Biology subjects

Chazot, N.

Publications and source records attributed to Chazot, N..

14 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↗

bioRxiv2026

A meta-analysis of Neotropical butterflies reveals long-term signature of the Pebas wetland on Neotropical biogeography

The palaeogeographic history of the Neotropics has inspired multiple diversification models, emphasizing either the Andes or Amazonia as primary drivers of biodiversity. However, the extensive Pebas wetland system that covered western Amazonia during the Miocene has gained attention as a potential barrier to dispersal and diversification. This system may have contributed to the modern diversity imbalance between the Andes and Amazonia, known as the Gentry pattern. Most evidence for this hypothesis comes from isolated case studies and fossils. Here, we conduct a meta-analysis including 1,361 species of Neotropical butterflies to evaluate the role of the Pebas wetland on the dynamics of dispersal and diversification. We found that Andean- and Amazonian-centred clades show complementary but temporarily distinct biogeographic patterns. High Andean diversity reflects an Andean origin, with early occupancy and diversification occurring within the Andes, followed by post-Pebas dispersal into Amazonia. By contrast, Amazonian-centred clades, reflect an Amazonian origin, with an early diversification in Amazonia and subsequent post-Pebas colonization of the Andes. While several clades show little evidence of Pebas influence, our study suggests that the Pebas wetland did not uniformly restrict diversification, but rather selectively constrained dispersal between the Andes and Amazonia, preventing simultaneous diversification in both regions and thereby creating a diversity imbalance that characterizes the modern Neotropic.

evolutionary biology↗

bioRxiv2025

Discovering the unseen: a performance comparison of taxonomic classification methods under unknown DNA barcodes

O_LIDNA barcoding and metabarcoding have emerged as cost-efficient, standardized methods for characterizing local biodiversity. Based on the sequencing of a small targeted gene fragment, it is theoretically possible to identify a wide diversity of taxa by comparing them with reference sequence databases. However, a key challenge for accurate taxonomic classification is the incompleteness of such databases, leading to most query sequences lacking species-level matches. C_LIO_LIWhere species-level matches are missing, it may be possible to classify query sequences to a higher taxonomic level, such as genus or family, based on the similarity of related reference taxa. The challenge then lies in confidently recognizing whether the sequence belongs to an unobserved (here, "novel") taxon on a given taxonomic level. C_LIO_LIIn this study, we evaluated the performance and utility of several methods for taxonomic classification. Methods were assessed based on the classification accuracy of both observed and novel taxa, training time, space requirements, and run time. We did this for two cases: the COI barcode for arthropods, and the ITS barcode for fungi, with the latter representing an instance with substantially greater sequence similarity variation within classes. To test classification of novel taxa, we used well-curated datasets with partially distinct taxonomic distribution between the training and test set. Novel taxa occurred at all evaluated taxonomic levels, such as novel species in observed genera and novel genera in observed families. We further assessed the effect on performance when shifting from full-length barcodes to shorter sequences as generated through metabarcoding in the testing dataset. C_LIO_LIThis study sheds light on the strengths and limitations of different classification algorithms across varied ecological contexts and provides guidance for researchers in selecting suitable algorithms for DNA barcoding and metabarcoding applications. In particular, it demonstrates the supreme performance of phylogenetic placement methods such as EPA-ng for classification of arthropod COI barcodes, and composition-based classifiers such as SINTAX, RDP-NBC, and IDTAXA for fungal ITS. C_LI

bioinformatics↗

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

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

Large-scale assessment of habitat quality and quantity change on declining European butterflies

The rapid decline of biodiversity as a consequence of increased environmental impact by human activities requires urgent action against the ongoing crisis. At the heart of conservation policy is the debate on quality versus quantity: should the post-2020 Global Biodiversity Framework focus on maximising total protected area, or pursue instead qualitative targets? To improve conservation practices, we need to both understand the current trend of biodiversity and the factors driving the decline. We address this by: (1) projecting current European butterfly population trends for the next 50 and 100 years, (2) comparing the decline in species richness, phylogenetic diversity and habitat specialization among different habitats, and (3) estimating the relationship between recent trends in habitat quality or quantity and the decline of multiple metrics of European butterfly diversity. We do not find any significant relationship between diversity loss and habitat area loss and conclude that neither long-term nor short-term changes in habitat area are primary predictors of large-scale butterfly decline in Europe. Habitat quality emerges as the most crucial factor in our analyses - both the area affected and the severity of habitat quality reduction. Habitat degradation causes vegetation changes in structure and diversity, which affect butterfly survival. We also estimate a higher decline of habitat specialists than generalist species. We conclude that habitat protection cannot focus solely on the maximization of habitat area but urgently needs to restore high quality ecosystems to provide the full range of ecological requirements for biodiversity.

ecology↗

bioRxiv2021

Fruit colour and range size interact to influence diversification

AimDifferent fruit colours are associated with dispersal by different frugivores, largely based on colour vision type. Frugivore mobility affects overall range size for the plant being dispersed. Here we determine the interaction between different fruit colours, range sizes, and diversification rates by testing two hypotheses: That (1) fruit colours attractive to birds have larger range sizes due to their higher dispersal ability, and that (2) different frugivore disperser groups, bird or mammal, leads to different diversification rate at different range size, where intermediate range size leads to the highest diversification rate. LocationGlobal. Time periodContemporary (or present) Major taxa studiedPalms (Arecaceae) MethodsUsing model selection, we identified three groups of colours with similar diversification rate and likely disperser. Range sizes were estimated and categorized species as small, intermediate, or large-ranged. For model selection and to determine the relationship beween fruit color, range size and diversification rate we used Multi-State Speciation and Extinction (MuSSE) models. ResultsSpecies with intermediate range size had the highest net diversification for all three fruit colour groups. Bird-dispersed palms more likely diversified at small than at large range size while mammal-dispersed palms more likely diversified at larger range size than small. Fruit colours associated with mammal dispersal had more large-ranged species than colours associated with bird dispersal. Main conclusionsThe associated between intermediate range size and higher diversification rate indicates that spatial factors that affect diversification at small and large range sizes result in higher diversification at intermediate ranges. We find striking differences in diversification rate within each range size category between fruit color groups. This suggests that the relationship between diversification rate and range size depends on the specific frugivorous dispersers and their dispersal patterns. This study reveals how fruit traits alter dispersal patterns and how that, in turn, influences diversification.

ecology↗

bioRxiv2021

Temperature predicts leaf shape in palms (Arecaceae)

AimLeaves display a remarkable variety of shapes, each with potential ecological advantages in specific climates. While relations between leaf shape and either climate or height has been relatively well studied in eudicots, the macroecological drivers of shape remain poorly known in monocots. Here, we investigated associations between climate and plant height with the evolution of leaf shape in a clade with high species and morphological diversity. LocationGlobal. Time periodCretaceous to contemporary. Major taxa studiedPalms (Arecaceae). MethodsWe apply a Bayesian phylogenetic mixed model to test for associations between climate and leaf shape (all entire-leaved, pinnate-dissected, palmate-dissected, and costapalmate). We further reconstruct the ancestral leaf shape using multistate speciation and extinction models and compare the frequency of shapes with global temperatures through time. ResultsWe find that plant height associates with dissected leaves and that annual precipitation associates with pinnate shapes. The ancestral leaf shape is unclear but early diversification was dominated by pinnate-dissected palms, which has remained the most species-rich form of leaves throughout palm history. Main conclusionsPalms that are tall and live in humid regions are more likely to have pinnate leaves. Through geological time scales, temperature did not play an obvious role in determining leaf shapes. This study contributes to our understanding of how the diversity of leaf shapes is linked to biological and climatic factors.

evolutionary biology↗

bioRxiv2021

Punctual ecological changes rather than global factors drive species diversification and the evolution of wing phenotypes in Morpho butterflies.

Assessing the relative importance of geographical and ecological drivers of evolution is paramount to understand the diversification of species and traits at the macroevolutionary scale. Here, we use an integrative approach, combining phylogenetics, biogeography, ecology, and quantified phenotypes to investigate the drivers of both species and phenotypic diversification of the iconic Neotropical butterfly genus Morpho. We generated a time-calibrated phylogeny for all known species and inferred historical biogeography. We fitted models of time-dependent (accounting for rate heterogeneity across the phylogeny) and paleoenvironment-dependent diversification (accounting for global effect on the phylogeny). We used geometric morphometrics to assess variation of wing size and shape across the tree, and investigated their dynamics of evolution. We found that the diversification of Morpho is best explained when considering multiple independent diversification dynamics across the tree, possibly associated with lineages occupying different microhabitat conditions. First, a shift from understory to canopy was characterized by an increased speciation rate partially coupled with an increasing rate of wing shape evolution. Second, the occupation of dense bamboo thickets accompanying a major host-plant shift from dicotyledons towards monocotyledons was associated with a simultaneous diversification rate shift and an evolutionary "jump" of wing size. Our study points to a diversification pattern driven by punctual ecological changes instead of a global driver or biogeographic history.

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↗

bioRxiv2019

A complete time-calibrated multi-gene phylogeny of the European butterflies

With the aim of supporting ecological analyses in butterflies, the third most species-rich superfamily of Lepidoptera, this paper presents the first time-calibrated phylogeny of all 496 extant butterfly species in Europe, including 18 very localized endemics for which no public DNA sequences had been available previously. It is based on a concatenated alignment of the mitochondrial gene COI and up to 11 nuclear gene fragments, using Bayesian inference of phylogeny. To avoid analytical biases that could result from our region-focus sampling, our European tree was grafted upon a global genuslevel backbone butterfly phylogeny for analyses. In addition to a consensus tree, we provide the posterior distribution of trees and the fully-concatenated alignment for future analyses.

evolutionary biology↗

bioRxiv2018

The Trials and Tribulations of Priors and Posteriors in Bayesian Timing of Divergence Analyses: the Age of Butterflies Revisited

The need for robust estimates of times of divergence is essential for downstream analyses, yet assessing this robustness is still rare. We generated a time-calibrated genus-level phylogeny of butterflies (Papilionoidea), including 994 taxa, up to 10 gene fragments and an unprecedented set of 12 fossils and 10 host-plant node calibration points. We compared marginal priors and posterior distributions to assess the relative importance of the former on the latter. This approach revealed a strong influence of the set of priors on the root age but for most calibrated nodes posterior distributions shifted from the marginal prior, indicating significant information in the molecular dataset. We also tested the effects of changing assumptions for fossil calibration priors and the tree prior. Using a very conservative approach we estimated an origin of butterflies at 107.6 Ma, approximately equivalent to the Early Cretaceous-Late Cretaceous boundary, with a credibility interval ranging from 89.5 Ma (mid Late Cretaceous) to 129.5 Ma (mid Early Cretaceous). This estimate was robust to alternative analyses changing core assumptions. With 994 genera, this tree provides a comprehensive source of secondary calibrations for studies on butterflies.

evolutionary biology↗

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↗