciation is a key aim of evolutionary biology. Here we focus on two rapid radiations of Neotropical butterflies. Our genomic data with broad taxonomic and geographic coverage reveal rampant hybridisation and chromosomal rearrangements, each likely contributing to the high diversification rates. Our study highlights the use of genomic data to resolve taxonomically challenging species groups and elucidate drivers of diversification in rapid radiations. We show that for biodiversity hotspots with recent radiations, barcoding is insufficient to characterise species richness due to gene flow and recent speciation. The taxonomic implications of both introgression and karyotype diversity for species delimitation are important to consider during monitoring and management of biodiversity in these vulnerable habitats.

evolutionary biology↗

bioRxiv2024

Time-dependent effects of species interactions at macroevolutionary scale shape the diversity of mimetic butterflies.

Evidence that species interactions can affect macroevolutionary dynamics of trait and species diversification is scarce. Mutualistic Mullerian mimicry is a compelling example of example of ecological interactions that has been shown to drive evolutionary convergence, Here, we test how mutualistic Mullerian mimicry shapes macroevolutionary patterns of diversification in the Ithomiini butterflies. We show that the age of color patterns is the primary determinant of species richness within mimicry rings but not phylogenetic diversity. We find pervasive phylogenetic signal in mimicry rings and in color patterns associated within polymorphic species. Only a small set of mimicry rings show high phylogenetic diversity. We identify patterns of saturation in the accumulation of new mimicry rings and in the number of evolutionary convergences towards the most species-rich mimicry rings. Those saturation patterns are likely caused by niche filling along various ecological dimensions, within and among the mimetic communities living in sympatry. The time-dependent effects detected in our study illustrate how neutral processes and ecological interactions interact and shape species and phenotypic diversification.

evolutionary biology↗

bioRxiv2022

Diversification is correlated with temperature in white and sulfur butterflies

Temperature is thought to be a key variable explaining global patterns of species richness. However, to investigate this relationship carefully, it is necessary to study clades with broad geographic ranges that are comprised of species inhabiting diverse biomes with well- characterized species ranges. In the present study, we investigate the link between temperature and diversification in the butterfly family Pieridae (sulfurs and whites) by combining Next Generation sequences and published molecular data with fine-grained distribution information. After building the most comprehensive phylogeny of the group, with almost 600 species and all higher taxa (subfamilies, tribes and subtribes), we found strong support for the following relationships within the family: Dismorphiinae + (Coliadinae + (Pseudopontiinae + Pierinae)). With a curated distribution dataset of over 800,000 occurrences, we conducted multiple comparative phylogenetic analyses that provided strong evidence that species in environments with more stable daily temperatures or with cooler maximum temperatures in the warm seasons have higher diversification rates. We also found a positive correlation between speciation and extinction with paleotemperature: as global temperature decreased through geological time, so did diversification rates. Although many studies demonstrate higher diversity in the tropics, we have been able to identify specific climate variables associated with changes in diversification, while also inferring the most robust and well sampled phylogenomic framework for Pieridae to date.

evolutionary biology↗

bioRxiv2020

Hard to catch: Experimental evidence supports evasive mimicry

Most research on aposematism has focused on chemically defended prey, but signalling difficulty of capture remains poorly explored. Similarly to classical Batesian and Mullerian mimicry related to distastefulness, such "evasive aposematism" may also lead to convergence in warning colours, known as evasive mimicry. A prime candidate group for evasive mimicry are Adelpha butterflies, which are agile insects and show remarkable colour pattern convergence. We tested the ability of naive blue tits to learn to avoid and generalise Adelpha wing patterns associated with difficulty of capture, and compared their response to that of birds that learned to associate the same wing patterns with distastefulness. Birds learned to avoid all wing patterns tested, but learning was faster with evasive prey compared with distasteful prey. Birds generalised their learned avoidance from evasive models to imperfect mimics if the mimic shared colours with the model. Despite imperfect mimics gaining protection from birds generalisation, perfect mimics always had the best fitness, supporting selection for accurate mimicry. Faster avoidance learning and broader generalisation of evasive prey suggest that being hard to catch may deter predators at least as effectively as distastefulness. Our results provide empirical evidence for a potentially widespread alternative scenario, evasive mimicry, for the evolution of similar aposematic colour patterns.

evolutionary biology↗

bioRxiv2020

The latitudinal diversity gradient in brush-footed butterflies (Nymphalidae): conserved ancestral tropical niche but different continental histories.

The latitudinal diversity gradient (LDG) is arguably one of the most striking patterns in nature. The global increase in species richness toward the tropics across continents and taxonomic groups stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluated several of these hypotheses to explain spatial diversity patterns in the butterfly family, Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal to the LDG, and also the extent to which these processes differ among regions at the same latitude. We generated a new, time-calibrated phylogeny of Nymphalidae based on 10 gene fragments and containing ca. 2,800 species ([~]45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the LDG among regions because temporal diversification dynamics differ greatly across longitude. For example, we found that Neotropical nymphalid diversity results from low extinction rates, not high speciation rates, and that biotic interchanges with other regions were rare. Southeast Asia was also characterized by a low speciation rate but, unlike the Neotropics, was the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, particularly during the Eocene-Oligocene transition, combined with the conserved ancestral tropical niches, played a major role in generating the modern LDG of butterflies.

ecology↗

bioRxiv2017

Renewed diversification following Miocene landscape turnover in a Neotropical butterfly radiation

\"This preprint has been reviewed and recommended by Peer Community In Evolutionary Biology (http://dx.doi.org/10.24072/pci.evolbiol.100032)\"\n\nThe Neotropical region has experienced a dynamic landscape evolution throughout the Miocene, with the large wetland Pebas occupying western Amazonia until 11-8 my ago and continuous uplift of the Andes mountains along the western edge of South America. Although the complex dynamics between the Andes and Amazonia may have strongly affected the trajectory of Neotropical biodiversity, there is little evidence for such an influence from time-calibrated phylogenies of groups that diversified during this period. Here, we generate one of the most comprehensive time-calibrated molecular phylogenies of a group of Neotropical insects: the butterfly tribe Ithomiini. Our tree includes 340 species (87% of extant species), spanning 26 million years of diversification in the Neotropics. We investigate temporal and spatial patterns of diversification, focusing on the influence of Miocene landscape tranformations on the dynamics of speciation, extinction and biotic interchanges at the Amazonia/Andes interface. We find that Ithomiini likely began diversifying at the interface between the Andes and the Amazonia around 26.4 my ago. Five subtribes with a very low extant diversity started diversifying early in western Amazonia, but a rapid decrease in diversification rate due to increased extinction rate between 20 and 10 my ago suggests a negative impact of the Pebas wetland system on these early lineages. By contrast, the clade containing the five most species-rich subtribes (85% of extant species) was characterized by a high, positive net diversification rate. This clade diversified exclusively in the Central Andes from 20 to 10 my ago. After the demise of the Pebas system (11-8 my ago), we found a sudden increase of interchanges with the Northern Andes and Amazonia, followed by local diversification, which led to a substantial renewal of diversification. In general, ecological turnovers throughout the Miocene strongly determined the dynamics of speciation, and extinction and interchanges, and appear as a key driving force shaping the regions current extraordinary biodiversity.

evolutionary biology↗

bioRxiv2017

North Andean origin and diversification of the largest ithomiine butterfly genus

The Neotropics harbour the most diverse flora and fauna on Earth. The Andes are a major centre of diversification and source of diversity for adjacent areas in plants and vertebrates, but studies on insects remain scarce, even though they constitute the largest fraction of terrestrial biodiversity. Here, we combine molecular and morphological characters to generate a dated phylogeny of the butterfly genus Pteronymia (Nymphalidae: Danainae), which we use to infer spatial, elevational and temporal diversification patterns. We first propose six taxonomic changes that raise the generic species total to 53, making Pteronymia the most diverse genus of the tribe Ithomiini. Our biogeographic reconstruction shows that Pteronymia originated in the Northern Andes, where it diversified extensively. Some lineages colonized lowlands and adjacent montane areas, but diversification here remained scarce. The recent colonization of lowland areas was reflected by an increase in the rate of evolution of species elevational ranges towards present. By contrast, speciation rate decelerated with time, with no extinction. The geological history of the Andes and adjacent regions have likely contributed to Pteronymia diversification by providing compartmentalized habitats and an array of biotic and abiotic conditions, and by limiting dispersal between some areas while promoting interchange across others.

evolutionary biology↗