Evolutionary ecomorphology for the twenty-first century: examples from mammalian carnivores.

Carnivores (cats, dogs and kin) are a diverse group of mammals that inhabit a remarkable range of ecological niches. While the relationship between ecology and morphology has long been of interest in carnivorans, the application of quantitative techniques has resulted in a recent explosion of work in the field. Therefore, they provide a case study of how quantitative techniques, such as geometric morphometrics (GMM), have impacted our ability to tease apart complex ecological signals from skeletal anatomy, and the implications for our understanding of the relationships between form, function and ecological specialization. This review provides a synthesis of current research on carnivoran ecomorphology, with the goal of illustrating the complex interaction between ecology and morphology in the skeleton. We explore the ecomorphological diversity across major carnivoran lineages and anatomical systems. We examine cranial elements (skull, sensory systems) and postcranial elements (limbs, vertebral column) to reveal mosaic patterns of adaptation related to feeding and hunting strategies, locomotion and habitat preference. We highlight the crucial role that new approaches have played in advancing our understanding of carnivoran ecomorphology, while addressing challenges that remain in the field, such as ecological classifications, form-function relationships and multi-element analysis, offering new avenues for future research.

Strategies of vascularization of the ethmoid labyrinth in selected even-toed ungulates (Artiodactyla) and carnivores (Carnivora).

The anatomy of the nasal cavity and its structures, as well as other elements building a scaffold for olfactory organs, differs significantly among various groups of mammals. Understanding anatomical conditions of quality of olfaction are being studied worldwide and is a complex problem. Among many studies regarding bone and epithelial structures of turbinates and connected anatomical structures, few studies describe the vascularization of turbinates. Ethmoid turbinates are above all covered in olfactory epithelium containing branched axons that receive olfactory stimuli and as olfactory nerves penetrate the cribriform lamina of the ethmoid bone conveying information from smell receptors to the brain. Differences in vascularization of the cribriform plate and turbinates may add crucial information complementing studies regarding the olfactory organ's bone and soft tissue structures. In the study, we describe the vascularization of the cribriform plate of the ethmoid bone of 54 Artiodactyla and Carnivora.

Topographically distinct adaptive landscapes for teeth, skeletons, and size explain the adaptive radiation of Carnivora (Mammalia).

Models of adaptive radiation were originally developed to explain the early, rapid appearance of distinct modes of life within diversifying clades. Phylogenetic tests of this hypothesis have yielded limited support for temporally declining rates of phenotypic evolution across diverse clades, but the concept of an adaptive landscape that links form to fitness, while also crucial to these models, has received more limited attention. Using methods that assess the temporal accumulation of morphological variation and estimate the topography of the underlying adaptive landscape, I found evidence of an early partitioning of mandibulo-dental morphological variation in Carnivora (Mammalia) that occurs on an adaptive landscape with multiple peaks, consistent with classic ideas about adaptive radiation. Although strong support for this mode of adaptive radiation is present in traits related to diet, its signal is not present in body mass data or for traits related to locomotor behavior and substrate use. These findings suggest that adaptive radiations may occur along some axes of ecomorphological variation without leaving a signal in others and that their dynamics are more complex than simple univariate tests might suggest.

A new gigantic carnivore (Carnivora, Amphicyonidae) from the late middle Miocene of France.

Serravallian terrestrial vertebrates are very uncommon in the northern margin of the Pyrenean Mountains. A mandible of a new large sized amphicyonid (ca. 200 kg) is here described from the marine deposits of Sallepisse (12.8-12.0 Mya). Despite that this new taxon is close in size to some European amphicyonids from the Miocene (e.g., Amphicyon, Megamphicyon, and Magericyon), the unique morphology of its p4, unknown in this clade, allows the erection of the new genus Tartarocyon cazanavei nov. gen. & sp. This taxon may be derived from a Cynelos-type amphicyonine. The description of this new taxon highlights the erosion of the ecological and morphological diversity of the Amphicyonidae in response to well-known Miocene events (i.e., Proboscidean Datum Event, Middle Miocene Climatic Transition, Vallesian Crisis).

Why big brains? A comparison of models for both primate and carnivore brain size evolution.

Despite decades of research, much uncertainty remains regarding the selection pressures responsible for brain size variation. Whilst the influential social brain hypothesis once garnered extensive support, more recent studies have failed to find support for a link between brain size and sociality. Instead, it appears there is now substantial evidence suggesting ecology better predicts brain size in both primates and carnivores. Here, different models of brain evolution were tested, and the relative importance of social, ecological, and life-history traits were assessed on both overall encephalisation and specific brain regions. In primates, evidence is found for consistent associations between brain size and ecological factors, particularly diet; however, evidence was also found advocating sociality as a selection pressure driving brain size. In carnivores, evidence suggests ecological variables, most notably home range size, are influencing brain size; whereas, no support is found for the social brain hypothesis, perhaps reflecting the fact sociality appears to be limited to a select few taxa. Life-history associations reveal complex selection mechanisms to be counterbalancing the costs associated with expensive brain tissue through extended developmental periods, reduced fertility, and extended maximum lifespan. Future studies should give careful consideration of the methods chosen for measuring brain size, investigate both whole brain and specific brain regions where possible, and look to integrate multiple variables, thus fully capturing all of the potential factors influencing brain size.

Giant Hyena Pachycrocuta brevirostris (Hyaenidae, Carnivora) from the Lower Pleistocene of Taurida Cave, Crimea.

The dental remains of a giant hyena Pachycrocuta brevirostris (Gervais, 1850) from the Early Pleistocene locality of the Taurida cave (Crimea, Late Villafranchian, 1.8-1.5 Ma) are described. This species was a typical representative of the Villafranchian fauna of the Eastern Mediterranean. The Taurida cave was occasionally used by hyenas and other carnivorans as a den and retreat.

Evolutionary history or function? Which preponderates in the expression of the muscle mass of the thoracic limb in wild carnivorans?

Thoracic limbs are extremely versatile and exhibit informative characteristics about habits of the Carnivora order in the wild. Despite this relevance, comparative studies with quantitative variables on thoracic limb muscles are still scarce in carnivorans. The aims of this study were to measure the mass of the intrinsic muscles of the thoracic limb of neotropical species of the Carnivora order and to establish comparative conjectures. For this purpose, 39 thoracic limbs of 10 neotropical carnivorans species were dissected. The mass of each muscle was measured on a digital scale, muscles were grouped by function, and the mass average percentage that each functional group of muscles occupied in the thoracic limb was calculated. The data of the present study was added to that available in the literature for 22 other carnivoran species. Three groups of species were considered: Canidae, Musteloidea, and Feliformia. Comparatively, the eight canid species included in this analyses concentrate muscle mass proximally in the thoracic limb to prioritize essential cursoriality. The nine musteloids had more muscle mass in the distal muscles due to the demand for versatility and manual strength, and the 14 Feliformia species exhibited an intermediate trend. The analysis of clusters revealed a great overlap of the percentage distribution of muscle mass with the phylogeny previously established for carnivorans. It could be verified that the distribution of muscle masses meets the demand of the locomotor habits of the species up to a certain level, from which phylogeny begins to limit morphological adaptations.

Os membros torácicos são extremamente versáteis e exibem características informativas sobre os hábitos das espécies da ordem Carnivora em vida livre. A despeito desta relevência, estudos comparativos com variéveis quantitativas dos músculos do membro torácico são escassos em carnívoros. Os objetivos deste estudo foram aferir a massa dos músculos intrínsecos do membro torácico de espécies neotropicais da ordem Carnivora e estabelecer conjecturas comparativas. Para este fim, foram dissecados 39 membros torácicos de dez espécies de carnívoros neotropicais. A massa de cada músculo foi mensurada em uma balança digital, os músculos foram agrupados por função e calculou-se a massa percentual média a que cada grupo funcional de músculos correspondia no membro torácico. Os dados do presente estudo foram analisados conjuntamente às informações disponíveis na literatura para outras 22 espécies da ordem Carnivora. Três grupos de espécies foram considerados: Canidae, Musteloidea e Feliformia. Comparativamente, as oito espécies de canídeos incluídas nesta análise demonstraram concentração de massa muscular mais proximalmente no membro torácico em razão dos hábitos essencialmente cursoriais. As nove espécies da superfamília Musteloidea exibiram mais massa nos músculos distais devido à maior demanda por versatilidade e força manual. As 14 espécies da subordem Feliformia apresentaram um padrão intermediário. A análise de clusters revelou nítida sobreposição entre o percentual de distribuição da massa muscular com a filogenia previamente estabelecida para a ordem Carnivora. Verificou-se que a distribuição da massa dos músculos atende à demanda dos hábitos locomotores das espécies até um certo nível, a partir do qual a filogenia começa a limitar as adaptações morfológicas.

Quantifying morphological adaptations using direct measurements: The carnivoran appendicular skeleton as a case study.

Here, I study whether locomotor adaptations can be detected in limb bones using a univariate approach, and whether those results are affected by size and/or shared evolutionary history. Ultimately, it tests whether classical papers on locomotor adaptations should be trusted. To do that, I analyzed the effect of several factors (size, taxonomic group, and locomotor habit) on limb bone morphology using a set of 43 measurements of the scapula, long bones, and calcaneus, of 435 specimens belonging to 143 carnivoran species. Size was the main factor affecting limb morphology. Size-corrected analyses revealed artifactual differences between various locomotion-related categories in the analyses of raw data. Additionally, several between-group differences were new to the size-corrected analyses, suggesting that they were masked by the size-effect. Phylogeny had also an important effect, although it only became apparent after removing the effect of size, probably due to the strong covariation of both factors. Regarding locomotor adaptations, locomotor type was used to represent locomotor specialization, and utilized habitat as an indicator of the capacity to adopt different modes of locomotion (running, swimming, climbing, and digging) and thus maximize resource exploitation by being capable of navigating all the substrates in the habitat they use. Locomotor type produced better results than utilized habitat, suggesting that carnivorans use locomotor specialization to minimize locomotion costs. The characteristic limb bone morphology for each locomotor type studied is described, including several adaptations and trends that are novel to the present study. Finally, the results presented here support the hypothesis of a "viverrid-like", forest-dwelling carnivoran ancestor, either arboreal or terrestrial.

The hippocampal formation of two carnivore species: The feliform banded mongoose and the caniform domestic ferret.

Employing cyto-, myelo-, and chemoarchitectural staining techniques, we analyzed the structure of the hippocampal formation in the banded mongoose and domestic ferret, species belonging to the two carnivoran superfamilies, which have had independent evolutionary trajectories for the past 55 million years. Our observations indicate that, despite the time since sharing a last common ancestor, these species show extensive similarities. The four major portions of the hippocampal formation (cornu Ammonis, dentate gyrus, subicular complex, and entorhinal cortex) were readily observed, contained the same internal subdivisions, and maintained the topological relationships of these subdivisions that could be considered typically mammalian. In addition, adult hippocampal neurogenesis was observed in both species, occurring at a rate similar to that observed in other mammals. Despite the overall similarities, several differences to each other, and to other mammalian species, were observed. We could not find evidence for the presence of the CA2 and CA4 fields of the cornu Ammonis region. In the banded mongoose the dentate gyrus appears to be comprised of up to seven lamina, through the sublamination of the molecular and granule cell layers, which is not observed in the domestic ferret. In addition, numerous subtle variations in chemoarchitecture between the two species were observed. These differences may contribute to an overall variation in the functionality of the hippocampal formation between the species, and in comparison to other mammalian species. These similarities and variations are important to understanding to what extent phylogenetic affinities and constraints affect potential adaptive evolutionary plasticity of the hippocampal formation.

The amygdaloid body of two carnivore species: The feliform banded mongoose and the caniform domestic ferret.

The current study provides an analysis of the cytoarchitecture, myeloarchitecture, and chemoarchitecture of the amygdaloid body of the banded mongoose (Mungos mungo) and domestic ferret (Mustela putorius furo). Using architectural and immunohistochemical stains, we observe that the organization of the nuclear and cortical portions of the amygdaloid complex is very similar in both species. The one major difference is the presence of a cortex-amygdala transition zone observed in the domestic ferret that is absent in the banded mongoose. In addition, the chemoarchitecture is, for the most part, quite similar in the two species, but several variances, such as differing densities of neurons expressing the calcium-binding proteins in specific nuclei are noted. Despite this, certain aspects of the chemoarchitecture, such as the cholinergic innervation of the magnocellular division of the basal nuclear cluster and the presence of doublecortin expressing neurons in the shell division of the accessory basal nuclear cluster, appear to be consistent features of the Eutherian mammal amygdala. The domestic ferret presented with an overall lower myelin density throughout the amygdaloid body than the banded mongoose, a feature that may reflect artificial selection in the process of domestication for increased juvenile-like behavior in the adult domestic ferret, such as a muted fear response. The shared, but temporally distant, ancestry of the banded mongoose and domestic ferret allows us to generate observations relevant to understanding the relative influence that phylogenetic constraints, adaptive evolutionary plasticity, and the domestication process may play in the organization and chemoarchitecture of the amygdaloid body.

The diencephalon of two carnivore species: The feliform banded mongoose and the caniform domestic ferret.

This study provides an analysis of the cytoarchitecture, myeloarchitecture, and chemoarchitecture of the diencephalon (dorsal thalamus, ventral thalamus, and epithalamus) of the banded mongoose (Mungos mungo) and domestic ferret (Mustela putorius furo). Using architectural and immunohistochemical stains, we observe that the nuclear organization of the diencephalon is very similar in the two species, and similar to that reported in other carnivores, such as the domestic cat and dog. The same complement of putatively homologous nuclei were identified in both species, with only one variance, that being the presence of the perireticular nucleus in the domestic ferret, that was not observed in the banded mongoose. The chemoarchitecture was also mostly consistent between species, although there were a number of minor variations across a range of nuclei in the density of structures expressing the calcium-binding proteins parvalbumin, calbindin, and calretinin. Thus, despite almost 53 million years since these two species of carnivores shared a common ancestor, strong phylogenetic constraints appear to limit the potential for adaptive evolutionary plasticity within the carnivore order. Apart from the presence of the perireticular nucleus, the most notable difference between the species studied was the physical inversion of the dorsal lateral geniculate nucleus, as well as the lateral posterior and pulvinar nuclei in the domestic ferret compared to the banded mongoose and other carnivores, although this inversion appears to be a feature of the Mustelidae family. While no functional sequelae are suggested, this inversion is likely to result from the altricial birth of Mustelidae species.

Evolutionary and morphological patterns underlying carnivoran body shape diversity.

The diversity of body shapes is one of the most prominent features of phenotypic variation in vertebrates. Biologists, however, still lack a full understanding of the underlying morphological components that contribute to its diversity, particularly in endothermic vertebrates such as mammals. In this study, hypotheses pertaining to the evolution of the cranial and axial components that contribute to the diversity of carnivoran body shapes were tested. Three trends were found in the evolution of carnivoran body shapes: (1) carnivorans exhibit diverse body shapes with intrafamilial variation predicted best by family clade age, (2) body shape is driven by strong allometric effects of body size where species become more elongate with decreasing size, and (3) the thoracic and lumbar regions and rib length contribute the most to body shape variation, albeit pathways differ between different families. These results reveal the morphological patterns that led to increased diversity in carnivoran body shapes and elucidate the similarities and dissimilarities that govern body shape diversity across vertebrates.

Brain Volume Fractions in Mammals in Relation to Behavior in Carnivores, Primates, Ungulates, and Rodents.

The volume fraction (VF) of a given brain region, or the proper mass, ought to reflect the importance of that region in the life of a given species. This study sought to examine the VF of various brain regions across 61 different species of mammals to discern if there were regularities or differences among mammalian orders. We examined the brains of carnivores (n = 17), ungulates (n = 8), rodents (n = 7), primates (n = 11), and other mammals (n = 18) from the online collections at the National Museum of Health and Medicine. We measured and obtained the VF of several brain regions: the striatum, thalamus, neocortex, cerebellum, hippocampus, and piriform area. We refined our analyses by using phylogenetic size correction, yielding the corrected (c)VF. Our groups showed marked differences in gross brain architecture. Primates and carnivores were divergent in some measures, particularly the cVF of the striatum, even though their overall brain size range was roughly the same. Rodents predictably had relatively large cVFs of subcortical structures due to the fact that their neocortical cVF was smaller, particularly when compared to primates. Not so predictably, rodents had the largest cerebellar cVF, and there were marked discrepancies in cerebellar data across groups. Ungulates had a larger piriform area than primates, perhaps due to their olfactory processing abilities. We provide interpretations of our results in the light of the comparative behavioral and neuroanatomical literature.

Morphological evolution of the carnivoran sacrum.

The sacrum is a key piece of the vertebrate skeleton, since it connects the caudal region with the presacral region of the vertebral column and the hind limbs through the pelvis. Therefore, understanding its form and function is of great relevance in vertebrate ecomorphology. However, it is striking that morphometric studies that quantify its morphological evolution in relation to function are scarce. The main goal of this study is to investigate the morphological evolution of the sacrum in relation to its function in the mammalian order Carnivora, using three-dimensional (3D) geometric morphometrics. Principal component analysis under a phylogenetic background indicated that changes in sacrum morphology are mainly focused on the joint areas where it articulates with other parts of the skeleton allowing resistance to stress at these joints caused by increasing muscle loadings. In addition, we demonstrated that sacrum morphology is related to both the length of the tail relativised to the length of the body, and the length of the body relativised to body mass. We conclude that the sacrum in carnivores has evolved in response to the locomotor requirements of the species analysed, but in locomotion, each family has followed alternative morphological solutions to address the same functional demands.

Clavicle in Carnivorans: A Forgotten Bone.

The clavicle is a bone whose development is related to the versatility of the thoracic limbs in mammals. Studies with vestigial structures are scarce and controversial, especially in the order Carnivora. The objective of this study was to verify the presence and to investigate the shape and constitution of the clavicle in neotropical carnivores. In order to do this, 108 cadavers of 19 different species were collected dead on highways and were analyzed. The clavicles were submitted to dissections, longitudinal length measurements, radiographs, histological sections and, in some cases, diaphanization. Sixteen of the 19 species had clavicles in both sides, being significantly larger (P < 0.05) and distinctly more radiopaque in the felids than in the other families. There were no macro or microscopic evidence of clavicle in the specimens of Nasua nasua (n = 6), Conepatus semistriatus (n = 2), and Conepatus chinga (n = 1). The clavicle of the males of Lycalopex gymnocercus, Galictis cuja, and Leopardus geoffroyi was significantly larger (P < 0.05). The predominating contour of the clavicles was a thin stick with cranial convexity. The histological sections demonstrated compact bone consisting of trabecula and lamellae filled by bone marrow and different levels of occupation by chondroid matrix. It can be proposed the clavicles of the order Carnivora, although vestigial and rarely absent, have their presence, constitution and shape more associated with the phylogenetic proximity and evolutionary history of the species than to the variety of movements the thoracic limbs perform in free-living conditions. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:1831-1841, 2020. © 2019 American Association for Anatomy.

Measuring the magnitude of morphological integration: The effect of differences in morphometric representations and the inclusion of size.

The magnitude of morphological integration is a major aspect of multivariate evolution, providing a simple measure of the intensity of association between morphological traits. Studies concerned with morphological integration usually translate phenotypes into morphometric representations to quantify how different morphological elements covary. Geometric and classic morphometric representations translate biological form in different ways, raising the question if magnitudes of morphological integration estimates obtained from different morphometric representations are compatible. Here we sought to answer this question using the relative eigenvalue variance of the covariance matrix obtained for both geometric and classical representations of empirical and simulated datasets. We quantified the magnitude of morphological integration for both shape and form and compared results between representations. Furthermore, we compared integration values between shape and form to evaluate the effect of the inclusion or not of size on the quantification of the magnitude of morphological integration. Results show that the choice of morphological representation has significant impact on the integration magnitude estimate, either for shape or form. Despite this, ordination of the integration values within representations is relatively the same, allowing for similar conclusions to be reached using different methods. However, the inclusion of size in the dataset significantly changes the estimates of magnitude of morphological integration, hindering the comparison of this statistic obtained from different spaces. Morphometricians should be aware of these differences and must consider how biological hypothesis translate into predictions about integration in each particular choice of representation.

Bite Force and Masticatory Muscle Architecture Adaptations in the Dietarily Diverse Musteloidea (Carnivora).

Dietary ecology and its relationship with both muscle architecture and bite force potential has been studied in many mammalian (and non-mammalian) taxa. However, despite the diversity of dietary niches that characterizes the superfamily Musteloidea, the masticatory muscle fiber architecture of its members has yet to be investigated anatomically. In this study, we present myological data from the jaw adductors in combination with biomechanical data derived from craniomandibular measurements for 17 species representing all four families (Ailuridae, Mephitidae, Mustelidae, and Procyonidae) of Musteloid. These data are combined to calculate bite force potential at each of three bite points along the dental row. Across our sample as a whole, masticatory muscle mass scaled with isometry or slight positive allometry against both body mass and skull size (measured via a cranial geometric mean). Total jaw adductor physiological cross-sectional area scaled with positive allometry against both body mass and skull size, while weighted fiber length scaled with negative allometry. From a dietary perspective, fiber length is strongly correlated with dietary size such that taxa that exploit larger foods demonstrated myological adaptations toward gape maximization. However, no consistent relationship between bite force potential and dietary mechanical resistance was observed. These trends confirm previous findings observed within the carnivoran family Felidae (as well as within primates), suggesting that the mechanisms by which masticatory anatomy adapts to dietary ecology may be more universally consistent than previously recognized. Anat Rec, 302:2287-2299, 2019. © 2019 American Association for Anatomy.

Bayesian Estimation of Species Divergence Times Using Correlated Quantitative Characters.

Discrete morphological data have been widely used to study species evolution, but the use of quantitative (or continuous) morphological characters is less common. Here, we implement a Bayesian method to estimate species divergence times using quantitative characters. Quantitative character evolution is modeled using Brownian diffusion with character correlation and character variation within populations. Through simulations, we demonstrate that ignoring the population variation (or population "noise") and the correlation among characters leads to biased estimates of divergence times and rate, especially if the correlation and population noise are high. We apply our new method to the analysis of quantitative characters (cranium landmarks) and molecular data from carnivoran mammals. Our results show that time estimates are affected by whether the correlations and population noise are accounted for or ignored in the analysis. The estimates are also affected by the type of data analyzed, with analyses of morphological characters only, molecular data only, or a combination of both; showing noticeable differences among the time estimates. Rate variation of morphological characters among the carnivoran species appears to be very high, with Bayesian model selection indicating that the independent-rates model fits the morphological data better than the autocorrelated-rates model. We suggest that using morphological continuous characters, together with molecular data, can bring a new perspective to the study of species evolution. Our new model is implemented in the MCMCtree computer program for Bayesian inference of divergence times.

Taking a look into the orbit of mammalian carnivorans.

In this study, we explore the relationship between orbit anatomy and different ecological factors in carnivorous mammals from a phylogenetic perspective. We calculated the frontation (α), convergence (ß), and orbitotemporal (Ω) angles of the orbit from 3D coordinates of anatomical landmarks in a wide sample of carnivores with different kinds of visual strategy (i.e. photopic, scotopic, and mesopic), habitat (i.e. open, mixed, and closed), and substrate use (i.e. arboreal, terrestrial, and aquatic). We used Bloomberg's K and Pagel's λ to assess phylogenetic signal in frontation, convergence, and orbitotemporal angles. The association of orbit orientation with skull length and ecology was explored using phylogenetic generalized least squares and phylogenetic manova, respectively. Moreover, we also computed phylomorphospaces from orbit orientation. Our results indicate that there is not a clear association between orbit orientation and the ecology of living carnivorans. We hypothesize that the evolution of the orbit in mammalian carnivores represents a new case of an ecological bottleneck specific to carnivorans. New directions for future research are discussed in light of this new evidence.

The effect of the masticatory muscle physiological cross-sectional area on the structure of the temporomandibular joint in Carnivora.

We compared the temporomandibular joint structure between species of the order Carnivora and investigated its variation among family lineages. We also investigated the effect of the masticatory muscle physiological cross-sectional area (PCSA) on temporomandibular joint structure. The masticatory muscle is composed of multiple muscles, which contract in different directions and exert pressure on the temporomandibular joint. We investigated the effect of the ratio of each muscle's PCSA-an indicator of muscle force-and muscle size relative to body size on temporomandibular joint structure. The temporalis PCSA relative to body size showed the highest correlation with temporomandibular joint structure. When the temporalis PCSA is large relative to body size, the preglenoid projects caudally, the postglenoid projects rostrally and the pre-postglenoid angle interval is small, indicating that the condyle is locked in the fossa to reinforce the temporomandibular joint. Most Carnivora use blade-like carnassial teeth when slicing food. However, dislocation occurs when the carnassial teeth are used by the temporalis muscle. Our results suggest that the temporomandibular joint is reinforced to prevent dislocation caused by the temporalis muscle. In Mustelidae, the temporomandibular joint with a rostrally projecting postglenoid is suitable for carnassial biting using the temporalis muscle. In Felidae, the force of the masseter onto the carnassial teeth is diverted to the canine by tightening the temporomandibular joint. In Canidae, the masticatory muscle arrangement is well-balanced, enabling combined action. Hence, reinforcement of the temporomandibular joint by bone structure is unnecessary.