Biology subjects

Herrel, A.

Publications and source records attributed to Herrel, A..

At least 19 recordsLinked to original sources

bioRxiv2026

A Novel Computational Framework to Investigate the Biomechanics of Burrowing

Head-first burrowing likely imposes strong mechanical demands on the skull of fossorial vertebrates, but how cranial tissues respond to these loads remains obscure. Here, we developed a computational framework integrating the discrete element method (DEM) with finite element analysis (FEA) to investigate cranial biomechanics during burrowing. We used Blanus cinereus, a round-headed amphisbaenian, as a model to establish DEM predictions that were validated and calibrated against in vitro experiments across three substrates and compared with in vivo push forces. DEM-calibrated particle contact forces were then transferred to an anatomically detailed finite element model to quantify cranial loading. The analyses suggested that soft tissues accommodated substantial deformation, while the skull transmitted burrowing loads from the premaxilla towards the occipital region. Cranial sutures further redistributed stresses and strains within the skull. This validated DEM-FEA framework links particle-scale substrate interactions with internal cranial biomechanics and provides a quantitative approach for investigating the mechanics of head-first burrowing across fossorial vertebrates.

bioengineering↗

bioRxiv2026

Elevated genetic load is associated with reduced bite force in insular lizard populations

Demographic history can profoundly shape genomic variation by altering levels of inbreeding, the strength of genetic drift, and the efficacy of purifying selection, thereby influencing both the accumulation and expression of deleterious variation. Yet the extent to which these genomic consequences translate into organismal performance remains poorly understood. Here, we investigated how contrasting demographic histories shape genomic diversity, inbreeding, genetic load, and performance across mainland and insular populations of the Italian wall lizard, Podarcis siculus. Using whole-genome sequence data from 263 wild-caught individuals and field-based performance measurements from 134 lizards, we characterised population structure and demographic history, quantified genomic diversity, runs of homozygosity, and deleterious variation, and tested associations between genomic inbreeding and performance traits. Island populations showed reduced genetic diversity, increased autozygosity, and elevated genetic load relative to mainland populations, consistent with stronger genetic drift and reduced efficacy of purifying selection. Patterns of deleterious variation were consistent with purging of strongly, but not mildly, deleterious variants. Greater genomic autozygosity was associated with reduced bite force in island lizards, a pattern consistent with inbreeding depression, whereas sprint speed, sperm quality, immunocompetence, and spatial cognition showed no significant association with inbreeding. Genome-wide mapping of runs of homozygosity further identified multiple genomic regions associated with reduced bite performance. Together, these results show how demographic history shapes genomic diversity, homozygosity, and deleterious variation in wild vertebrate populations, while revealing that phenotypic consequences can be trait-specific, a pattern that may reflect differences in genetic architecture and the extent of environmental influence on trait variation.

evolutionary biology↗

bioRxiv2026

Coordinated beak-tongue mechanics enable dexterous seed manipulation in songbirds

Dexterous manipulation of objects relies on precise coordination between anatomical elements. In seed-eating birds, seeds are manipulated and dehusked using both the beak and tongue, but the functional roles and coordination of these structures remain unresolved. Here, we quantified the 3D movements of the upper beak, lower beak, tongue, and seed in a hard-biting and a weak-biting songbird species using X-ray Reconstruction of Moving Morphology (XROMM) and measured contractile properties of their primary jaw muscles. We show that the tongue serves as the main tool for seed rotation, transport, and stabilization. Multi-dimensional, high-frequency movements of the upper and lower beaks reveal that efficient seed processing depends on high mobility of the kinetic avian skull. Strong and weak biters differ in feeding kinematics and jaw muscle speeds, suggesting ecological specialization of cranial mechanics. The complexity, precision, and tight coordination of beak and tongue motions show that the avian cranium rivals the dexterity of the primate hand despite limited degrees of freedom.

ecology↗

bioRxiv2025

Brain size reduction in dogs was already established at least by the Late Neolithic of western Europe, 5,000 years ago

The timing and causes of brain size reduction in domestic dogs remain uncertain. Using endocasts volume as a proxy for brain size, this study provides a first insight into long-term brain size evolution in the wolf-dog lineage. We compared endocranial volumes of 185 modern and 22 prehistoric wolves and dogs ranging from Western Europe to Australia, and spanning the Pleniglacial (35 Ky BP) to the Late Neolithic (5 Ky BP). Our results reveal that Pleistocene so called "protodogs" show no brain size reduction compared to coeval Pleistocene wolves. Instead, we observed a slightly larger relative endocranial volume in the 35,000-year-old protodog from Goyet, which could suggest increased behavioural flexibility in the presence of humans. This hypothesis needs to be tested further. In contrast, Late Neolithic dogs show a drastic 46% brain size reduction with an endocranial volumes comparable to modern small terrier and toy breeds. The anxious and wary temperaments of these Late Neolithic dogs, induced by the brain tissue reorganization associated with such a size reduction, could have served an alerting purpose, among the many other potential roles dogs could have played within this Late Neolithic socio-ecosystems.

zoology↗

bioRxiv2025

Anatomical description of the jaw muscles and theoretical bite force assessment in South-American opossums using manual and virtual dissection methods.

Marsupials (Marsupialia, Mammalia) represent a clade with highly varied ecologies. This is particularly true for opossums (South American marsupials), which are difficult to observe and collect. Consequently, few studies have focused on their bite force and the muscles of their masticatory apparatus, and there exists only scant information about the diet of certain species. Here we describe the masticatory muscles of several previously unstudied opossum species including Caenolestes fuliginosus, Dromiciops gliroides, and Monodelphis touan. We calculate the bite force of these species using data from both manual and virtual dissections and compare their theoretical bite forces with literature data. Additionally, we explore the differences between manual and virtual dissection to determine muscle PCSA (Physiological Cross-Sectional Area). We tried two virtual methods (VPDE: "virtual physiological data estimating method" and SM: "slicing muscle method") to calculate the PCSA, determine the differences induced by the inter-fiber void in the virtual volume, and calculate a correction post-treatment with the contrast agent. The results highlighted variation in the position of the muscular attachments of the M. zygomaticomandibularis, whose insertion area is the largest in Monodelphis touan and the smallest in Caenolestes fuliginosus. The bite forces are coherent with estimates from the literature suggesting that the biomechanical model is reliable. The comparison between manual and virtual dissection showed that while virtual dissection allows an overall description of the masticatory muscles, it is more complex to accurately describe the different subdivisions of the muscle bundles. Virtual dissection data could potentially complete manual dissection data with the association of the VPDE method, the exclusion of inter-fiber voids, and a correction for the treatment with contrast-agents.

zoology↗

bioRxiv2025

Divergent paths, convergent heads: morphological adaptation of head shape to ecological niches in snakes.

Morphological convergence--where distantly related species evolve similar traits in response to shared ecological pressures--is a hallmark of adaptive evolution. In snakes, head shape is a key functional trait linked to habitat use and diet, yet the extent and drivers of its convergence across habitat types and diet remain poorly understood. Here, we used geometric morphometric analyses to analyze dorsal head shape from photographs of over 266 snake species, representing [~]5% of global snake diversity. Our analyses reveal that head shape variation is only weakly structured by phylogeny and allometry, but shaped by ecological specialization. Morphospace patterns reflect distinct differences in species with different ecologies: fossorial species exhibit compact, posteriorly wider heads suited for burrowing; aquatic species show streamlined profiles for hydrodynamic efficiency; and arboreal snakes tend to possess elongated heads for maneuvering in complex habitats. Terrestrial and semi-fossorial snakes display broad morphospace overlap and an elevated shape disparity highlighting morphological versatility. While diet covaried significantly with head shape, species with similar diets did not exhibit strong morphological convergence. These findings underscore the dominant role of ecology over phylogeny in shaping the evolution of head shape in gape-limited predators like snakes.

evolutionary biology↗

bioRxiv2025

From head to tail: does habitat use drive morphological variation in snakes?

Convergence is a hallmark of adaptive evolution, yet its extent across different taxa remains unclear. Snakes, with over 4,100 species worldwide, provide a unique model to explore how habitat use influences morphology in a group with an at first sight uniform body plan. We here quantified body, head, and tail shape in over 400 species ([~]10% of the global snake diversity) to assess the role of habitat use in shaping morphological variation. Our results reveal significant differences across ecological groups, with habitat use acting as a key driver of phenotypic diversity. Terrestrial species display the highest morphological diversity in contrast to other habitat groups which appear morphologically specialized. For example, whereas arboreal and semi-arboreal species exhibit elongated heads and slender necks, aquatic and semi-aquatic snakes share streamlined bodies and narrow heads. Fossorial and semi-fossorial species, on the other hand, have compact bodies. Surprisingly, morphological convergence remains limited to arboreal, semi-arboreal, and terrestrial habitat groups. Thus, despite strong functional constraints, evolutionary convergence in fossorial and aquatic species is weak, indicating multiple adaptive solutions rather than a single morphological trajectory. Morphological disparity patterns show that non-specialist ecologies generally exhibit greater disparity than highly specialized ones, in accordance with the need of these species to move in different habitats. Our findings underscore the role of ecological constraints in shaping snake morphology and highlight the complexity of adaptation beyond strict convergent evolution. Lay summaryThe living environment of a species has been suggested to play a key role in shaping its morphology. Our study aimed to understand whether and how habitat use has influenced the evolution of external body and head shape in snakes. To do so, we gathered a comprehensive dataset of morphological traits from 436 snake species. Our findings reveal that species living in different habitats are morphologically distinct with species living in similar habitats displaying similar morphologies despite the conserved morphology of snakes.

evolutionary biology↗

bioRxiv2024

CO-OPTION OF NECK MUSCLES SUPPORTED THE VERTEBRATE WATER-TO-LAND TRANSITION

A major event in vertebrate evolution was the separation of the skull from the pectoral girdle and the acquisition of a functional neck, transitions which required profound developmental rearrangements of the musculoskeletal system. The neck is a hallmark of the tetrapod body plan and allows for complex head movements on land. While head and trunk muscles arise from distinct embryonic mesoderm populations, the origins of neck muscles remained elusive. Here, we combine comparative embryology and anatomy to reconstruct the mesodermal contribution to neck evolution. We demonstrate that head/trunk-connecting muscle groups have conserved mesodermal origins in fishes and tetrapods and that the neck evolved from muscle groups present in fishes. We propose that expansions of mesodermal populations into head and trunk domains during embryonic development underpinned the emergence and adaptation of the tetrapod neck. Our results provide evidence for exaptation of archetypal muscle groups in ancestral fishes, which were co-opted to acquire novel functions adapted to terrestrial lifestyle.

evolutionary biology↗

bioRxiv2024

Gekko gecko as a model organism for understanding aspects of laryngeal vocal evolution

The ability to communicate through vocalization plays a key role in the survival of animals across all vertebrate groups. While avian reptiles have received much attention relating to their stunning sound repertoire, non-avian reptiles have been wrongfully assumed to have less elaborate vocalization types and little is known about the biomechanics of sound production and their underlying neural pathways. We investigated alarm calls of Gekko gecko using audio and cineradiographic recordings of their alarm calls. Acoustic analysis revealed three distinct call types: a sinusoidal call type (type 1), a train-like call type, characterized by distinct pulse trains (type 3), and an intermediary type, which showed both sinusoidal and pulse train components (type 2). Kinematic analysis of cineradiographic recordings showed that laryngeal movements differ significantly between respiratory and vocal behavior: during respiration, animals repeatedly moved their jaws to partially open their mouths, which was accompanied by small glottal movements. During vocalization, the glottis was pulled back, contrasting with what has previously been reported. In-vitro retrograde tracing of the nerve innervating the laryngeal constrictor and dilator muscles revealed round to fusiform motoneurons in the hindbrain-spinal cord transition ipsilateral to the labeled nerve. Taken together, our observations provide insight into the alarm calls generated by G. gecko, the biomechanics of this sound generation and the underlying organization of motoneurons involved in the generation of vocalizations. Our observations suggest that G. gecko may be an excellent non-avian reptile model organism for enhancing our understanding of the evolution of vertebrate vocalization. Summary StatementInvestigation of Gekko gecko alarm calls revealed distinct call types, during which the larynx is being pulled back by muscles innervated by motoneurons located in the hindbrain.

animal behavior and cognition↗

bioRxiv2023

The pharynx of the iconic stem-group chondrichthyan Acanthodes (Agassiz, 1833) revisited with micro computed tomography.

Acanthodes has long been the primary source of information on the pharyngeal skeleton of acanthodians, a stem-group chondrichthyan grade. Because of this its anatomy has played an outsized role in attempts to understand the evolution of the jawed vertebrate pharynx and the clade as a whole. However, the anatomy of the pharynx of Acanthodes remains poorly understood and subject to several competing interpretations. We use computed tomography (CT) to image the articulated pharyngeal skeletons of three specimens of Acanthodes confusus from Lebach, Germany. Acanthodes had a melange of osteichthyan-like and chondrichthyan-like morphologies in its pharyngeal skeleton. Like other stem-chondrichthyans, Acanthodes had a basihyal with no hypohyals, and four pairs of posteriorly oriented pharyngobranchials. Like osteichthyans, Acanthodes possessed an interhyal, but lacked the separate infra- and supra-pharyngobranchial elements present in osteichthyans and the crown-chondrichthyan Ozarcus. Using this new data we build and animated a digital 3D model of the pharyngeal endoskeleton in Acanthodes, showing that the jaws would have swung outwards during the opening cycle, increasing the anteriorly facing area of the gape for suspension feeding. These new data provide a more definitive picture of the anatomy of a taxon that has long been of great significance in early vertebrate palaeontology.

paleontology↗

bioRxiv2022

Armed to the teeth: the underestimated diversity in tooth shape in snakes and its relationship to feeding constraints

Teeth are one of the most studied hard tissues in vertebrates. Their structure, composition and shape are related to dietary specialization in many species. At first glance, snake teeth all look similar; conical, sharp, curved. Yet, snakes, like other vertebrates, have very diverse diets that may have affected their shape. We compared the morphology of the teeth of 63 species that cover both the phylogenetic and dietary diversity of snakes. We predicted that prey properties play a role in shaping snakes teeth along with their feeding behavior. Limblessness combined to the peculiar feeding behavior of snakes impose strong functional constraints on their teeth, especially during arboreal or aquatic feeding. Our results show that prey hardness, foraging substrate and the main feeding constraint are drivers of tooth shape, size, and curvature. We highlight two main morphotypes: long, slender, curved with a thin layer of hard tissue for snakes that need a good grip on their prey and short, stout, less curved teeth in snakes that eat hard or long prey. Our study demonstrates the diversity of tooth morphology in snakes and the need to investigate the underlying functional implications to better understand the evolution of teeth in vertebrates.

evolutionary biology↗

bioRxiv2022

Measuring the 3D wake of swimming dice snakes using volumetric velocimetry

Experimental observations of the 3-dimensional wake of swimmers are scarce. This study provides the first experimental measurements of the three-dimensional vortical structures of an anguilliform swimmer. A volumetric velocimetry (DDPTV) setup was used to quantify the wake of freely swimming dice snakes (Natrix tessellata) in a tank. Multiple swimming trials were recorded with three snakes swimming at a forward swimming speed ranging from 0.2 to 0.8L.s-1. The volumetric measurements show that during swimming, Natrix tessellata sheds vortex tubes from different parts of its body with oscillating lateral undulations. The vortex tubes are then linked to form a hairpin-like structure as predicted by computational fluid dynamic analyses. Quantitative measurements show that the vortex size gradually increases over time. The vortex circulation decreases after attaining a maximum. These results provide an experimental demonstration that the 3D wake structure of an anguilliform swimmer takes well organized dynamic shapes as predicted and provide baseline data for future studies comparing the wake structure of snakes with different locomotor ecologies.

zoology↗

bioRxiv2022

The mammalian forelimb diversity as a morphological gradient of increasing evolutionary versatility

Vertebrate limb morphology often reflects the environment, due to variation in locomotor requirements and other ecological traits. However, proximal and distal limb segments may evolve differently to each other, reflecting an anatomical gradient of functional specialization that has been suggested to be impacted by the timing of bone condensation during ontogeny. Here we explore whether the temporal sequence of bone condensation predicts variation in the capacity of evolution to generate morphological diversity between proximal and distal forelimb segments across more than 600 species of mammals. Our findings are consistent with the hypothesis that late developing distal limb elements should display greater morphological variation than more proximal limb elements, which condense earlier during morphogenesis. Distal limb elements, belonging to the autopod, not only exhibit higher diversity of form, but are also more integrated and, on average, show greater evolutionary versatility than intermediate and upper limb segments. Our findings indicate that the macroevolutionary patterns of proximal and distal limb segments are not the same, suggesting that strong functional selection, combined with the higher potential of development to generate variation in more distal limb structures, facilitate the evolution of high autopodial disparity in mammals.

evolutionary biology↗

bioRxiv2022

Evidence of attack deflection suggests adaptive evolution of wing tails in butterflies

Predation is a powerful selective force shaping many behavioural and morphological traits in prey species. The deflection of predator attacks from vital parts of the prey usually involves the coordinated evolution of prey body shape and colour. Here, we test the deflection effect of hindwing tails in the swallowtail butterfly Iphiclides podalirius. In this species, hindwings display long tails associated with a conspicuous colour pattern. By surveying the wings within a wild population of I. podalirius, we observed that wing damage was much more frequent on the tails. We then used a standardised behavioural assay employing dummy butterflies with real I. podalirius wings to study the location of attacks by great tits Parus major. Wing tails and conspicuous coloration of the hindwings were struck more often than the rest of the body by birds. Finally, we characterised the mechanical properties of fresh wings and found that the tail vein was more fragile than the others, suggesting facilitated escape ability of butterflies attacked at this location. Our results clearly support the deflective effect of hindwing tails and suggest that predation is an important selective driver of the evolution of wing tails and colour pattern in butterflies.

evolutionary biology↗

bioRxiv2020

Do substrate type and gap distance impact gap-bridging strategies in arboreal chameleons?

Chameleons are well-equipped for an arboreal lifestyle, having zygodactylous hands and feet as well as a fully prehensile tail. However, to what degree tail use is preferred over autopod prehension has been largely neglected. Using an indoor experimental set-up, where chameleons had to cross gaps of varying distances, we tested the effect of substrate diameter and roughness on tail use in Chamaeleo calyptratus. Our results show that when crossing greater distances, C. calyptratus is more likely to use its tail for additional stability. The animals were able to cross greater distances (up to 1 75 times the shoulder-hip length) on perches with a rougher surface. We saw that depending on the distance of the gap, chameleons would change how they use their prehensile tails when crossing. With shorter gaps the tails either do not touch, or only touch the perch without coiling around it. With larger distances the tails are fully coiled around the perch, and with the largest distances additionally they reposition the hind legs, shifting them towards the end of the perch. Males were able to cross relatively greater distances than females, likely due to their larger size and strength.

animal behavior and cognition↗

bioRxiv2020

The morphology and evolution of chondrichthyan cranial muscles: a digital dissection of the elephantfish Callorhinchus milii and the catshark Scyliorhinus canicula.

The anatomy of sharks, rays, and chimaeras (chondrichthyans) is crucial to understanding the evolution of the cranial system in vertebrates, due to their position as the sister group to bony fishes (osteichthyans). Strikingly different arrangements of the head in the two constituent chondrichthyan groups - holocephalans and elasmobranchs - have played a pivotal role in the formation of evolutionary hypotheses targeting major cranial structures such as the jaws and pharynx. However, despite the advent of digital dissections as a means of easily visualizing and sharing the results of anatomical studies in three dimensions, information on the musculoskeletal systems of the chondrichthyan head remains largely limited to traditional accounts, many of which are at least a century old. Here we use synchrotron tomography acquire 3D data which we used to carry out a digital dissection of a holocephalan and an elasmobranch widely used as model species: the elephantfish, Callorhinchus milii, and the small-spotted catshark, Scyliorhinus canicula. We describe and figure the skeletal anatomy of the head, labial, mandibular, hyoid, and branchial cartilages in both taxa as well as the muscles of the head and pharynx. We make new observations, particularly regarding the branchial musculature of Callorhinchus, revealing several previously unreported or previously ambiguous structures. Finally, we review what is known about the evolution of chondrichthyan cranial muscles from their fossil record and discuss the implications for muscle homology and evolution, broadly concluding that the holocephalan pharynx is likely derived from a more elasmobranch-like form. This dataset has great potential as a resource, particularly for researchers using these model species for zoological research, functional morphologists requiring models of musculature and skeletons, as well as for palaeontologists seeking comparative models for extinct taxa.

evolutionary biology↗

bioRxiv2020

Exploring the functional meaning of head shape disparity in aquatic snakes.

Phenotypic diversity, or disparity, can be explained by simple genetic drift or, if functional constraints are strong, by selection for ecologically relevant phenotypes. We here studied phenotypic disparity in head shape in aquatic snakes. We investigated whether conflicting selective pressures related to different functions have driven shape diversity and explore whether similar phenotypes may give rise to the same functional output (i.e. many-to-one mapping of form to function). We focused on the head shape of aquatically foraging snakes as they fulfil several fitness-relevant functions and show a large amount of morphological variability. We used 3D surface scanning and 3D geometric-morphometrics to compare the head shape of 62 species in a phylogenetic context. We first tested whether diet specialization and size are drivers of head shape diversification. Next, we tested for many-to-one mapping by comparing the hydrodynamic efficiency of head shapes characteristic of the main axis of variation in the dataset. We 3D printed these shapes and measured the forces at play during a frontal strike. Our results show that diet and size explain only a small amount of shape variation. Shapes did not functionally converge as more specialized aquatic species evolved a more efficient head shape than others. The shape disparity observed could thus reflect a process of niche specialization under a stabilizing selective regime.

evolutionary biology↗

bioRxiv2019

Tooth Shape Adaptations in Aglyphous Colubrid Snakes Inferred from 3D Geometric Morphometrics and Finite Element Analysis

To date there are few detailed and quantitative studies investigating the evolution of the tooth shape and function in Aglyphous snakes in relation to diet. To study dental adaptations to diet, a lineage that is of particular interest due to its large range of adult body sizes, is the one including dwarfed snakes of the genus Eirenis and their immediate sister group, whip snakes of the genus Dolichophis. A considerable evolutionary decrease in the size is observed from a Dolichophis-like ancestor to the miniature Eirenis, coupled with a considerable shift in their diet from a regime consisting mainly of endotherms with endoskeleton to ectotherms bearing a hard exoskeleton. Maxilla, palatine, pterygoid and dentary teeth were examined in an adult and a juvenile of Dolichophis schmidti, one Eirenis punctolineatus and one Eirenis persicus. 3D Geometric Morphometrics comparison revealed maxilla and palatine teeth of the E. persicus are blunt and conical shape while those teeth are sharp and elongated in E. punctatolineatus as well as the adult and juvenile D. schmidti. A similar difference could be noted for the pterygoid teeth. In contrast, the dentary teeth are not as different among the examined snakes. Blunt and conically shaped teeth, as observed in E. persicus, seem to be more adapted for biting hard bodied, arthropod prey, while sharp and elongated teeth in Dolichophis and E. punctatolineatus, are specialized for puncturing endotherm prey. The results of a finite element analysis confirms that during biting a hard bodied prey, the generated stresses in E. persicus tooth is mostly confined to the tip of the tooth and mostly well below the von Mises yield criterion the tooth. In contrary, D. schmidti tooth appears less well suited for biting a hard prey since the generated stresses widely distribute across the tooth with values roughly 2 to 3 times higher than the von Mises yield criterion of the tooth. A lower degree of specialization that was observed among the dentary teeth in the examined snakes suggest a similar functional constraint in pushing the prey against the upper tooth rows.

evolutionary biology↗