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Biological Journal of the Linnean Society

Oxford University Press (OUP)

Preprints posted in the last 90 days, ranked by how well they match Biological Journal of the Linnean Society's content profile, based on 24 papers previously published here. The average preprint has a 0.02% match score for this journal, so anything above that is already an above-average fit.

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Distribution and heritable shell differentiation among populations of the sole lymnaeid snail across freshwater habitats of southern Patagonia

Muller Baigorria, M. A.; Abafatori, M.; Chapuis, E.; Juillet, N.; Faugere, D.; Jarne, P.; David, P.; Pointier, J.-P.; Hurtrez-Bousses, S.; Alda, P.; Bonel, N.

2026-05-16 evolutionary biology 10.64898/2026.05.14.725217 medRxiv
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AO_SCPLOWBSTRACTC_SCPLOWEnvironmental heterogeneity across freshwater systems often promotes phenotypic variation, yet disentangling environmentally induced variation from heritable differentiation remains a central goal in evolutionary ecology. We investigated the geographic distribution and morphological differentiation, and heritability of shell traits among populations of the freshwater lymnaeid snail Pectinidens diaphanus in Patagonia. Extensive field surveys across 196 freshwater sites revealed that the species occupies a broad range of lentic and lotic habitats and constitutes the only lymnaeid inhabiting southern Patagonia. While reproductive anatomical structures were conserved across populations, shell shape differed markedly among populations from contrasting habitat types, with population identity explaining nearly 50% of total shape variation. Populations from hydrologically unstable habitats (ponds and streams) exhibited more elongated shells and relatively smaller apertures, a pattern consistent with functional responses to hydroperiod variability and desiccation risk. To assess the heritability of this differentiation, we conducted a common-garden experiment across two generations. Shell shape differences between permanent- (lagoon) and temporary- (pond) habitat-derived populations persisted into the G2 generation reared under standardized laboratory conditions, indicating that the observed variation is not solely a response to local environmental conditions but includes a heritable component. Together, our findings demonstrate that P. diaphanus constitutes the sole lymnaeid across southern Patagonia, occupying a broader range than previously documented, and that populations show heritable shell differentiation potentially associated with contrasting freshwater habitats. By integrating large-scale biogeographic surveys with morphometric and experimental approaches, this study provides new insight into how habitat variation may contribute to ecological and evolutionary differentiation in freshwater gastropods.

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Scaling and ecomorphology of lagomorph body shape and appendicular skeleton

Huizenga, C.; Brice, N.; Law, C. J.

2026-05-12 evolutionary biology 10.64898/2026.05.07.723560 medRxiv
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The diversity of body shapes is one of the most prominent features of phenotypic variation in mammals. Yet, mammalian body shapes are poorly quantified and the underlying components contributing to its diversity as well as its relationship to other components of the skeleton are rarely tested. Here, we use lagomorphs (hares, rabbits and pikas) as a model system to (1) investigate which components of the skeleton contributed the most to body shape diversity, (2) examine the relationships between body shape and relative limb lengths, and (3) test how body size, ecotype, burrowing behavior, and locomotor mode influenced variation in lagomorph body shape and appendicular morphology. We quantified the body shape and functional proxies of the appendicular skeleton in 40 lagomorph species from osteological specimens held at museum collections. Using phylogenetic comparative methods, we found the relative length of the ribs and elongation or shortening of the thoracic and lumbar regions contributed the most to body shape evolution across lagomorphs. Second, we found that only leporids (hares and rabbits) exhibited a significant relationship between limb length and body shape, where more elongate species exhibit relatively shorter forelimbs and hindlimbs. Lastly, we found that models incorporating body size were the best predictors of lagomorph body shape and the majority of the appendicular traits, whereas models incorporating burrowing behavior and locomotor mode were largely poor fits. Broadly, these results indicate that larger lagomorphs tend to exhibit more robust body shapes with longer, more gracile forelimbs, whereas smaller lagomorphs tend to exhibit more elongate body shapes with shorter, more robust forelimbs. Overall, this work contributes to the growing understanding of mammalian body shape evolution and demonstrates the importance of not omitting body size in ecomorphological analyses.

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Female oviposition site selection influences hatching success and embryonic development in a Neotropical glass frog

Curaca-Fierro, J. S.; Goyes Vallejos, J.

2026-06-18 animal behavior and cognition 10.64898/2026.06.14.732161 medRxiv
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For oviparous animals, the decision of where to lay eggs is critical, as offspring remain sessile from oviposition through hatching and are thus unable to escape unfavorable conditions. Consequently, females are expected to select oviposition sites that benefit embryo development and survival. This may be particularly relevant for arboreal frogs, which typically lay eggs on leaves overhanging water, where embryos are exposed to predation, desiccation, and other risks until hatching. Yet studies directly linking maternal substrate choice to embryo survival remain scarce. Here, we examine how oviposition substrate influences embryo survival in the Emerald glass frog (Espadarana prosoblepon), a species in which females deposit eggs on multiple substrates, providing a rare opportunity to test how oviposition decisions affect reproductive success. Monitoring clutches in situ, we compared microclimatic conditions, hatching success, and sources of embryo mortality between the most used substrates: the spike moss Selaginella diffusa and leaves. Additionally, we conducted a two-choice experiment in semi-captivity to test whether females preferentially select one substrate over the other. Although microclimatic conditions did not differ between substrates, hatching success was significantly higher on S. diffusa, which also experienced less predation. In the two-choice experiment, all females laid their eggs on S. diffusa, and those clutches had higher hatching success and faster embryonic development rates than those on leaves. Together, these results support the hypothesis that non-random oviposition site selection in E. prosoblepon is driven by the maximization of embryo survival, demonstrating that substrate choice has measurable fitness consequences for the offspring.

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Socioecological differentiation and the evolution of brain size and synaptic architecture in predatory ants, Neoponera

Azorsa, F.; Traniello, J. F. A.

2026-07-02 evolutionary biology 10.64898/2026.06.27.735026 medRxiv
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Brain size and structure are hypothesized to be adaptively designed to satisfy the behavioral requirements of securing food and living socially. The importance of these socioecological and sociobiological selective forces in brain evolution is constantly debated. Socioecological divergence is striking in the Neotropical ant genus Neoponera: N. apicalis is a generalist solitary predator forming small colonies of ~100 whereas N. commutata colonies are approximately 10 times as large and workers pheromonally organize cooperatively raids only on Syntermes termite colonies. We interspecifically compared the size and structure of the compound eyes, size and number of antennal glomeruli, mosaic brain scaling and synaptic processing (microglomeruli-MG). Our results indicate that N. apicalis workers have a larger number of ommatidia, antennal lobe glomeruli, and allometrically larger antennal and optic lobes than N. commutata. These sensory traits were associated with differences in higher-order processing architectures in the mushroom body (MB) microglomeruli (MG). N. commutata workers had an allometrically larger MB, perhaps due to their socially complex chemical foraging communication, although MG density in N. apicalis was higher in both the MB lip and collar, regions associated with processing olfactory and visual information, respectively. The increase in MG density in N. apicalis may be associated with higher demands for navigation, learning, and memory, as well as a higher density of antennal lobe glomeruli to support prey odor discrimination. In contrast, N. commutata workers had larger ommatidia and antennal lobe glomeruli. Larger ommatidia correlate with their diurnal/nocturnal habits and a larger MB Our findings indicate that differences in behavioral performance demands associated with socioecological differentiation are reflected in variation in visual and olfactory system structure, brain size, mosaicism, and synaptic organization. Our results support both social and ecological brain hypothesis as drivers of mosaic brain evolution.

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Temporal persistence of admixed phenotypes in a damselfly hybrid zone

Velasquez-Velez, M. I.; Sanchez-Guillen, R. A.; Pulido-Rios, L.; Medina-Villarreal, A.; Cordero-Rivera, A.; Realpe, E.; Saldamando-Benjumea, C. I.

2026-05-31 evolutionary biology 10.64898/2026.05.29.728807 medRxiv
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In odonates, hybridization has been documented in several species pairs, yet the long-term persistence of hybrid phenotypes remains poorly understood, particularly in Neotropical systems. Here, we investigate a putative hybrid zone between Ischnura capreolus and Ischnura cyane, which overlap along an altitudinal gradient in the Colombian Andes. We combined temporal sampling across sympatric and allopatric localities with multilocus genetic analyses and geometric morphometrics of wings and male caudal appendages to evaluate patterns of admixture and phenotypic intermediacy. Morphometric analyses revealed pronounced differentiation between I. capreolus and I. cyane, especially in allopatric populations, whereas individuals from sympatric localities displayed increased morphological overlap and intermediate phenotypes. Genetic analyses based on nuclear and mitochondrial genes identified two main genetic clusters corresponding to the parental species, with evidence of admixture and shared haplotypes in sympatric localities. Patterns of differentiation varied among loci, with mitochondrial markers showing broader haplotype sharing than nuclear loci. Across sympatric localities, morphologically intermediate individuals remained consistently present through time, particularly in Anolaima, where they represented the dominant phenotype across sampling periods. These results support concordant morphological and genetic evidence consistent with persistent hybridization and introgression between I. capreolus and I. cyane in their zone of contact and suggest that incomplete reproductive isolation contributes to the long-term maintenance of admixed phenotypes in this hybrid zone.

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Extreme spatial plasticity in Italian wolves: ecological determinants of range size and movement patterns across heterogeneous landscapes

Cavazza, S.; Brogi, R.; Zanni, M.; Buelli, C.; Berzi, D.; Luccarini, S.; Costanzi, L.; Tirapelle, C.; Cappai, N.; Bongi, P.; Del Frate, M.; Vettorazzo, E.; Apollonio, M.

2026-06-16 animal behavior and cognition 10.64898/2026.06.12.731796 medRxiv
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As wolves (Canis lupus) recolonize diverse environments, the extent to which ecological and individual factors shape their spatial behaviour remains insufficiently understood. Italy, where wolf recovery began earlier than elsewhere in Europe and spans highly heterogeneous landscapes, offers an opportunity to explore patterns relevant to future European scenarios. We analysed variation in home range size and movement patterns using GPS data from 19 wolves monitored between 2019 and 2025 across five study areas representing a broad environmental gradient. Continuous-time movement modelling was used to estimate home range size and distance travelled. Home range size varied markedly among areas, with annual averages for pack-member wolves ranging from 35 to 185 km2. The smallest ranges occurred where wolves had been established for longer and/or where prey availability was higher, suggesting that both intraspecific competition and resource abundance influence space use. Wolves expanded their ranges in winter and in more urbanized landscapes. Pack members consistently had smaller home ranges than floaters, and among floaters, females showed slightly larger ranges than males. Distance travelled also differed significantly across areas (local averages 25-53 km/day) but showed no significant association with individual traits. Three dispersal events were documented, two of which resulted in successful pair formation and subsequent reproduction. Overall, our results provide a broad overview of wolf spatial behaviour in contemporary Italian landscapes and offer insights relevant to the management of wolf populations across Europe.

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Genomics Analysis Illuminates Morphology, Ecology, Phenology and Distribution of Two Cryptic Atrytonopsis Skippers (Hesperiidae: Hesperiinae)

Cary, S. J.; Doneski, S. M.; Zhang, J.; Cong, Q.; Grishin, N. V.

2026-06-19 evolutionary biology 10.64898/2026.06.16.732465 medRxiv
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The Hesperiine genus Atrytonopsis Godman, 1900, occurs broadly across the American Southwest. Atrytonopsis margarita (Skinner, 1913) and Atrytonopsis python (W. H. Edwards, 1882) have look-alike appearances, concurrent flights, and geographic distributions which converge in New Mexico. Their similar wing markings and intertwined taxonomic history has made it challenging to fully understand the identity and occurrence of each. Burns (2015) revealed differences in genitalia, clarifying that they are distinct species. Genomic DNA analysis of more than 100 specimens now illuminates their genetic uniqueness, phylogenetic relationship, field identification challenges and details of their geographic distributions.

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How Robust are Multispecies Coalescent Species Delimitations in Taxonomically Complex Systems? A Genomic Assessment Using Mediterranean Tethya Sponges

van der Sprong, J.; Cardone, F.; Hoehna, S.; Schaetzle, S.; Deister, F.; Erpenbeck, D.; Woerheide, G.; Vargas, S.

2026-07-05 evolutionary biology 10.64898/2026.07.04.735074 medRxiv
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Reliable species delimitation underpins biodiversity assessment but remains difficult for organisms with plastic morphology and few diagnostic characters. Multispecies coalescent (MSC) methods can delimit species from genomic data, yet they are rarely tested in taxonomically complex, marine invertebrate groups where they are arguably most needed. We used the three Mediterranean species of the genus Tethya, a rare, well-characterised system within the otherwise taxonomically difficult phylum Porifera-distinguished by multiple independent morphological and ecological characters-to evaluate how robust MSC-based delimitation is in such groups. Analysing 64 single-copy nuclear loci in BEAST2 and BPP, we compared constrained, hypothesis-testing approaches (BFD*, BFdriver, A10) with freer, heuristic ones (SPEEDEMON, A11), and examined their sensitivity to data type, clock model, priors, and the species-collapse threshold. All methods recovered the three recognised Mediterranean species, but the resolution of within-lineage structure was method-dependent. The hypothesis-testing approaches consistently supported six lineages, robustly across data types and model assumptions, whereas the heuristic approaches proved less stable. Configurations without a priori species hypotheses often failed to converge or were computationally intractable, a problem compounded by the relaxed clock. In SPEEDEMON the outcome changed with the collapse threshold. Because our system lacks an independent reference point to calibrate this threshold, any delimitation based on it is poorly constrained. We conclude that constrained, hypothesis-testing delimitation is the most robust and reproducible MSC approach, yielding a quantitative, model-based hypothesis that can be weighed against other lines of evidence to inform taxonomic decisions. By clarifying how these methods behave and how their outcomes should be interpreted, our study offers a practical guide for researchers working on comparably complex systems.

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The vocal repertoire of white-nosed coatis: structural features, temporal dynamics, and social variations

Grout, E. M.; Hass, C. C.; Minasandra, P.; Thomas, M.; Ashbury, A. M.; Ortega, J.; Crofoot, M. C.; Hirsch, B. T.; Strandburg-Peshkin, A.

2026-06-22 animal behavior and cognition 10.64898/2026.06.17.732959 medRxiv
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For many group living species, vocal signals are a vital form of communication needed to coordinate social behaviours. Investigating these processes first requires a detailed description of the species vocal repertoire. White-nosed coatis (Nasua narica), which forage and move together in forest habitats, are thought to rely on vocalisations to coordinate movements and maintain group cohesion. However, quantitative studies of their vocal repertoire are lacking. We examined the vocal repertoire of white-nosed coatis from wild populations in Panama and Arizona, USA, to gain a more comprehensive view of their calling behaviours. By combining traditional acoustic analyses with an unsupervised approach based on spectrogram structure, we characterised the diversity of calls in this species and described the temporal and structural features of their vocalisations. We identified 19 call types, with some of these calls emitted in multi-syllable call sequences or in fast succession. We found variability in call rates among group members, which may be driven by differences in social status within the group. In addition, our results indicate that white-nosed coatis likely have individually-recognisable vocalisations. This study provides a foundational description of the white-nosed coati vocal repertoire, laying the groundwork for future research on vocal communication in this species.

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Microplastics disrupt adaptive predator-induced behavioural plasticity in Rana dalmatina tadpoles

Herczeg, D.; Horvath, G.; Miko, Z.; Kovacs, B.; Hettyey, A.; Herczeg, G.

2026-06-16 animal behavior and cognition 10.64898/2026.06.11.731648 medRxiv
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The accumulation of microplastics (MP; plastic particles 1 m - 1 mm in diameter) in the environment is an increasing global concern. Although the physiological effects of MP on organisms, including humans, are increasingly documented, their impacts on behaviour are far less understood. Further, little is known about how MP affect adaptive phenotypic plasticity - the ability of a genotype to adaptively modify its phenotype in response to environmental cues. Anuran tadpoles are key models for studying phenotypic plasticity, with well-established evidence for predator-induced behavioural adjustments. Tadpoles typically reduce their activity and risk-taking when exposed to chemical cues released by predators, which has been proven to be adaptive. We investigated whether MP exposure from the fertilised egg stage alters the behaviour of agile frog (Rana dalmatina) tadpoles, and whether it interferes with their predator-induced behavioural plasticity. Tadpoles exposed to chemical cues from dragonfly larvae showed the expected antipredator response: these larvae showed significantly reduced movement activity and risk-taking. Although exposure to MP did not influence the behaviour of tadpoles that had not been exposed to predator cues, and also did not alter predator-induced changes in movement activity, it entirely abolished the predator-induced reduction in risk-taking. These results indicate that MP can compromise antipredator behaviour, increasing the vulnerability of individuals and natural populations to predation without causing visible developmental abnormalities. We recommend that future MP research targets behavioural traits with direct relevance to survival and reproduction, and examines how adaptive phenotypic plasticity is affected.

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Contrasting Mitochondrial Diversity of Endemic Corbicula Clams in Sulawesis Ancient Lakes: Phylogeography and Implications for Conservation

Muhammad, G.; Sumarto, B. K. A.; Dwiyanto, D.; Dewana, I. G. J.; Chadijah, A.; Astuti, S. S.; Sahidin, A.; von Rintelen, T.

2026-07-03 evolutionary biology 10.64898/2026.07.02.735996 medRxiv
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The global study of freshwater clams in the genus Corbicula is frequently confounded by invasive androgenetic lineages that experience mitochondrial DNA capture and clonal propagation. In contrast, the endemic Corbicula of Sulawesi's ancient lakes reproduce sexually, offering a uniquely reliable system for mitochondrial population genetics. This study provides the first population-genetic framework for two endemic species, Corbicula possoensis (Lake Poso) and C. linduensis (Lake Lindu), using the cytochrome c oxidase subunit I (COI) marker. We analysed 90 newly generated COI sequences from C. possoensis (six stations) and C. linduensis (three stations), integrated with reference sequences from GenBank, to assess genetic diversity, population structure, and phylogeographic patterns. Hierarchical AMOVA revealed deep divergence between the two lakes ({Phi}_CT = 0.607), consistent with prolonged independent isolation rather than a single shared vicariance event, as the two species do not form a sister pair in the phylogeny. Within Lake Poso, C. possoensis exhibited exceptionally high genetic diversity (24 haplotypes; h = 0.876; {pi} = 0.016) and pronounced micro-geographic structuring into three phylogeographic zones (North: Tentena and Siuri; East: Tando Nceppo and Busogo Beach; Southwest: Bancea and Pendolo), each characterised by distinct haplogroups. Remarkably, the maximum divergence between zones (K2P = 2.33%) approached the interspecific distance between C. possoensis and C. linduensis (K2P = 2.42%), indicating that within-lake mitochondrial divergence has reached near-interspecific levels. Conversely, C. linduensis displayed near-panmixia and extreme genetic depauperation (3 haplotypes; h = 0.246; {pi} = 0.0004), indicating long-term demographic stasis within a restricted habitat. The deep phylogeographic zonation in C. possoensis suggests that its discrete populations should be treated as separate Management Units (MUs) in conservation planning to preserve locally adapted gene complexes, whereas the severely depauperate gene pool of C. linduensis renders it critically vulnerable to environmental disturbance and invasive species, warranting urgent IUCN Red List assessment. To validate these mitochondrial boundaries and inform future conservation strategies, multi-marker and genome-wide reassessments are strongly recommended.

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Seeing and smelling mates: multimodal integration and visual gating of chemical cues in female mate-location behavior in the prawn Macrobrachium rosenbergii

da Costa, F. P.; Arruda, M. d. F.; Ribeiro, K.; Pessoa, D. M. d. A.

2026-05-15 animal behavior and cognition 10.64898/2026.05.12.723903 medRxiv
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Multimodal communication plays a central role in animal behavior, particularly when individuals must integrate information from different sensory channels to make rapid decisions. In aquatic environments, chemical and visual cues differ markedly in their spatial and temporal properties, such that chemical signals may be constrained by limited spatial resolution and temporal instability, potentially requiring visual information to reliably guide social decisions. In decapod crustaceans, both cue types are known to mediate reproduction, yet their relative contribution to mate-location behavior remains unclear. Here, we tested how visual and chemical cues from males influence mate-location behavior in females of the prawn Macrobrachium rosenbergii. Females were placed in a central arena and exposed to four stimulus configurations combining visual cues (a life-size photograph of a male or a control background) and chemical cues (water from an aquarium with or without a male). Attraction was quantified as the time spent in each half of the arena. Females showed no directional preference when exposed to chemical cues alone or when visual and chemical cues were spatially incongruent. In contrast, females spent significantly more time near male-associated stimuli only when visual and chemical cues were spatially congruent. These results indicate that mate-location behavior in this species depends on multimodal integration with a strong contextual dependence on visual information, which appears to gate the effectiveness of chemical cues. Spatially congruent multimodal signals are therefore necessary to guide orientation during mate search, suggesting that disruption of visual or chemical information in aquaculture systems may impair mating efficiency.

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Isolation and introductions disrupt the homogeneity of Argentine ants in Europe.

Narasimhan, S.; Csata, E.; Trapero-Camesella, L.; Cordonnier, M.; Mannino, G.; Casacci, L. P.; Witek, M.; Sanmartin-Villar, I.

2026-06-11 evolutionary biology 10.64898/2026.06.09.731050 medRxiv
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The introduction of alien species into new habitats stands as a pressing economic and ecological challenge but it is also essential for unveiling evolutionary processes. The introduction of the Argentine ant (Linepithema humile) led to the spread of a single supercolony through different continents and thousands of kilometres like in Europe, from Northwest Spain to Greece. It was assumed that the high invasiveness of the species mainly relied on the lack of agonism among colonies, an effect derived from its introduction. However, recent studies suggest that local adaptations and evolutionary divergence could involve the disruption of the Argentine ant "empire" into a mosaic of competitive colonies. We investigated how isolation affects population divergence by comparing mainland and island populations in two distant regions colonized in Spain and in Greece with morphology, agonism, cuticular hydrocarbons, and genetic diversity of ant workers. Our results showed that all colonies sampled belonged to the most spread supercolony in Europe (main supercolony) except one sampled in Crete (Heraklion; Greece), which resulted to be a supercolony not registered in Europe. The Heraklian supercolony showed a different chemical and genetic profile and hostile agonism towards the other Greek colonies. Differences between islands and mainland colonies belonging to the main supercolony were higher in Galiza than in Greece. Surprisingly, the chemical profile of the Cretan colony belonging to the main supercolony showed more similarity with the Galizan colonies than with the Greek mainland, suggesting that L. humile may have been introduced into Greece through this island instead of the mainland. Our study suggests that local adaptations in Argentine ant colonies can trigger competition between colonies. Our data strongly support the existence of a candidate supercolony which highlights either ongoing introductions of L. humile in Europe or gaps in our understanding of its metapopulation dynamics.

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Rapid speciation in Harlequin Toads (Anura: Bufonidae) endemic to the Sierra Nevada de Santa Marta

Ramirez-Romero, J. P.; Eslava, L.; Salgado-Roa, F.; Barros-Castaneda, J. D.; Barrientos, L. S.; Crawford, A. J.; Pardo-Diaz, C.; Rueda-Solano, L. A.; Salazar, C.

2026-05-29 evolutionary biology 10.64898/2026.05.28.728521 medRxiv
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The Neotropical genus Atelopus has experienced a drastic population decline in recent decades. Despite this, a knowledge gap remains regarding the conservation genetic status, phylogeography, and demographic history of most of its species, especially those endemic to areas with limited access like the Sierra Nevada de Santa Marta in Colombia. In this genomic study, we inferred phylogenetic relationships, demographic history, and gene flow among four endemic Atelopus morphospecies in the Sierra Nevada de Santa Marta-SNSM. Additionally, we compared the effective population size (Ne) estimates with the available population census data. NextRAD was used to obtain genomic data from 95 individuals collected at five sites in the SNSM and two in the Colombian Pacific (outgroup). The morphospecies recently diverged in a scenario without gene flow and were recovered as monophyletic. Their phylogenetic relationships were discordant, which is attributed to the presence of incomplete lineage sorting-ILS, which would also explain their shared ancestry among them. The lack of gene flow as well as the recent divergence times given by demography suggests a recent and rapid speciation. However, the reproductive isolation mechanisms that promote or maintain the species boundaries in this group remain unknown and require further investigation. We suggest that this process may have been influenced by the complex topography of the SNSM, traits such as high philopatry, low dispersal ability, and behavioral factors such as habitat preference or to factors related to genetic architecture that influenced the rapid formation of reproductive barriers among populations. Additionally, a pattern of population decline was observed around 200.000 years ago, with recent increases in three morphospecies. Despite the reduction in effective population size, no signs of inbreeding were detected. However, for A. laetissimus, the only species surveyed, the estimated value of Ne and its implications should be interpreted with caution. Ultimately, our findings reveal an evolutionary history shaped by a burst of diversification and abrupt reproductive isolation, highlighting how the resulting endemism and restricted genetic connectivity shape the unique evolutionary trajectory and vulnerability of this threatened montane species. Significance StatementThe mechanisms driving rapid speciation in montane ecosystems remain a central question in evolutionary biology. This study provides crucial genomic insights into the diversification of four endemic Atelopus species in an isolated Neotropical massif. We reveal a compelling evolutionary scenario where species diverge rapidly in absence of gene flow. This rapid speciation was likely facilitated by complex topography, environmental heterogeneity, and ecological and behavioral differences among species. Nevertheless, the evolutionary processes that limited gene flow among SNSM species have not yet been identified.

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Song evolution in light of ecosystem differences: exploring effects of urbanization and ecology on temporal and frequency traits of Spotted and Eastern towhee songs

Leon Du'Mottuchi, X.; Mesfin, M.; Creanza, N.

2026-06-02 evolutionary biology 10.64898/2026.05.30.728953 medRxiv
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The Eastern towhee (Pipilo erythrophthalmus) and Spotted towhee (Pipilo maculatus) are large New World sparrows found across North America. These two species were previously classified as a single species, the Rufous-sided towhee, which was separated in 1995 based on differences in plumage, geographic range, and song. Previous studies have shown that ecological factors, such as urbanization and climate, can affect learned vocalizations, particularly frequency-related song characteristics such as minimum frequency; however, most studies have been conducted on one species in a specific location. The extensive geographic distributions of these towhee species, along with their ample publicly available song-recording data, give us the unique opportunity to assess whether ecological pressures influence song variation. Here, we extract frequency-related song features from 2916 Spotted and Eastern towhee recordings and investigate whether geography and ecology--including recording location (latitude and longitude), tree cover, urbanization (nighttime lights, human population density, and distance to road), climate zones, elevation, and ecoregions--explain patterns in these song frequency variables of this sister-species pair. Our results show that geographic location, particularly longitude, contributes more strongly to the variation of song frequency features than urbanization, environment, and climate, suggesting that culturally transmitted differences in learned song, not ecology or anthropogenic structure, drive this variation. However, there was not a clear pattern of isolation-by-distance despite the geographic patterns that we found in their songs. Further, we did not find strong support for behavioral adaptation to habitat structure, but we did find a weak signal that urbanization was associated with increased frequency in Spotted towhees. Overall, we provide a detailed study on the interactions between ecology and song evolution, and provide new insight into the evolution of birdsong.

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Predator stimulus and habitat structure jointly shape antipredator behavior in frog species

Provete, D. B.; Citadini, J. M.; Gomes, F. R.

2026-05-29 animal behavior and cognition 10.64898/2026.05.26.728007 medRxiv
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When encountering a predator, prey must choose between immobility and flight, a decision shaped by predator proximity, habitat structure, body size, and evolutionary history. Despite extensive work on optimal escape theory, few studies have jointly modelled both the decision to flee and the intensity of flight in a phylogenetic comparative framework. Here, we used a Bayesian phylogenetic hurdle log-normal model to analyze the probability of immobility and the conditional jump distance in 534 trials from 89 males across 17 Neotropical frog species (seven families), exposed to a simulated snake predator in three arenas of varying structural complexity. Physical contact with the predator (touch) was the strongest predictor, reducing immobility probability from 94% to 15% and increasing jump distance by 26%. Habitat complexity increased immobility (bush > leaf litter > empty), and frogs in open arenas jumped 31% farther. Larger-bodied species were substantially more likely to remain immobile but did not jump farther, indicating that body size determines strategy choice rather than locomotor magnitude. Phylogenetic signal was strong for both components (Pagels {lambda} = 0.80 for jump distance; {lambda} = 0.71 for immobility), with phylogeny accounting for 69-75% of variance. Species random effects formed a phylomorphospace in which fossorial species showed highest immobility and arboreal/torrent species favored active escape. Substantial individual-level variation in immobility tendency (25% of variance) provides a heritable substrate for ongoing selection. Frog antipredator strategies are jointly shaped by ecological context and evolutionary history; hurdle models offer a powerful framework for decomposing behavioral decisions. LAY SUMMARYWhen a predator approaches, prey must choose between staying still or fleeing -- a decision depending on body size, habitat, and shared evolutionary history. We exposed 17 frog species to a snake in arenas of varying vegetation cover. Frogs fled when touched but mostly stayed still when only approached; larger species and those in dense vegetation were most likely to remain immobile. These antipredator strategies were strongly shaped by frogs shared ancestry.

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Static allometry of the horn and pronotal depression in adult Oryctes rhinoceros supports continuous nonlinearity, sexual dimorphism, and size-independent covariation

Okahara, M.; Niimi, T.; Morita, S.

2026-06-01 zoology 10.64898/2026.05.28.728611 medRxiv
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Exaggerated insect traits often show positive allometry, yet nonlinear scaling can reflect either continuous curvature or discrete morphs. Distinguishing between these alternatives is important because they imply different developmental and evolutionary scenarios. Using cross-sectional data from 1,000 adult Oryctes rhinoceros, we analyzed the static allometry of nine traits with pronotum width as the primary body-size proxy and body length for sensitivity analyses. Cross-validated comparisons among linear, continuous nonlinear, and two-component mixture models showed that continuous nonlinear models improved predictive performance over linear models for horn length and pronotal depression width in both sexes. By contrast, mixture regression did not outperform the best continuous model in either sex, providing no positive support for discrete within-sex dimorphism under the tested model set. After accounting for body size, horn length remained positively associated with pronotal depression width, indicating size-independent covariation. These associations were retained when body length was used instead of pronotum width, supporting robustness to body-size proxy choice. Together, these results support continuous nonlinear adult scaling of the horn and pronotal depression in O. rhinoceros and indicate covariation not attributable solely to body size under the tested model set.

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Comparative morphology of silk-spinning systems in amphipods

McKim, S.; Turner, T. L.

2026-05-12 evolutionary biology 10.64898/2026.05.07.723571 medRxiv
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Silk glands have been found in two groups of amphipods: the Corophiida and the Ampeliscidae. The silk glands in Ampeliscidae, however, have yet to be examined in detail. Here we report, for the first time, the morphology and distribution of pereopodal glands in the Ampeliscidae, in non-thread producing Synopiidae, and in the Paragammaropsidae. In the Ampeliscidae we found two gland types distributed throughout all pereopods which have the ability to create threads. Pereopods three and four have additional silk extrusion morphology at the tip of the dactylus in which silk is transformed into semi-cylindrical threads used for building domiciles. Synopiid outgroup species have one of the gland types but lack silk extrusion morphology. Using ancestral state reconstruction analysis, we find that glands in the Synopiidae are likely ancestral and hypothesize that silk glands in Ampeliscidae are derived from these ancestral glands. Silk-spinning pereopods in the Paragammaropsidae had similarities with both Corophiida and Ampeliscidae but had distinctions. Ampeliscidae silk-spinning systems bear surprising resemblance to the Corophiida which presents one to reconsider the taxonomic placement of Ampeliscidae and the origins of silk-spinning in amphipods. This is the first comprehensive study on the glandular systems of Ampeliscidae, Synopiidae, and Paragammaropsidae using advanced microscopy, providing pertinent morphological data to the study of arthropod silk gland evolution and complex traits.

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Timing matters: age-dependent female receptivity and preference patterns

Yan, L.; Elias, D. O.

2026-06-29 animal behavior and cognition 10.64898/2026.06.23.734080 medRxiv
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The timing of reproductive processes can influence how mating interactions shape reproductive strategies. We examined how female age correlates with receptivity and preference in the jumping spider Habronattus formosus, a system characterized by elaborate male courtship and strong female choice. To test how receptivity changes across the post-maturation period, we paired females of different ages with males and quantified male courtship displays and mating outcomes. Females were not receptive immediately after maturation and instead exhibited receptivity primarily 2-3 weeks after maturation. We further found that the full male courtship predicted mating success in older females, whereas only the second stage of the courtship predicted success in younger females. The delay in female receptivity is consistent with expectations for strong female choice systems, where females may have a greater opportunity to evaluate male courtship before mating. These results highlight the importance of age-dependent changes in female reproductive state and suggest that the timing of receptivity may shape how different components of male courtship influence mating success.

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Social control, not service quality, explains fast growth in the cleaner wrasse Labroides dimidiatus.

Pessina, L.; Bshary, R.

2026-05-19 animal behavior and cognition 10.64898/2026.05.16.725469 medRxiv
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Interactions between cleaner fish Labroides dimidiatus and client fish, from which cleaners remove ectoparasites and mucus, represent a textbook example of mutualism involving sophisticated strategic decision-making. However, cleaners must also face intraspecific social challenges within a size-based hierarchy, where the largest females may eventually change sex and become males with higher reproductive rates. Following 540 individuals over 11 months, we found that, contrary to expectations, slow-growing females spent more time cleaning and cheated more frequently, without causing more negative client responses than fast-growing females did. Instead, variation in growth was best explained by social factors: fast-growing individuals experienced reduced social control, while slow growers spent more time in proximity to dominant individuals. As there was no evidence that spawning activity affected growth patterns, it appears that fast growth as a viable strategy for becoming a male largely depends on the lack of control by dominants.