Biology subjects

Matos-Maravi, P.

Publications and source records attributed to Matos-Maravi, P..

12 recordsLinked to original sources

bioRxiv2026

Contrasting signatures of interspecific gene flow in the Atlantic Forest endemic butterfly Heliconius besckei

Gene flow between diverging species can reshuffle genomic variation and greatly influence the speciation process, sometimes reinforcing, sometimes eroding species boundaries. However, the effect of ecological and historical processes on the patterns of introgression remains poorly understood. Here, we focus on the evolutionary history of Heliconius besckei, a geographically restricted butterfly endemic to the Brazilian Atlantic Forest, together with the mimetic H. melpomene nanna, the sympatric non-mimetic H. ethilla narcaea, and its closely related H. numata robigus. Using whole-genome sequencing, genome-wide tests of introgression, demographic modelling and palaeoclimatic species distribution models, we tested the extent and timing of interspecific gene flow in relation to mimicry, ecological context, and changes in geographic overlap. Despite strong genomic differentiation among the four focal Heliconius species, we detected widespread signatures of interspecific gene flow, greatly influenced by geographical overlap. Demographic models including post-divergence gene flow were better supported than strict-isolation models. In addition, genome-wide introgression analyses showed more numerous and larger candidate introgressed regions between H. besckei and H. ethilla narcaea, consistent with recurrent opportunities for exchange between these currently sympatric species. Species distribution models further recovered repeated expansions of suitable habitat for H. besckei during cold periods, providing a potential geographic context for secondary contact within the Atlantic Forest. Together, our results reveal recurrent gene flow occurring under different temporal, geographic, and potentially selective contexts, and show that the evolutionary history of a geographically restricted lineage can be shaped by repeated genetic exchange while substantial species differentiation is maintained.

evolutionary biology↗

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↗

bioRxiv2026

Heated debate: Is the seasonal polyphenism of Araschnia levana (Lepidoptera: Nymphalidae) driven by thermoregulation?

The seasonal forms of the temperate butterfly Araschnia levana (Nymphalidae: Nymphalinae) differ in morphology (weight, wing area, and wing loading) and colouration. Spring individuals are predominantly orange with higher weight per wing area, (i.e. wing loading) while summer individuals are black with a white stripe and have lower wing loading. However, it remains unclear if and how these seasonal differences affect heating and cooling dynamics. We compared thermal responses of seasonal forms, focusing on the roles of morphology and colouration. Further, we assessed whether live butterflies heat and cool differently from dead individuals to detect the presence of active thermoregulation. Morphology and colouration influenced the thermal dynamics of the thorax and wings as expected from heat-transfer principles, but we found no evidence of active thermoregulation on the thorax. Based on aligned temperature curves, seasonal forms showed similar thermal dynamics. This similarity was driven by morphology and colouration, with larger wing area accelerating thermal change and higher body weight (or wing loading) reducing it, thereby masking underlying form-specific patterns. After accounting for significant morphological differences between forms, the thorax of spring individuals heated and cooled faster than that of summer ones. This trend suggests form-specific optimisation of thermal performance, likely as a response to temperate climates. Thermal responses differ between forms in ways not directly explained by the polyphenism itself, potentially reflecting a broader trait of multivoltine ectotherms to cope with seasonal temperature changes.

ecology↗

bioRxiv2025

Environmental Variation Promotes Convergent Evolution and Rapid Diversification of Wing Shape and Color in Skipper Butterflies

Predation is a key driver of speciation and phenotypic diversification, yet how antipredator traits evolve and persist over evolutionary time remains poorly understood. We generated target sequence capture and whole-genome sequencing data for the skipper butterfly subfamily Eudaminae (Hesperiidae) to test whether antipredator defenses, in interaction with environmental variation, promote diversification. We focus on two antipredation wing traits: hindwing tails, which deflect predator attacks, and blue-green coloration, which may enhance motion dazzle and be used as warning coloration. Applying phylogenomics, morphometrics, and comparative methods, we model trait evolution in relation to diel activity and geographic distribution and find that hindwing tails repeatedly evolved at least seven times and blue-green coloration at least fifteen times. Both traits are associated with elevated speciation rates, but evolutionary transitions toward tailless wings and non-iridescent coloration occurred more frequently than trait gains, indicating high evolutionary lability of these antipredator defenses. Trait loss, particularly pronounced in tropical diurnal species, may reflect trade-offs in flight performance, shifts in predation guilds, or the evolution of alternative defensive traits. Our findings highlight rampant convergent evolution of wing traits that are under strong predator-mediated selection. By identifying how antipredator defenses and environmental contexts influence phenotypic and species diversification, this study provides new insights into the ecological and evolutionary processes underlying insect diversity.

evolutionary biology↗

bioRxiv2025

Association of iridescence and tails with flight proxies suggests that predation drives convergence in skipper butterflies

1. Phenotypic evolution is shaped by phylogenetic, ecological and biomechanical constraints. Butterfly wings offer an ideal system to explore these constraints as their evolution is strongly influenced by natural and sexual selection affecting colour patterns and wing shape in relation to visual signalling and flight performance. 2. We focus on Eudamina skippers, butterflies with large-scale phenotypic convergences involving wing shape and colour pattern. Using a comprehensive morphometric dataset encompassing 176 species, we assess their morphological evolution in a phylogenetic framework. 3. We show that hindwing tails and dorsal iridescence, proposed to signal high escape ability, have repeatedly evolved in tropical species. Hindwing tails are associated with intermediate body size, low wing loading and a forewing shape that facilitates manoeuvrability; iridescence is more common in large species with significantly high wing loading and aspect ratio, indicating high flight speed and manoeuvrability. 4. The evolution of all tested morphometric traits is best described by Ornstein-Uhlenbeck models, indicating stabilising selection around fitness optima in Eudamina. Models testing for random drift or early burst dynamics were rejected in all cases. 5. We reveal a clear co-evolution between the fore- and hindwing shapes, despite both wings having different functional constraints. Hindwing tails have a significant recurrent effect on forewing shape in lineages evolving them independently, suggesting the existence of evolutionary trade-offs between flight performance and antipredator defences. 6. Our study shows that the convergent morphologies across Eudamina have likely arisen not by neutral evolution but by ecological selection influenced by biomechanical constraints.

evolutionary biology↗

bioRxiv2024

Quantifying the success of prey crypsis, aposematism and evasiveness in avoiding avian predators attack

Antipredator defences typically act at distinct stages of the predation sequence--encounter, identification, approach, and subjugation. However, their effectiveness has rarely been quantified and compared simultaneously in wild predator-prey systems. We conducted a study in Peru, where we installed aviaries at two localities and recorded the responses of wild avian predators to three types of antipredator defences--crypsis, aposematism, and evasiveness--expressed by three butterfly species. The study included both immature and adult birds from insectivorous species in forest and urban environments. We tested the theoretical expectations that cryptic butterflies (Nymphalidae: Euptychiina) were rarely detected, aposematic Heliconius (Nymphalidae: Heliconiinae) were often sighted but seldom attacked, and evasive Spicauda (Hesperiidae: Eudaminae) were frequently detected and attacked but evaded capture at higher rates. Despite these distinct defensive strategies, mortality rates among prey types were largely similar, but predator life stage strongly influenced defence effectiveness. Immature birds were more likely to attack Heliconius, possibly reflecting a lack of learned avoidance for aposematic signals. Additionally, predator family influenced predation patterns, with more skilled insectivores (e.g., Vireonidae) showing higher capture success against defended prey. These findings illuminate the evolutionary pressures that shape predator behaviour and prey defences in tropical ecosystems, where high predator diversity possibly maintains the coexistence of multiple defensive strategies. The observed similar mortality rates underscore the adaptive value of these defences, as they collectively distribute the total predation pressure across prey species.

ecology↗

bioRxiv2023

Reduced palatability, fast flight, and tails: Decoding the defence arsenal of Eudaminae skipper butterflies in a Neotropical locality

O_LIPrey often relies on multiple defences to avoid predators, such as difficulty of capture, attack deflection, or unpleasant taste. Eudaminae skipper butterflies rely on behavioural (fast flight) and mechanical defences (hindwing tails), but other defences like unpalatability (consumption deterrence), and the interaction among defences, have not been assessed. C_LIO_LIWe test the palatability of 12 abundant Eudaminae species from a Neotropical locality, using training and feeding experiments with domestic chicks. Further, we approximate the difficulty of capture explained by flight speed and quantified by wing loading. To test how multiple defences are associated and explained by body size and habitat preference, we perform phylogenetic Bayesian regression analyses. C_LIO_LIWe found a broad range of palatability in Eudaminae, within and among species. Palatability was negatively correlated with wing loading, indicating that butterflies with higher flight speed tend to have lower palatability. C_LIO_LIThe presence or absence of tails did not explain the level of butterfly palatability, showing that attack deflection and consumption deterrence are not mutually exclusive defences. Habitat preference (open or forested environments) did not explain the level of palatability in Eudaminae, but it seems that butterflies with high wing loading (fast flight) tended to occupy semi-closed (e.g., hilltops) or closed habitats (e.g., secondary forest). C_LIO_LIFinally, the level of unpalatability in Eudaminae is size dependent, larger butterflies are less palatable, perhaps due to their more likely detectability/preference by predators. Altogether, our findings shed light on the contexts favouring the prevalence of single vs. multiple defensive strategies in prey. C_LI

ecology↗

bioRxiv2023

Disturbance increases functional diversity but decreases phylogenetic diversity of an arboreal tropical ant community

O_LITropical rainforest canopies host a highly diverse arthropod fauna, which contribute to ecosystem function through their functional (FD) and phylogenetic diversity (PD). While a lot of previous research has documented the severe negative impacts of disturbance on the FD and PD of ground invertebrate communities, our understanding of arboreal counterparts is limited. C_LIO_LIHere, we studied the effects of forest disturbance on an ecologically important invertebrate group, the ants, in a lowland rainforest in New Guinea. We exhaustively sampled 4000 m2 area of a primary and a secondary forest for canopy ants. We report > 2800 occurrences of 128 ant species in 852 trees, one of the most comprehensive arboreal collections to date. C_LIO_LITo test how ant PD and FD differ between the two forests, we constructed the ant species-level community phylogeny and measured 10 functional traits. Furthermore, we assessed by data exclusion the influence of species which were not nesting in individual trees (visitors) or only nesting (nesters), and of non-native species on FD and PD values. We expected that disturbance would decrease FD and PD in tree dwelling ants. We hypothesized that traits in primary forests would be more overdispersed due to the greater availability of ecological niches, while secondary forests would have stronger trait clustering due to a a stronger habitat filtering caused by more extreme microclimate. C_LIO_LIPrimary forests had higher species richness and PD than secondary forest. Surprisingly, we found higher FD in secondary forest. This pattern was robust even if we decoupled functional and phylogenetic signals or if non-native ant species were excluded from the data. Visitors did not contribute strongly to FD, but they increased PD. Community trait means further corroborate the functional distinctiveness of arboreal ants among secondary and primary forest, with almost all traits being impacted by disturbance and forest succession. C_LIO_LIWe conclude that the most plausible explanation is increased competition among closely related ant species in the secondary forest, which drives trait divergence. In the primary forest, abiotic habitat filters leads to more similar morphology and thus lower FD of phylogenetically more diverse ant assemblages. C_LI

ecology↗

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↗

bioRxiv2019

Mesoamerica is a cradle and the Brazilian Atlantic Forest is a museum of Neotropical butterfly diversity (Lepidoptera: Nymphalidae: Brassolini)

Regional species diversity is ultimately explained by speciation, extinction, and dispersal. Here we estimate dispersal and speciation rates of Neotropical butterflies to propose an explanation for their distribution and diversity of extant species. We focus on the tribe Brassolini (owl butterflies and allies): a Neotropical group that comprises 17 genera and 108 species, most of them endemic to rainforest biomes. We infer a robust species tree using the multispecies coalescent framework and a dataset including molecular and morphological characters. This formed the basis for three changes in Brassolini classification: 1) Naropina, SYN. NOV. is subsumed within Brassolina; 2) Aponarope, SYN. NOV. is subsumed within Narope; 3) Selenophanes orgetorix, COMB. NOV. is reassigned from Catoblepia to Selenophanes. By applying biogeographical stochastic mapping, we found contrasting species diversification and dispersal dynamics across rainforest biomes, which might be partly explained by the geological and environmental history of each bioregion. Our results reveal a mosaic of biome-specific evolutionary histories within the Neotropics, where butterfly species have diversified rapidly (cradles: Mesoamerica), have accumulated gradually (museums: Atlantic Forest), or have alternately diversified and accumulated (Amazonia). Our study contributes evidence from a major butterfly lineage that the Neotropics are a museum and cradle of species diversity.

evolutionary biology↗

bioRxiv2018

Species limits in butterflies (Lepidoptera: Nymphalidae): Reconciling classical taxonomy with the multispecies coalescent

Species delimitation is at the core of biological sciences. During the last decade, molecular-based approaches have advanced the field by providing additional sources of evidence to classical, morphology-based taxonomy. However, taxonomy has not yet fully embraced molecular species delimitation beyond threshold-based, single-gene approaches, and taxonomic knowledge is not commonly integrated to multi-locus species delimitation models. Here we aim to bridge empirical data (taxonomic and genetic) with recently developed coalescent-based species delimitation approaches. We use the multispecies coalescent model as implemented in two Bayesian methods (DISSECT/STACEY and BP&P) to infer species hypotheses. In both cases, we account for phylogenetic uncertainty (by not using any guide tree) and taxonomic uncertainty (by measuring the impact of using or not a priori taxonomic assignment to specimens). We focus on an entire Neotropical tribe of butterflies, the Haeterini (Nymphalidae: Satyrinae). We contrast divergent taxonomic opinion--splitting, lumping and misclassifying species--in the light of different phenotypic classifications proposed to date. Our results provide a solid background for the recognition of 22 species. The synergistic approach presented here overcomes limitations in both traditional taxonomy (e.g. by recognizing cryptic species) and molecular-based methods (e.g. by recognizing structured populations, and not raise them to species). Our framework provides a step forward towards standardization and increasing reproducibility of species delimitations.

evolutionary biology↗