Rensch's rule: linking intraspecific to evolutionary allometry.

Sexual dimorphism describes phenotypic differences between the sexes; the most prominent of which is sexual size dimorphism (SSD). Rensch's rule (RR) is an allometric trend in which SSD increases in male-larger taxa and decreases in female-larger ones. Covariation between a trait and overall size within and across species can both be affected by sexual and natural selection. Thus, intraspecific allometric variation could influence the expression of RR. Here we used computer simulations to dissect how RR emerges under specific allometric patterns of intraspecific sexual differentiation in a trait. We found that sexual differentiation in static allometric slopes is the main determinant of RR. Based on our findings, RR and its converse can manifest in both body size and other traits. As a realistic showcase, we also examined RR and static allometry of different body parts in Mediterranean green lizards to establish whether intraspecific and evolutionary allometry are linked. Here, we identified RR and its converse for different traits, where the amount of sexual differentiation in static allometric slopes within species had a significant contribution to RR. Integrating the simulations and the empirical case we corroborate that sexual differentiation in static allometric slopes is a major parameter affecting evolutionary allometry.

The Non-Dereliction in Evolution: Trophic Specialisation Drives Convergence in the Radiation of Red Devil Spiders (Araneae: Dysderidae) in the Canary Islands.

Natural selection plays a key role in deterministic evolution, as clearly illustrated by the multiple cases of repeated evolution of ecomorphological characters observed in adaptive radiations. Unlike most spiders, Dysdera species display a high variability of cheliceral morphologies, which has been suggested to reflect different levels of specialization to feed on isopods. In this study, we integrate geometric morphometrics and experimental trials with a fully resolved phylogeny of the highly diverse endemic species from the Canary Islands to 1) quantitatively delimit the different cheliceral morphotypes present in the archipelago, 2) test their association with trophic specialization, as reported for continental species, 3) reconstruct the evolution of these ecomorphs throughout the diversification of the group, 4) test the hypothesis of convergent evolution of the different morphotypes, and 5) examine whether specialization constitutes a case of evolutionary irreversibility in this group. We show the existence of 9 cheliceral morphotypes and uncovered their significance for trophic ecology. Further, we demonstrate that similar ecomorphs evolved multiple times in the archipelago, providing a novel study system to explain how convergent evolution and irreversibility due to specialization may be combined to shape phenotypic diversification in adaptive radiations.

Head Shape Heritability in the Hungarian Meadow Viper Vipera ursinii rakosiensis.

Understanding heritability patterns in functionally relevant traits is a cornerstone for evaluating their evolutionary potential and their role in local adaptation. In this study, we investigated patterns of heritability in the head shape of the Hungarian meadow viper (Vipera ursinii rakosiensis). To this end, we used geometric morphometric data from 12 families composed of 8 mothers, 6 fathers and 221 offspring, bred in captivity at the Hungarian Meadow Viper Conservation Centre (Hungary). We separately evaluated maternal and paternal contributions to the offspring phenotype, in addition to additive genetic effects, all determined using a mixed animal model. Our results indicate a strong genetic and maternal contribution to head shape variations. In contrast, the paternal effects-which are rarely evaluated in wild-ranging species-as well as residual environmental variance, were minimal. Overall, our results indicate a high evolutionary potential for head shape in the Hungarian meadow viper, which suggests a strong contribution of this ecologically important trait in shaping the ability of this endangered species to adapt to changing conditions and/or habitats. Furthermore, our results suggest that maternal phenotypes should be carefully considered when designing captive breeding parental pairs for reinforcing the adaptive capacity of threatened populations, whereas the paternal phenotypes seem less relevant.

The Vitruvian spider: Segmenting and integrating over different body parts to describe ecophenotypic variation.

Understanding what drives the existing phenotypic variability has been a major topic of interest for biologists for generations. However, the study of the phenotype may not be straightforward. Indeed, organisms may be interpreted as composite objects, comprising different ecophenotypic traits, which are neither necessarily independent from each other nor do they respond to the same evolutionary pressures. For this reason, a deep biological understanding of the focal organism is essential for any morphological analysis. The spider genus Dysdera provides a particularly well-suited system for setting up protocols for morphological analyses that encompass a suit of morphological structures in any nonmodel system. This genus has undergone a remarkable diversification in the Canary Islands, where different species perform different ecological roles, exhibiting different levels of trophic specialization or troglomorphic adaptations, which translate into a remarkable interspecific morphological variability. Here, we seek to develop a broad guide, of which morphological characters must be considered, to study the effect of different ecological pressures in spiders and propose a general workflow that will be useful whenever researchers set out to investigate variation in the body plans of different organisms, with data sets comprising a set of morphological traits. We use geometric morphometric methods to quantify variation in different body structures, all of them with diverse phenotypic modifications in their chelicera, prosoma, and legs. We explore the effect of analyzing different combined landmark (LM) configurations of these characters and the degree of morphological integration that they exhibit. Our results suggest that different LM configurations of each of these body parts exhibit a higher degree of integration compared to LM configurations from different structures and that the analysis of each of these body parts captures different aspects of morphological variation, potentially related to different ecological factors.

Niche expansion and adaptive divergence in the global radiation of crows and ravens.

The processes that allow some lineages to diversify rapidly at a global scale remain poorly understood. Although earlier studies emphasized the importance of dispersal, global expansions expose populations to novel environments and may also require adaptation and diversification across new niches. In this study, we investigated the contributions of these processes to the global radiation of crows and ravens (genus Corvus). Combining a new phylogeny with comprehensive phenotypic and climatic data, we show that Corvus experienced a massive expansion of the climatic niche that was coupled with a substantial increase in the rates of species and phenotypic diversification. The initiation of these processes coincided with the evolution of traits that promoted dispersal and niche expansion. Our findings suggest that rapid global radiations may be better understood as processes in which high dispersal abilities synergise with traits that, like cognition, facilitate persistence in new environments.

Reevaluating scorpion ecomorphs using a naïve approach.

BACKGROUND: Ecomorphs create the opportunity to investigate ecological adaptation because they encompass organisms that evolved characteristic morphologies under similar ecological demands. For over 50 years, scorpions have been empirically assigned to ecomorphs based on the characteristic morphologies that rock, sand, vegetation, underground, and surface dwellers assume. This study aims to independently test the existence of scorpion ecomorphs by quantifying the association between their morphology and ecology across 61 species, representing 14 families of the Scorpiones order. RESULTS: Without a priori categorization of species into ecomorphs, we identified four groups based on microhabitat descriptors, which reflect how scorpion ecospace is clustered. Moreover, these microhabitat groups, i.e., ecotypes, have significantly divergent morphologies; therefore, they represent ecomorphs. These ecomorphs largely correspond with the ones previously described in the literature. Therefore, we retained the names Lithophilous, Psammophilous, and Pelophilous, and proposed the name Phytophilous for vegetation dwellers. Finally, we sought to map the morphology-ecology association in scorpions and found that the morphological regions most tightly associated with ecology are at the extremities. Moreover, the major trend in ecomorphological covariation is that longer walking legs and relatively slender pedipalps (pincers) are associated with sandy microhabitats, while the inverse morphological proportions are associated with rocky microhabitats. CONCLUSIONS: Scorpion ecomorphs are validated in a naïve approach, from ecological descriptors and whole body anatomy. This places them on a more solid quantitative footing for future studies of ecological adaptation in scorpions. Our results verify most of the previously defined ecomorphotypes and could be used as a current practice to understand the adaptive significance of ecological morphology.

Extensive introgression and mosaic genomes of Mediterranean endemic lizards.

The Mediterranean basin is a hotspot of biodiversity, fuelled by climatic oscillation and geological change over the past 20 million years. Wall lizards of the genus Podarcis are among the most abundant, diverse, and conspicuous Mediterranean fauna. Here, we unravel the remarkably entangled evolutionary history of wall lizards by sequencing genomes of 34 major lineages covering 26 species. We demonstrate an early (>11 MYA) separation into two clades centred on the Iberian and Balkan Peninsulas, and two clades of Mediterranean island endemics. Diversification within these clades was pronounced between 6.5-4.0 MYA, a period spanning the Messinian Salinity Crisis, during which the Mediterranean Sea nearly dried up before rapidly refilling. However, genetic exchange between lineages has been a pervasive feature throughout the entire history of wall lizards. This has resulted in a highly reticulated pattern of evolution across the group, characterised by mosaic genomes with major contributions from two or more parental taxa. These hybrid lineages gave rise to several of the extant species that are endemic to Mediterranean islands. The mosaic genomes of island endemics may have promoted their extraordinary adaptability and striking diversity in body size, shape and colouration, which have puzzled biologists for centuries.

Fluctuating asymmetry as biomarker of pesticides exposure in the Italian wall lizards (Podarcis siculus).

The extensive use of pesticides in agricultural environments produces drastic effects on wildlife, hence the need for less invasive indicators of environmental stress to monitoring the impact of agriculture treatments on biological systems. Fluctuating asymmetry (FA), as measure of developmental instability, has recently been proposed as reliable biomarker of populations stress due to environmental disturbance. We investigate femoral pores (FP) and dorsal head shape (HS) traits in populations of the Italian wall lizard inhabiting agricultural environments to examine whether different pesticide exposures (conventional, organic and control) can cause distinctive degree of FA. High-resolution photographs of FP and HS were taken in the field with a digital camera. The number of FP were counted twice on both sides and HS was analysed using geometric morphometrics with 25 landmarks and 12 semilandmarks. Individuals under conventional management showed higher levels of FA compared to control ones, and females exhibited higher FA levels than males for the FP. However, no significant difference was found for the HS trait. Our study provided evidence that FA may have a real potential as biomarker of population stress in wall lizards, highlighting the importance in the choice of the experimental design and the traits adopted for estimating DI.

Variable levels of introgression between the endangered Podarcis carbonelli and highly divergent congeneric species.

Recent empirical studies have demonstrated that speciation with gene flow is more common than previously thought. From a conservation perspective, the potential negative effects of hybridization raise concerns on the genetic integrity of endangered species. However, introgressive hybridization has also been growingly recognized as a source of diversity and new advantageous alleles. Carbonell's wall lizard (Podarcis carbonelli) is an endangered species whose distribution overlaps with four other congeneric species. Our goal here was to determine whether P. carbonelli is completely reproductively isolated from its congeners and to evaluate the relevance of hybridization and interspecific gene flow for developing a conservation plan. We used restriction site associated DNA (RAD) sequencing to discover SNPs in samples from four contact zones between P. carbonelli and four other species. Principal component analysis, multilocus genotype assignment and interspecific heterozygosity suggest incomplete reproductive isolation and ongoing gene flow between species. However, hybridization dynamics vary across all pairs, suggesting complex interactions between multiple intrinsic and extrinsic barriers. Despite seemingly ubiquitous interspecific gene flow, we found evidence of strong reproductive isolation across most contact zones. Instead, indirect effects of hybridization like waste of reproductive effort in small isolated populations may be more problematic. Our results highlight the need to further evaluate the consequences of introgression for P. carbonelli, both on a geographic and genomic level and included in a comprehensive and urgently needed conservation plan. Besides, those findings will add important insights on the potential effects of hybridization and introgression for endangered species.

Genetic and morphological divergence between Littorina fabalis ecotypes in Northern Europe.

Low dispersal marine intertidal species facing strong divergent selective pressures associated with steep environmental gradients have a great potential to inform us about local adaptation and reproductive isolation. Among these, gastropods of the genus Littorina offer a unique system to study parallel phenotypic divergence resulting from adaptation to different habitats related with wave exposure. In this study, we focused on two Littorina fabalis ecotypes from Northern European shores and compared patterns of habitat-related phenotypic and genetic divergence across three different geographic levels (local, regional and global). Geometric morphometric analyses revealed that individuals from habitats moderately exposed to waves usually present a larger shell size with a wider aperture than those from sheltered habitats. The phenotypic clustering of L. fabalis by habitat across most locations (mainly in terms of shell size) support an important role of ecology in morphological divergence. A genome scan based on amplified fragment length polymorphisms (AFLPs) revealed a heterogeneous pattern of differentiation across the genome between populations from the two different habitats, suggesting ecotype divergence in the presence of gene flow. The contrasting patterns of genetic structure between nonoutlier and outlier loci, and the decreased sharing of outlier loci with geographic distance among locations are compatible with parallel evolution of phenotypic divergence, with an important contribution of gene flow and/or ancestral variation. In the future, model-based inference studies based on sequence data across the entire genome will help unravelling these evolutionary hypotheses, improving our knowledge about adaptation and its influence on diversification within the marine realm.

Thermal melanism explains macroevolutionary variation of dorsal pigmentation in Eurasian vipers.

Colouration may endorse thermoregulatory and antipredatory functions in snakes. The thermal melanism hypothesis predicts that dark-coloured individuals are ecologically favoured in cool climates. However, the loss of aposematic and cryptic colourations may imply high predation for melanistic snakes. Here, we used the monophyletic group of Eurasian vipers (subfamily Viperinae) to test whether an increase in the extent of dark area inside the characteristic zigzag dorsal pattern is associated to colder environments. We measured two colouration traits in zigzag-patterned individuals (number of dorsal marks and weighted pigmentation index) and used a phylogenetic comparative approach to explore macroevolutionary patterns of dorsal pigmentation and test whether its extent is associated to ecogeographic characteristics of lineages' ranges. Phylogenetically-naïve and phylogenetically-informed analyses yielded a significant association between the degree of pigmentation of the zigzag pattern and environmental variables such as solar radiation, elevation and latitude. The degree of pigmentation of the zigzag pattern is highlighted as an adaptive trait that matches range attributes mirroring cold environments irrespective of the phylogeny. These results constitute the first large-scale evidence supporting the thermal melanism hypothesis in snakes, opening new avenues of inquiry for the mechanisms that shape the evolution of colour phenotypes.

Interspecific allometry for sexual shape dimorphism: Macroevolution of multivariate sexual phenotypes with application to Rensch's rule.

Allometric trends in the degree of sexual dimorphism with body size have long fascinated evolutionary biologists. Many male-biased clades display more prominent sexual dimorphism in larger taxa (Rensch's rule), with most examples documenting this pattern for body size dimorphism. Although sexual dimorphism in traits other than body size is equally functionally relevant, characterizing allometric patterns of sexual dimorphism in such traits is hampered by lack of an analytical framework that can accommodate multivariate phenotypes. In this article, we derive a multivariate equivalency for investigating trends in sexual dimorphism-relative to overall body size-across taxa and provide a generalized test to determine whether such allometric patterns correspond with Rensch's rule. For univariate linear traits such as body size, our approach yields equivalent results to those from standard procedures, but our test is also capable of detecting trends in multivariate datasets such as shape. Computer simulations reveal that the method displays appropriate statistical properties, and an empirical example in Mediterranean lizards provides the first demonstration of Rensch's rule in a multivariate phenotype (head shape). Our generalized procedure substantially extends the analytical toolkit for investigating macroevolutionary patterns of sexual dimorphism and seeking a better understanding of the processes that underlie them.

New Insights into Bite Performance: Morphological Trade-Offs Underlying the Duration and Magnitude of Bite Force.

Biting performance is important for feeding, territory defense, and mating in many animals. While maximal bite force is a well-studied trait, other aspects of biting and their variation depending on behavior are rarely considered. Here, we took an innovative approach, where (1) we quantified a novel trait, bite duration in lizards; (2) we examined variation across setups that simulate feeding and antagonistic behavior; and (3) we used F-matrix statistics to investigate how different functional components are optimized for ecological and social demands with respect to individual morphology. Our results did not show differences between the sexes in bite duration, but bite performance varied across experimental setups in males, suggesting a higher functional flexibility in this sex. The investigation of form-function associations revealed that trade-offs, facilitations, and one-to-one relationships are simultaneously involved in the morphological optimization of bite force and duration. Put together, our integrated analysis of two different components of bite performance-force and duration-demonstrates their importance for males in both ecological and social tasks. Our findings also suggest the existence of trade-offs in the morphological optimization of functional components, possibly due to physiological constraints on muscle composition, insertion, and orientation.

Hybridization patterns between two marine snails, Littorina fabalis and L. obtusata.

Characterizing the patterns of hybridization between closely related species is crucial to understand the role of gene flow in speciation. In particular, systems comprising multiple contacts between sister species offer an outstanding opportunity to investigate how reproductive isolation varies with environmental conditions, demography and geographic contexts of divergence. The flat periwinkles, Littorina obtusata and L. fabalis (Gastropoda), are two intertidal sister species with marked ecological differences compatible with late stages of speciation. Although hybridization between the two was previously suggested, its extent across the Atlantic shores of Europe remained largely unknown. Here, we combined genetic (microsatellites and mtDNA) and morphological data (shell and male genital morphology) from multiple populations of flat periwinkles in north-western Iberia to assess the extent of current and past hybridization between L. obtusata and L. fabalis under two contrasting geographic settings of divergence (sympatry and allopatry). Hybridization signatures based on both mtDNA and microsatellites were stronger in sympatric sites, although evidence for recent extensive admixture was found in a single location. Misidentification of individuals into species based on shell morphology was higher in sympatric than in allopatric sites. However, despite hybridization, species distinctiveness based on this phenotypic trait together with male genital morphology remained relatively high. The observed variation in the extent of hybridization among locations provides a rare opportunity for future studies on the consequences of different levels of gene flow for reinforcement, thus informing about the mechanisms underlying the completion of speciation.

Where does diversity come from? Linking geographical patterns of morphological, genetic, and environmental variation in wall lizards.

BACKGROUND: Understanding how phenotypic variation scales from individuals, through populations, up to species, and how it relates to genetic and environmental factors, is essential for deciphering the evolutionary mechanisms that drive biodiversity. We used two species of Podarcis wall lizards to test whether phenotypic diversity within and divergence across populations follow concordant patterns, and to examine how phenotypic variation responds to genetic and environmental variability across different hierarchical levels of biological organization, in an explicit geographic framework. RESULTS: We found a general concordance of phenotypic variation across hierarchical levels (i.e. individuals and populations). However, we also found that within-population diversity does not exhibit a coherent geographic structure for most traits, while among-population divergence does, suggesting that different mechanisms may underlie the generation of diversity at these two levels. Furthermore, the association of phenotypic variation with genetic and environmental factors varied extensively between hierarchical levels and across traits, hampering the identification of simple rules to explain what yields diversity. CONCLUSIONS: Our results in some cases comply with general ecological and evolutionary predictions, but in others they are difficult to explain in the geographic framework used, suggesting that habitat characteristics and other regulatory mechanisms may have a more substantial contribution in shaping phenotypic diversity.

Run for your life, but bite for your rights? How interactions between natural and sexual selection shape functional morphology across habitats.

A central issue in evolutionary biology is how morphology, performance, and habitat use coevolve. If morphological variation is tightly associated with habitat use, then differences in morphology should affect fitness through their effect on performance within specific habitats. In this study, we investigate how evolutionary forces mold morphological traits and performance differently given the surrounding environment, at the intraspecific level. For this purpose, we selected populations of the lizard Podarcis bocagei from two different habitat types, agricultural walls and dunes, which we expected to reflect saxicolous vs ground-dwelling habits. In the laboratory, we recorded morphological traits as well as performance traits by measuring sprint speed, climbing capacity, maneuverability, and bite force. Our results revealed fast-evolving ecomorphological variation among populations of P. bocagei, where a direct association existed between head morphology and bite performance. However, we could not establish links between limb morphology and locomotor performance at the individual level. Lizards from walls were better climbers than those from dunes, suggesting a very fast evolutionary response. Interestingly, a significant interaction between habitat and sex was detected in climbing performance. In addition, lizards from dunes bit harder than those from walls, although sexual differentiation was definitely the main factor driving variation in head functional morphology. Taking into account all the results, we found a complex interaction between natural and sexual selection on whole-organism performance, which are, in some cases, reflected in morphological variation.

The ontogeny of developmental buffering in lizard head shape.

Canalization and developmental stability (DS) are important organismal properties involved in determining the level of phenotypic variation. Ontogenetic patterns of phenotypic variance components can shed light on the mechanistic basis of developmental buffering (DB). Here, we analyze how individual FA and among-individual variation in head shape change in ontogenetic series of three lizard species raised in laboratory. The degree of asymmetry increased slightly with size, suggesting that developmental mechanisms hypothesized to correct for deviations either do not exist, or that their efficiency is truncated with increasing size. Alternatively, they may need the disturbance as a trigger. The relationship between asymmetry and age was complex, with asymmetry being stable across the age range in two species but increased with age in the third. Lack of congruence in ontogenetic patterns of asymmetry might be due to intrinsic differences in buffering mechanisms or a result of species-specific growth patterns. Head shape was shown to be equally canalized across both size and age range in all species, probably as a result of a balance between the buffering mechanisms and mechanisms generating variance. The patterns of symmetric and asymmetric head shape variation were highly correlated across species meaning that DS and canalization may rely on similar mechanisms.

A non-invasive geometric morphometrics method for exploring variation in dorsal head shape in urodeles: sexual dimorphism and geographic variation in Salamandra salamandra.

The study of morphological variation among and within taxa can shed light on the evolution of phenotypic diversification. In the case of urodeles, the dorso-ventral view of the head captures most of the ontogenetic and evolutionary variation of the entire head, which is a structure with a high potential for being a target of selection due to its relevance in ecological and social functions. Here, we describe a non-invasive procedure of geometric morphometrics for exploring morphological variation in the external dorso-ventral view of urodeles' head. To explore the accuracy of the method and its potential for describing morphological patterns we applied it to two populations of Salamandra salamandra gallaica from NW Iberia. Using landmark-based geometric morphometrics, we detected differences in head shape between populations and sexes, and an allometric relationship between shape and size. We also determined that not all differences in head shape are due to size variation, suggesting intrinsic shape differences across sexes and populations. These morphological patterns had not been previously explored in S. salamandra, despite the high levels of intraspecific diversity within this species. The methodological procedure presented here allows to detect shape variation at a very fine scale, and solves the drawbacks of using cranial samples, thus increasing the possibilities of using collection specimens and alive animals for exploring dorsal head shape variation and its evolutionary and ecological implications in urodeles. J. Morphol. 278:475-485, 2017. © 2017 Wiley Periodicals, Inc.

Phylogenies, the Comparative Method, and the Conflation of Tempo and Mode.

The comparison of mathematical models that represent alternative hypotheses about the tempo and mode of evolutionary change is a common approach for assessing the evolutionary processes underlying phenotypic diversification. However, because model parameters are estimated simultaneously, they are inextricably linked, such that changes in tempo, the pace of evolution, and mode, the manner in which evolution occurs, may be difficult to assess separately. This may potentially complicate biological interpretation, but the extent to which this occurs has not yet been determined. In this study, we examined 160 phylogeny × trait empirical data sets, and conducted extensive numerical phylogenetic simulations, to investigate the efficacy of phylogenetic comparative methods to distinguish between models that represent different evolutionary processes in a phylogenetic context. We observed that, in some circumstances, a high uncertainty exists when attempting to distinguish between alternative evolutionary scenarios underlying phenotypic variation. When examining data sets simulated under known conditions, we found that evolutionary inference is straightforward when phenotypic patterns are generated by simple evolutionary processes that are represented by modifying a single model parameter at a time. However, inferring the exact nature of the evolutionary process that has yielded phenotypic variation when facing complex, potentially more realistic, mechanisms is more problematic. A detailed investigation of the influence of different model parameters showed that changes in evolutionary rates, marked changes in phylogenetic means, or the existence of a strong selective pull on the data, are all readily recovered by phenotypic model comparison. However, under evolutionary processes with a milder restraining pull acting on trait values, alternative models representing very different evolutionary processes may exhibit similar goodness-of-fit to the data, potentially leading to the conflation of interpretations that emphasize tempo and mode during empirical evolutionary inference. This is a mathematical and conceptual property of the considered models that, while not prohibitive for studying phenotypic evolution, should be taken into account and addressed when appropriate.

A geometric morphometric analysis of acetabular shape of the primate hip joint in relation to locomotor behaviour.

The description of acetabular shape variation among primates is essential for our understanding of the locomotor behaviour and ecology of both extant and fossil species. In this study, we use two-dimensional geometric morphometrics to examine variation in acetabular shape in human and non-human primates and to determine the degree to which it co-varies with locomotor behaviour, while taking both intra and inter-specific variation into account. To these ends, we examined the acetabulum of 303 left hip bones of 27 extant genera of primates (including humans) with different locomotor behaviours. After accounting for shape variation due to sex, size, and phylogeny, the results confirm that acetabular shape varies significantly across locomotor groups. The two most differentiated locomotor groups are leapers and slow-climbing quadrupeds, which exhibit a unique acetabular shape. Furthermore, the acetabulum of humans differed significantly from all other groups, while no significant differences existed between chimpanzees and gorillas. The most noticeable differences are detected in both cranial and dorsal areas and around the acetabular horns. This variation in acetabular morphology may have biomechanical implications at the level of the hip joint, potentially determining joint range motion and load distribution during locomotion. Given the increasing number of published studies on fossil pelves, our results are widely applicable to fossil analyses, with critical implications for paleoanthropological analyses about the complex locomotor behaviour of fossil specimens and their classification into locomotor groups, which may enhance our understanding of their ecological habits.