Skull Variation Research Articles

Morphological patterns of the European bison (Bison bonasus) skull

This study aimed to investigate the effects of environmental factors, sexual selection, and genetic variation on skull morphology by examining the skull structure of the European bison, a species at risk of extinction, and comparing it to other bovid species. The skull of the European bison was significantly bigger than that of other species of the tribe Bovini, and the results revealed considerable morphological differences in skull shape compared to other Bovini samples. The bison skull exhibited a broader shape in the frontal region and a more laterally oriented cornual process. The frontal bone also significantly influenced skull shape variation within the European bison subspecies. Also, the findings indicated that cornual processes size significantly affected skull shape. Male Bison had larger skulls and more developed cornual ridges, and these morphological changes in the frontal region are thought to have also influenced the nuchal region, maxillary region, and facial bones. Furthermore, the differences in skull size observed between the Lowland-Pszczyna and Lowland-Białowieża lines, which share a closer origin than the Lowland-Carpathian lines, may reflect environmental and genetic adaptations over time. This research provides a reference point for future studies on the ecological and evolutionary factors influencing bison skull variations, with significant implications for the conservation of this species.

Skull evolution and lineage diversification in endemic Malagasy carnivorans.

Madagascar is one of the world's foremost biodiversity hotspots with more than 90% of its species endemic to the island. Malagasy carnivorans are one of only four extant terrestrial mammalian clades endemic to Madagascar. Although there are only eight extant species, these carnivorans exhibit remarkable phenotypic and ecological diversity that is often hypothesized to have diversified through an adaptive radiation. Here, we investigated the evolution of skull diversity in Malagasy carnivorans and tested if they exhibited characteristics of convergence and an adaptive radiation. We found that their skull disparity exceeds that of any other feliform family, as their skulls vary widely and strikingly capture a large amount of the morphological variation found across all feliforms. We also found evidence of shared adaptive zones in cranial shape between euplerid subclades and felids, herpestids and viverrids. Lastly, contrary to predictions of adaptive radiation, we found that Malagasy carnivorans do not exhibit rapid lineage diversification and only marginally faster rates of mandibular shape evolution and to a lesser extent cranial shape evolution, compared to other feliforms. These results reveal that exceptional diversification rates are not necessary to generate the striking phenotypic diversity that evolved in carnivorans after their dispersal to and isolation on Madagascar.

Comparative morphology in the context of facial reduction: Modularity in primate, dog, and bat crania.

Biological variation in the mammalian skull is the product of a series of factors including changes in gene expression, developmental timing, and environmental pressures. When considering the diversity of extant mammalian crania, it is important to understand these mechanisms that contribute to cranial growth and in turn, how differences in cranial morphology have been attained. Various researchers, including Dr. Sue Herring, have proposed a variety of mechanisms to explain the process of cranial growth. This work has set the foundation on which modern analysis of craniofacial morphology happens today. This study focused on the analysis of modularity in three mammalian taxa, all of which exhibit facial reduction. Specifically, we examined facial reduction as a morphological phenomenon through the use of two-module and six-module modularity hypotheses. We recorded three-dimensional coordinate data for 55 cranial landmarks that allowed us to analyze differences in cranial shape in these three taxa (primates n = 88, bats n = 64, dogs n = 81). When assessing modularity within the two-module modularity hypothesis specifically, dogs exhibited the lowest levels of modularity, while bats and primates both showed a slightly more modular covariance structure. We further assessed modularity in the same sample using the Goswami six-module model, where again dogs exhibited a low degree of modularity, with bats and primates being more moderate. We then broke the sample into subsets by analyzing each morphotype separately. We hypothesized that the modularity would be more pronounced in the brachycephalic morphotype. Surprisingly, we found that in brachycephalic dogs, normocephalic dogs, brachycephalic primates, and normocephalic primates, there was a moderate degree of modularity. Brachycephalic bats had a low degree of modularity, while normocephalic bats were the most modular group observed in this study. Based on these results, it is evident that facial reduction is a complex and multifaceted phenomenon with unique morphological changes observed in each of the three taxa studied.

Morphological and molecular data combined reveal inter- and intraspecific cranial shape variations in bats of Artibeus Leach, 1821 (Chiroptera: Phyllostomidae)

Abstract Since morphological traits may overlap among species within a complex, the integration of morphological and mitochondrial data could provide crucial insights for distinguishing species, as observed in fruit-eating bats of the genus Artibeus. Therefore, the application of geometric morphometric (GM) techniques could yield more refined and robust analyses of inter- and intraspecific variations. In this study, we generated two datasets: one with molecular delimitation data based on the barcode region (COI) and the other with cranial size and shape data using GM methods. Our aims were to investigate variations between large and small species within the genus Artibeus, as well as to explore potential factors influencing such variations. The results from species delimitation revealed molecular operational taxonomic units (MOTUs) supporting the morphological identification. Analyses using GM techniques demonstrated significant inter- and intraspecific variations in cranial size and shape among Artibeus species. The combined outcomes suggest the absence of a significant phylogenetic signal influencing skull variation. This scenario indicates that potential historical ecological factors may have directly influenced the cranial morphology of these species, acting as significant selection forces in geographical space and generating intraspecific variations in Artibeus planirostris.

Determination of skull variation in Akkaraman and Kangal Akkaraman sheep by geometric morphometric method

Sheep breeding has an important place in the world and in Turkey, as sheep are farm animals that provide most of their nutritional needs from pasture, have good adaptability and are low in cost. Akkaraman sheep and the Kangal Akkaraman sheep, which was considered its sub-variety until 2012, constitute half of the sheep population in Turkey. In this study, morphometric diversity between the skulls of Akkaraman and Kangal Akkaraman breed sheep was examined by geometric morphometric method. 2D images obtained from 3D models of 24 male sheep skulls from two different provinces were analyzed using 10 landmarks from dorsal and lateral views. The results from the dorsal view, the first three principal components (PC1, PC2 and PC3) accounted for a total of 64.13% of the morphological variation and the largest changes in are associated with external occipital protuberance, midline nasofrontal suture and rostral edge of the incisive fissure. From the lateral view, three first principal components explained the morphological variation 27.56%, 22.32% and 10.65% respectively. In the comparison of the skull morphology of Akkaraman and Kangal Akkaraman sheep from dorsal and lateral views, no allometric components were identified, as most of the described shape variations were not size-dependent. As a result of these analyses, more detailed information was obtained about the shape differences of the skull. Geometric morphometric analysis is successful in detecting differences in skull shape of different sheep breeds and can be successfully used for classification.

Dynamic evolutionary interplay between ontogenetic skull patterning and whole-head integration.

The arrangement and morphology of the vertebrate skull reflect functional and ecological demands, making it a highly adaptable structure. However, the fundamental developmental and macroevolutionary mechanisms leading to different vertebrate skull phenotypes remain unclear. Here we exploit the morphological diversity of squamate reptiles to assess the developmental and evolutionary patterns of skull variation and covariation in the whole head. Our geometric morphometric analysis of a complex squamate ontogenetic dataset (209 specimens, 169 embryos, 44 species), covering stages from craniofacial primordia to fully ossified bones, reveals that morphological differences between snake and lizard skulls arose gradually through changes in spatial relationships (heterotopy) followed by alterations in developmental timing or rate (heterochrony). Along with dynamic spatiotemporal changes in the integration pattern of skull bone shape and topology with surrounding brain tissues and sensory organs, we identify a relatively higher phenotypic integration of the developing snake head compared with lizards. The eye, nasal cavity and Jacobson's organ are pivotal in skull morphogenesis, highlighting the importance of sensory rearrangements in snake evolution. Furthermore, our findings demonstrate the importance of early embryonic, ontogenetic and tissue interactions in shaping craniofacial evolution and ecological diversification in squamates, with implications for the nature of cranio-cerebral relations across vertebrates.

Ontogenetic skull variation in a shovel-headed amphisbaenian species.

Leposternon microcephalum is a species belonging to the Amphisbaenia, a group of burrowing reptiles. Amphisbaenia present various morphological and physiological adaptations that allow them to penetrate the ground and live underground, through a system of galleries and permanent chambers that they build themselves. Among the morphological adaptations in this group, those of the skull stand out as it serves as the main excavation tool. Four basic skull shapes are recognized: rounded, keeled, shovel-shaped, and spade-shaped. The skull of L. microcephalum belongs to this last type, which is considered the most specialized. The species inhabits soils that are highly compacted and difficult to penetrate. Among the species of Leposternon present in South America, L. microcephalum has the widest distribution, being found in all Brazilian biomes and neighboring countries such as Bolivia, Argentina, Paraguay, and Uruguay. The analysis of the skull of this species was carried out using three-dimensional geometric morphometrics (3D-GMM), a technique that allows comparative analysis, through robust statistical methods, of shape and its variations, using Cartesian coordinate data from a configuration of homologous landmarks. The technique allows the size and shape components of a structure to be analyzed separately. From an ontogenetic point of view, this methodology had also been used to investigate variations in Cynisca leucura, a member of the Amphisbaenidae with a rounded head. Our hypothesis is that the patterns of morphological differentiation in the skull, mainly in the intermediate and occipital regions, are similar in different Amphisbaenia species. Therefore, the objective of this study was to analyze cranial morphological variations in an ontogenetic series of L. microcephalum using 3D-GMM. Computed Tomographic scans of 13 specimens were analyzed: juveniles (N = 8) and adults (N = 5), based on 20 landmarks that characterize the skull. Principal components and regression analyses between shape (dependent variable) and size (independent variable) showed a clear difference between the cranial morphological pattern of juvenile individuals and that of adults. For instance, young specimens tend to have a dorsoventrally tall neurocranium, with the tip of the snout more anteriorly oriented and its dorsal border subtly curved. Dorsally, the parietal region is thicker and smoothly dome-shaped in juveniles. As in C. leucura, the variation was strongly correlated with the size change from juvenile to adult, indicating a dominant role for ontogenetic allometry in determining skull shape.

Digest: Ecological and life history drivers of avian skull evolution.

What are the factors that drive the patterns and evolutionary rates of morphological characteristics? To answer this question, Hunt et al. (2023) explore shape variation and rates of change in modern avian skulls using morphometric measurements and phylogenetic analyses. They find that habitat density and migration have the strongest influences on avian skull variation, and that denser habitats, longer migratory distances, and precocial developmental modes are all associated with faster rates of morphological evolution.

Genomic diversity, evolutionary history, and species limits of the endemic Ethiopian laminate-toothed rats (genus Otomys, Rodentia: Muridae)

Abstract The Ethiopian Highlands represent one of the most important centres of endemism on Earth. Six endemic species of laminate-toothed rats have been reported from Ethiopia in a previous study based mostly on morphological analyses. Largely missing genetic data for some of the species, insufficient sampling across Ethiopia, and presence of a pseudogene on cytochrome b created knowledge gaps in basic taxonomy, distribution, and phylogenetic position of the Ethiopian taxa. Here we perform an integrative taxonomic revision of the group by using the largest available sets of genomic and morphological data. We first reconstructed the mitochondrial phylogeny of the whole genus, delimited major clades, and evidenced two independent colonization events of Otomys to Ethiopia. By using genome-scale SNPs, we delimited putative Ethiopian species and analysed their phylogenetic relationships. In genetically characterized specimens we assessed morphological variation of skulls. We confirmed the presence of six previously reported species and significantly extended the known distributional range for some of them. Furthermore, we discovered a genetically and morphologically distinct lineage in northern Ethiopia, probably representing a new species. The speciation processes are similar to other Afromontane Ethiopian endemic rodent clades, with prevailing allopatric diversification combined with reticulate processes.

Comparative digital reconstruction of Pica pica and Struthio camelus and their cranial suture ontogenies.

To date, several studies describe post-hatching ontogenetic variation in birds; however, none of these studies document and compare ontogenetic variation of the entire skull in multiple avian species. Therefore, we studied ontogenetic skull variation of two bird species with very different ecologies, Pica pica, and Struthio camelus, using μCT based 3D reconstructions. For each specimen, we performed bone-by-bone segmentation in order to visualize and describe the morphological variation of each bone during ontogeny and estimated the average sutural closure of the skulls to identify different ontogenetic stages. Although bone fusion of P. pica occurs more rapidly than that of S. camelus the general sequence of bone fusion follows a similar trend from posterior to anterior, but a more detailed analysis reveals some interspecific variation in the fusion patterns. Although growth persists over a longer period in S. camelus than in P. pica and adults of the former species are significantly larger, the skull of the most mature S. camelus is still less fused than that of P. pica. Different growth and fusion patterns of the two species indicate that the interspecific ontogenetic variation could be related to heterochronic developments. Nevertheless, this hypothesis needs to be tested in a broader phylogenetic framework in order to detect the evolutionary direction of the potential heterochronic transformations.

The skull variation of the olive field mouse Abrothrix olivacea (Cricetidae: Abrotrichini) is localized and correlated to the ecogeographic features of its geographic distribution.

The relationship between phenotypic variation and landscape heterogeneity has been extensively studied to understand how the environment influences patterns of morphological variation and differentiation of populations. Several studies had partially addressed intraspecific variation in the sigmodontine rodent Abrothrix olivacea, focusing on the characterization of physiological aspects and cranial variation. However, these had been conducted based on geographically restricted populational samples, and in most cases, the aspects characterized were not explicitly contextualized with the environmental configurations in which the populations occurred. Here, the cranial variation of A. olivacea was characterized by recording twenty cranial measurements in 235 individuals from 64 localities in Argentina and Chile, which widely cover the geographic and environmental distribution of this species. The morphological variation was analyzed and ecogeographically contextualized using multivariate statistical analyses, which also included climatic and ecological variation at the localities where the individuals were sampled. Results indicate that the cranial variation of this species is mostly clustered in localized patterns associated to the types of environments, and that the levels of cranial differentiation are higher among the populations from arid and treeless zones. Additionally, the ecogeographical association of cranial size variation indicate that this species does not follow Bergmann's rule and that island populations exhibit larger cranial sizes compared to their continental counterparts distributed at the same latitudes. These results suggest that cranial differentiation among the populations of this species is not homogeneous throughout its geographic distribution, and that the patterns of morphological differentiation are also not completely consistent with the patterns of genetic structuring that have been described recently. Finally, the analyses performed to ponder morphological differentiation among populations suggest that the contribution of genetic drift in the formation of these patterns can be ruled out among Patagonian populations, and that the selective effect imposed by the environment could better explain them.

Skull variation in different breeds sheep from Balkan countries

The Balkan Peninsula region has a very diverse agricultural and livestock tradition, and almost every country has its own local breed of sheep. Different breeds of sheep and different breeding traditions, despite the small geographical distance, determine the morphological and morphometric variability among animal breeds. In this study, this morphological diversity among the skulls of sheep breeds of some countries in the Balkan region was examined by the geometric morphometric method. 2D images of 86 sheep skulls from five different countries were analyzed from the dorsal view.Sixteen landmarks were used. The Bardhoka and the Ivesi breed have the broadest distributions of skull shape amongst the sheep breeds. The Ruda sheep is the most morphologically conservative. The sheep from Turkey (Ivesi) and Kosovo (Bardhoka) seem to differ mainly from sheep from other Balkan countries. Bardhoka and Ruda differ most from each other (p < 0.0001). The next biggest differences were between Ivesi and Ruda (p < 0.0011) and between Bardhoka and Sharri sheep (p < 0.0016). The sheep breeds Dubska and Lara e Polisit differ the least from each other. Geometrics morphometric analysis is a useful tool to detect differences in the shape of the skull of different sheep breeds and can therefore be used successfully for taxonomic purposes.

Geographic Variation in Skull Morphology of the Japanese Crocodile Newt, Echinotriton andersoni (Amphibia: Urodela: Salamandridae), Inferred from Geometric Morphometrics

The Anderson's crocodile newt, Echinotriton andersoni, is an endangered species distributed in the islands of Central Ryukyus, Japan. This species is well known to show high intraspecific genetic variation among islands, but its morphological variation has not been explored. We examined the skull variation in this species including enigmatic specimens collected from Taiwan by using geometric morphometrics, and found sexual differences and geographic variation mainly between the Okinawa group (Okinawajima Island) and the Amami Group (Amamioshima and Tokunoshima Islands). We further found close resemblance between the Taiwanese specimens and the female specimens from Amamioshima Island.

Skull variation in populations of the Indian gerbil Tatera indica (Gerbillinae, Rodentia) sampled across its broad geographic range

Populations of broadly distributed species often exhibit geographic structuring, which is sometimes reflected in phenotype. The monotypic Indian gerbil (Tatera indica) is an example of a widely distributed species, with its range encompassing much of Asia. This study aims to determine if T. indica populations exhibit marked variation in skull morphology—this structure is particularly adaptable and thus could be amenable to show such variation. Furthermore, the potential drivers of skull variation are examined, including the role of climate and geography. To achieve these goals, 21 linear measurements were measured on the skulls of 509 specimens, coming from 111 different localities, across this species wide range. The specimens were then assigned into one of four broad geographic groups (≈ populations) based on their geographic proximity, and the overall and the pairwise differences in the 21 skull measurements among these groups were assessed. Specimens from Pakistan significantly differed from those belonging to the West Iran, East Iran, and India populations, which in turn did not significantly differ from each other. Pairwise bioclimatic and geographic distances between the localities explained a significant, yet small amount of variation in the measurements. Thus, while the Pakistani T. indica population was distinct in skull measurements, both climatic and non-climatic spatial factors seem not to account largely for its distinctiveness (from the other populations).

Phenotypic plasticity determines differences between the skulls of tigers from mainland Asia.

Tiger subspecific taxonomy is controversial because of morphological and genetic variation found between now fragmented populations, yet the extent to which phenotypic plasticity or genetic variation affects phenotypes of putative tiger subspecies has not been explicitly addressed. In order to assess the role of phenotypic plasticity in determining skull variation, we compared skull morphology among continental tigers from zoos and the wild. In turn, we examine continental tiger skulls from across their wild range, to evaluate how the different environmental conditions experienced by individuals in the wild can influence morphological variation. Fifty-seven measurements from 172 specimens were used to analyse size and shape differences among wild and captive continental tiger skulls. Captive specimens have broader skulls, and shorter rostral depths and mandible heights than wild specimens. In addition, sagittal crest size is larger in wild Amur tigers compared with those from captivity, and it is larger in wild Amur tigers compared with other wild continental tigers. The degree of phenotypic plasticity shown by the sagittal crest, skull width and rostral height suggests that the distinctive shape of Amur tiger skulls compared with that of other continental tigers is mostly a phenotypically plastic response to differences in their environments.

Intraspecific variation in the cranial osteology of Diplometopon zarudnyi (Squamata: Amphisbaenia: Trogonophidae).

A snake-like body plan and burrowing lifestyle characterize numerous vertebrate groups as a result of convergent evolution. One such group is the amphisbaenians, a clade of limbless, fossorial lizards that exhibit head-first burrowing behavior. Correlated with this behavior, amphisbaenian skulls are more rigid and coossified than those of nonburrowing lizards. However, due to their lifestyle, there are many gaps in our understanding of amphisbaenian anatomy, including how their cranial osteology varies among individuals of the same species and what that reveals about constraints on the skull morphology of head-first burrowing taxa. We investigated intraspecific variation in the cranial osteology of amphisbaenians using seven individuals of the trogonophid Diplometopon zarudnyi. Variation in both skull and individual skull element morphology was examined qualitatively and quantitatively through three-dimensional (3D) models created from microcomputed tomography data. Qualitative examination revealed differences in the number and position of foramina, the interdigitation between the frontals and parietal, and the extent of coossification among the occipital complex, fused basioccipital and parabasisphenoid ("parabasisphenoid-basioccipital complex"), and elements X. We performed 3D landmark-based geometric morphometrics for the quantitative assessment, revealing shape differences in the skull, premaxilla, maxilla, frontal, and parietal. The observed intraspecific variation may be the result of different stages of ontogenetic development or biomechanical optimization for head-first burrowing. For example, variation in the coossification of the occipital region suggests a potential ontogenetic coossification sequence. Examination of these areas of variation across other head-first burrowing taxa will help determine if the variation is clade-specific or part of a broader macroevolutionary pattern of head-first burrowing.

Multi-locus phylogeny and species delimitations of the striped-back shrew group (Eulipotyphla: Soricidae): Implications for cryptic diversity, taxonomy and multiple speciation patterns

The striped-back shrew group demonstrates remarkable variation in skull and body size, tail length, and brightness of the dorsal stripe; and karyotypic and DNA variation has been reported in recent years. In this study, we investigated the phylogenetic structure of the group, as well as speciation patterns and demographic history in Mountains of Southwestern China and adjacent mountains, including the southern Himalayas, Mts. Bashan, Wushan, and Qinling. We sequenced a total of 462 specimens from 126 localities in the known range of the group, which were sequenced and analyzed based on 6.2 kb of sequence data from two mitochondrial, six nuclear, and two Y chromosome markers. Phylogenetic analyses of the concatenated mtDNA data revealed 14 sympatric and independently evolving lineages within the striped-back shrew group, including Sorex bedfordiae, S. cylindricauda, S. excelsus, S. sinalis and several cryptic species. All concatenated data (ten genes) showed a consistent genetic structure compared to the mtDNA lineages for the group, whereas the nuclear and the Y chromosome data showed a discordant genetic structure compared to the mtDNA lineages for the striped-back shrew group. Species delimitation analyses and deep genetic distance clearly support the species status of the 14 evolving lineages. The divergence time estimation suggested that the striped-back shrew group began to diversify from the middle Pleistocene (2.34 Ma), then flourished at approximately 2.14 Ma, followed by a series of rapid diversifications through the Pleistocene. Our results also revealed multiple mechanisms of speciation in the Mountains of Southwestern China and Adjacent Mountains with complex landscapes and climate. The uplifting of the Qinghai-Tibetan Plateau, Quaternary climate oscillations, riverine barriers, ecological elevation gradients, topographical diversity, and their own low dispersal capacity may have driven the speciation, genetic structure, and phylogeographic patterns of the striped-back shrew group.

Skull ecomorphological variation of narwhals (Monodon monoceros, Linnaeus 1758) and belugas (Delphinapterus leucas, Pallas 1776) reveals phenotype of their hybrids.

Narwhals and belugas are toothed whales belonging to the Monodontidae. Belugas have a circumpolar Arctic and sub-Artic distribution while narwhals are restricted to the Atlantic Arctic. Their geographical ranges overlap during winter migrations in the Baffin Bay area (Canada/West Greenland) and successful interbreeding may occur. Here, we employed geometric morphometrics on museum specimens to explore the cranium and mandible morphology of a known hybrid (NHMD MCE 1356) and the cranium morphology of a putative hybrid (NHMD 1963.44.1.4) relative to skull morphological variation in the parental species. Specifically, we used 3D models of skulls from 69 belugas, 86 narwhals, and the two known/putative hybrids and 2D left hemi-mandibles from 20 belugas, 64 narwhals and the known hybrid. Skull shape analyses allowed clear discrimination between species. Narwhals are characterised by a relatively short rostrum and wide neurocranium while belugas show a more elongated and narrower cranium. Sexual size dimorphism was detected in narwhals, with males larger than females, but no sexual shape dimorphism was detected in either species (excluding presence/absence of tusks in narwhals). Morphological skull variation was also dependent on different allometric slopes between species and sexes in narwhals. Our analyses showed that the cranium of the known hybrid was phenotypically close to belugas but its 2D hemi-mandible had a narwhal shape and size morphology. Both cranium and mandible were strongly correlated, with the pattern of covariation being similar to belugas. The putative hybrid was a pure male narwhal with extruded teeth. Comparison of genomic DNA supported this result, and stable carbon and nitrogen isotope values suggested that the putative hybrid had a more benthic foraging strategy compared to narwhals. This work demonstrates that although the known hybrid could be discriminated from narwhals and belugas, detection of its affinities with these parental species was dependent on the part of the skull analysed.

Skull variation in Afro-Eurasian monkeys results from both adaptive and non-adaptive evolutionary processes

Afro-Eurasian monkeys originated in the Miocene and are the most species-rich modern primate family. Molecular and fossil data have provided considerable insight into their evolutionary divergence, but we know considerably less about the evolutionary processes that underlie these differences. Here, we apply tests developed from quantitative genetics theory to a large (n > 3000) cranio-mandibular morphometric dataset, investigating the relative importance of adaptation (natural selection) and neutral processes (genetic drift) in shaping diversity at different taxonomic levels, an approach applied previously to monkeys of the Americas, apes, hominins, and other vertebrate taxa. Results indicate that natural selection, particularly for differences in size, plays a significant role in diversifying Afro-Eurasian monkeys as a whole. However, drift appears to better explain skull divergence within the subfamily Colobinae, and in particular the African colobine clade, likely due to habitat fragmentation. Small and declining population sizes make it likely that drift will continue in this taxon, with potentially dire implications for genetic diversity and future resilience in the face of environmental change. For the other taxa, many of whom also have decreasing populations and are threatened, understanding adaptive pressures similarly helps identify relative vulnerability and may assist with prioritising scarce conservation resources.

Range-wide variation in grey seal (Halichoerus grypus) skull morphology

The large interspecific variation in marine mammal skull and dental morphology reflects ecological specialisations to foraging and communication. At the intraspecific level, the drivers of skull shape variation are less well understood, having implications for identifying putative local foraging adaptations and delineating populations and subspecies for taxonomy, systematics, management and conservation. Here, we assess the range-wide intraspecific variation in 71 grey seal skulls by 3D surface scanning, collection of cranial landmarks and geometric morphometric analysis. We find that skull shape differs slightly between populations in the Northwest Atlantic, Northeast Atlantic and Baltic Sea. However, there was a large shape overlap between populations and variation was substantially larger among animals within populations than between. We hypothesize that this pattern of intraspecific variation in grey seal skull shape results from balancing selection or phenotypic plasticity allowing for a remarkably generalist foraging behaviour. Moreover, the large overlap in skull shape between populations implies that the separate subspecies status of Atlantic and Baltic Sea grey seals is questionable from a morphological point of view.