Comparative bite forces and canine bending strength in feline and sabretooth felids: implications for predatory ecology

The sabretooth felids were widespread across much of the world in the Late Tertiary, and appear to have been an important group of large predators. Owing to the substantially different skull morphology of derived sabretooths compared with extant felids, there has been considerable debate over the killing mode, bite forces, and bending strength of the large upper canines, and over the implications of these characteristics on feeding ecology. Debates have, however, usually been based on indirect comparisons of force vectors. In this paper, I provide assessments of the estimated force output from the jaw adductor muscles, based on estimates of muscle cross-sectional areas and force vectors, along with canine bending strengths, in a variety of sabretooth felids, in comparison with extant felids. In general, sabretoothed felids had moderately powerful bites, albeit with less jaw adductor power for their body sizes compared with extant felids, sometimes markedly so. Less derived sabrecats appear to have had proportionally higher bite forces than derived forms. The length of the upper canines seemingly compromised their bending strength at any given body size, and again this was most marked in derived forms. However, compared with estimated jaw adductor forces, the canines of sabrecats appear, if anything, to have been stronger than those of extant conical-toothed felids. It has previously been suggested that large sabretoothed felids hunted large prey with a canine shearing bite, powered in part by the jaw adductors and in part by the muscles of the upper neck–occipital region. The present results of canine bending strengths versus the predicted bite force from the jaw adductors supports this suggestion.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 423–437.     

Directional selection in the evolution of elongated upper canines in clouded leopards and sabre‐toothed cats

Extremely developed or specialized traits such as the elongated upper canines of extinct sabre‐toothed cats are often not analogous to those of any extant species, which limits our understanding of their evolutionary cause. However, an extant species may have undergone directional selection for a similar extreme phenotype. Among living felids, the clouded leopard, Neofelis nebulosa, has exceptionally long upper canines for its body size. We hypothesized that directional selection generated the elongated upper canines of clouded leopards in a manner similar to the process in extinct sabre‐toothed cats. To test this, we developed an approach that compared the effect of directional selection among lineages in a phylogeny using a simulation of trait evolution and approximate Bayesian computation. This approach was applied to analyse the evolution of upper canine length in the Felidae phylogeny. Our analyses consistently showed directional selection favouring longer upper canines in the clouded leopard lineage and a lineage leading to the sabre‐toothed cat with the longest upper canines, Smilodon. Most of our analyses detected an effect of directional selection for longer upper canines in the lineage leading to another sabre‐toothed cat, Homotherium, although this selection may have occurred exclusively in the primitive species. In all the analyses, the clouded leopard and Smilodon lineages showed comparable directional selection. This implies that clouded leopards share a selection advantage with sabre‐toothed cats in having elongated upper canines.     

A dynamic model for the evolution of sabrecat predatory bite mechanics

The ability of sabretoothed felids to achieve sufficiently high bite forces for predation at extreme gape angles has been the subject of decades of debate. Previous studies have indicated that bite forces in derived sabretoothed felids would have been low, but that they were probably augmented by head depressing muscles. However, bite mechanics is a dynamic process, and mechanical properties change with changes in gape angles. In this study, I present the first comprehensive model of bite mechanics, vector angles, and forces about the temporomandibular joint at gape angles from occlusion to maximal inferred gape in sabretoothed felids. Primitive sabrecats (Machairodus, Paramachairodus) appear broadly comparable to extant large felids (Panthera, Puma), but derived sabrecats in the groups Homotherini (Amphimachairodus, Homotherium, Xenosmilus) and Smilodontini (Megantereon, Smilodon) are often substantially different from either of the former. The ability of the mandibular adductors to generate torque changes with gape angle, indicating that previous models fail to capture potentially important differences in bite function. Inferred muscle sizes and the angles of effective torque from individual adductor fibres in derived sabrecats are different from those of primitive sabrecats and extant large felids, but they had evolved a number of compensatory adaptations for maximizing force output at the canine and carnassial, primarily changes in muscle fibre angles and more compact crania. Inferred outforces at the canines and carnassials were comparable amongst all groups at low gape angles, but at extreme gape angles outforces would have been low, supporting previous hypotheses of head flexor contribution during initial stages of the killing bite in sabrecats. Mandibular adduction in extant carnivores is a complicated pattern of differences in twitch tension and electromyographical activity at different gape angles, and inference of maximal isotonic bite forces from reconstructed mandibular adductor sizes in fossils will give estimates primarily suitable for comparative purposes. Potentially, derived sabrecats could have evolved differences from extant felids in adductor histochemistry or pinnation angle of individual fibres. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 220–242.     

Mandibular shape correlates of tooth fracture in extant Carnivora: implications to inferring feeding behaviour of Pleistocene predators

Percentages of tooth fracture and mandible shape are robust predictors of feeding habits in Carnivora. If these parameters co‐vary above the species level, more robust palaeobiological inferences could be made on fossil species. A test of association is presented between mandible shape and tooth fracture in a subset of extant carnivorans together with large Pleistocene fossil predators from Rancho La Brea (Canis dirus, Panthera atrox, and Smilodon fatalis). Partial least square (PLS) and comparative methods are employed to validate co‐variation of these two parameters in extant carnivorans. Association between mandible shape and percentage of tooth fracture is strongly supported, even if both blocks of data exhibit a phylogenetic signal to a different degree. Dietary adaptations drive shape/fracture co‐variation in extant species, although no significant differences occur in the PLS scores between carnivores and bone/hard food consumers. The fossil species project into PLS morphospace as outliers. Their position suggests a unique feeding behaviour. The increase in the size of prey, together with consumption of skin and hair from carcasses in a cold environment, might have generated unusual tooth breakage patterns in large predators from Rancho La Brea. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 70–80.     

Sabretoothed carnivores and the killing of large prey

Sabre-like canines clearly have the potential to inflict grievous wounds leading to massive blood loss and rapid death. Hypotheses concerning sabretooth killing modes include attack to soft parts such as the belly or throat, where biting deep is essential to generate strikes reaching major blood vessels. Sabretoothed carnivorans are widely interpreted as hunters of larger and more powerful prey than that of their present-day nonsabretoothed relatives. However, the precise functional advantage of the sabretooth bite, particularly in relation to prey size, is unknown. Here, we present a new point-to-point bite model and show that, for sabretooths, depth of the killing bite decreases dramatically with increasing prey size. The extended gape of sabretooths only results in considerable increase in bite depth when biting into prey with a radius of less than ～10 cm. For sabretooths, this size-reversed functional advantage suggests predation on species within a similar size range to those attacked by present-day carnivorans, rather than "megaherbivores" as previously believed. The development of the sabretooth condition appears to represent a shift in function and killing behaviour, rather than one in predator-prey relations. Furthermore, our results demonstrate how sabretoothed carnivorans are likely to have evolved along a functionally continuous trajectory: beginning as an extension of a jaw-powered killing bite, as adopted by present-day pantherine cats, followed by neck-powered biting and thereafter shifting to neck-powered shear-biting. We anticipate this new insight to be a starting point for detailed study of the evolution of pathways that encompass extreme specialisation, for example, understanding how neck-powered biting shifts into shear-biting and its significance for predator-prey interactions. We also expect that our model for point-to-point biting and bite depth estimations will yield new insights into the behaviours of a broad range of extinct predators including therocephalians (gorgonopsian + cynodont, sabretoothed mammal-like reptiles), sauropterygians (marine reptiles) and theropod dinosaurs.     

Elbow-joint morphology as a guide to forearm function and foraging behaviour in mammalian carnivores

Among the hunting strategies employed by members of the order Carnivora (Mammalia), two, stalk and ambush and sustained pursuit, are particularly prevalent among larger species of the order. It has been difficult to identify morphological traits that support this distinction and ecological observations have shown that most carnivorans adopt a continuum of strategies, depending on available habitat and prey. In this paper, the shape of the distal humerus articulation is analysed, with the aim of exploring the use of the forelimb in prey procurement, and as a guide to such behaviour among fossil carnivorans. The results suggest that manual manipulation and locomotion are conflicting functions. Elbow‐joint morphology supports a division between grapplers (i.e. ambushers) and nongrapplers (i.e. pursuers). Joints of the former are characterized by being relatively wide and the latter, by being relatively narrow and box‐like with pronounced stabilizing features. At intermediate and large body sizes, carnivorans show a pattern suggesting mutually exclusive feeding strategies that involve either grappling with prey or sustained pursuit. The former allows for large body sizes, such as pantherine felids and ursids; the latter includes species of only moderate size, such as hyenids and canids. Elbow‐joint morphology is closely linked to phylogeny, but the morphology of the cheetah converges with that of nongrapplers, showing that strong selective forces may override the phylogenetic component. Two taxa of giant mustelids from the Miocene were analysed to test whether this sort of analysis is applicable to carnivorans of the past. The African Late Miocene species Ekorus ekakeran has a joint morphology comparable to that of modern‐day nongrapplers. Two joint morphologies were found in the North American Late Oligocene‐Early Miocene Megalictis ferox. The first morphology is comparable to that of modern pantherine cats and the second forms an intermediate between grapplers and nongrapplers that is not present in the modern carnivoran fauna. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142 , 91–104.     

Archaeocete-like jaws in a baleen whale

The titanic baleen whales (Cetacea, Mysticeti) have a bizarre skull morphology, including an elastic mandibular symphysis, which permits dynamic oral cavity expansion during bulk feeding. How this key innovation evolved from the sutured symphysis of archaeocetes has remained unclear. Now, mandibles of the Oligocene toothed mysticete Janjucetus hunderi show that basal mysticetes had an archaeocete-like sutured symphysis. This archaic morphology was paired with a wide rostrum typical of later-diverging baleen whales. This demonstrates that increased oral capacity via rostral widening preceded the evolution of mandibular innovations for filter feeding. Thus, the initial evolution of the mysticetes' unique cranial form and huge mouths was perhaps not linked to filtering plankton, but to enhancing suction feeding on individual prey.     

Maximum Bite Force and Prey Size of Tyrannosaurus rex and Their Relationships to the Inference of Feeding Behavior

The feeding behavior of the theropod dinosaur Tyrannosaurus rex is investigated through analysis of two variables that are critical to successful predation, bite force and prey body mass, as they scale with the size of the predator. These size-related variables have important deterministic effects on the predator’s feeding strategy, through their effects on lethal capacity and choice of prey. Bite force data compiled for extant predators (crocodylians, carnivorans, chelonians and squamates) are used to establish a relationship between bite force and body mass among extant predators. These data are used to estimate the maximum potential bite force of T. rex, which is between about 183,000 and 235,000 N for a bilateral bite. The relationship between maximum prey body mass and predator body mass among the same living vertebrates is used to infer the likely maximum size of prey taken by T. rex in the Late Cretaceous. This makes it possible to arrive at a more rigorous assessment of the role of T. rex as an active predator and/or scavenger than has hitherto been possible. The results of this analysis show that adult Triceratops horridus fall well within the size range of potential prey that are predicted to be available to a solitary, predaceous T. rex. This analysis establishes boundary conditions for possible predator/prey relationships among other dinosaurs, as well as between these two taxa.     

Craniodental indicators of prey size preference in the Felidae

In the present study, we used linear morphometrics of the crania, mandible and dentition to explore the association between craniodental shape and prey size among 35 species of living felids. To accomplish this, felids were divided into three prey-size groups: (1) large prey specialists; (2) small prey specialists; and (3) mixed prey feeders. From these linear measurements, large prey specialist felids can be distinguished from small and mixed prey feeders by their relatively robust canines and incisors and relatively wide muzzles. These cranial characters are advantageous when dispatching large prey, due to the stranglehold that cats employ during this activity. Robust canines resist the bending and torsional forces applied by struggling prey and a wider muzzle helps to stabilize grip and distribute bite forces more evenly during the killing bite. Small prey specialists had smaller canines, narrower muzzles and slightly longer jaws for a speed advantage when catching small, quick prey. Mixed prey feeders were intermediate between large and small prey specialists, indicating they are adapted to killing both sizes of prey. Given the success of this ecomorphological analysis of living felids that specialize on different prey sizes, we look forward to applying this same approach to extinct species.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 784–799.     

The morphology and systematics of Mammalodon colliveri (Cetacea: Mysticeti), a toothed mysticete from the Oligocene of Australia

Mammalodon colliveri is an unusual toothed archaic mysticete (Cetacea) from the Upper Oligocene Jan Juc Formation of south‐east Australia. The morphology of the holotype skull and postcrania are described in detail. Superimposed on the generally plesiomorphic archaeocete‐like morphology of Mammalodon are subtle mysticete synapomorphies. Derived features of Mammalodon include a short and bluntly rounded rostrum, reduced premaxillae, and anterodorsally directed orbits. Within Mysticeti, this suite of features is unique. The aberrant rostral morphology of Mammalodon suggests specialization for suction feeding. Janjucetus hunderi is placed in an expanded family Mammalodontidae. Phylogenetic analysis corroborates the monophyly of Basilosauridae, Neoceti, Odontoceti, and Mysticeti, and yields a novel hypothesis of toothed mysticete relationships: a basal clade of undescribed toothed mysticetes from South Carolina, a Llanocetidae + Mammalodontidae clade, and monophyletic Aetiocetidae are posited as successive sister taxa to edentulous baleen whales (Chaeomysticeti). Toothed archaic mysticetes clearly employed diverse prey capture strategies, with exaptations for filter feeding evolving sequentially in stem group Mysticeti. A stratigraphically calibrated phylogeny implies that the initial diversification of Mysticeti occurred during the Late Eocene. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 367–476.     

Mosaic functionality in a transitional ecomorphology: skull biomechanics in stem Hyaeninae compared to modern South African carnivorans

Ecomorphologies are categories of ecological adaptation and function, although intermediates are not always available to shed light on functionality at the transitional stages between them. We examined an intermediate bone‐cracking carnivoran ecomorphology, the stem hyaenine Ikelohyaena abronia, using finite element analysis. Skull models of Ikelohyaena, crown hyaenine Crocuta crocuta, and two other hypercarnivores were simulated with mastication and prey apprehension forces. The results obtained show that Ikelohyaena already possessed derived features in skull stress distribution and levels of strain energy, characteristic of the extant bone‐cracking Crocuta; however, the estimated bite forces in Ikelohyaena were significantly lower. Prey apprehension simulations showed similar patterns; the low skull strain energy and low bite force of the Ikelohyaena mandible indicate a poor individual ability to take down large prey. The mosaic features of craniodental function in Ikelohyaena suggest that initial evolution of the hyaenid bone‐cracking ecomorphology involved skull shape changes that increased stress dissipation, permitting incorporation of more hard food into the diet. Subsequent evolution of larger bite forces was then required to increase the size limit of bones that can be cracked and consumed. This mode of evolution would have allowed transitional hyaenid ecomorphologies to continuously increase the carcass processing ability both during competitive feeding and scavenging throughout their evolution. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 540–559.     

Tooth root morphology as an indicator for dietary specialization in carnivores (Mammalia: Carnivora)

The Carnivora occupy a wide range of feeding niches in concordance with the enormous diversity in their skull and dental form. It is well established that differences in crown morphology are linked to variations in the material properties of the foods ingested and masticated. However, how tooth root form is related to dietary specialization is less well known. In the present study, we investigate the relationship between tooth root morphology and dietary specialization in terrestrial carnivores (canids, felids, hyaenids, and ursids). We specifically address the question of how variation in tooth root surface area is related to bite force potentials as one of the crucial masticatory performance parameters in feeding ecology. We applied computed tomography imaging to reconstruct and quantify dental root surface area in 17 extant carnivore species. Moreover, we computed maximal bite force at several tooth positions based on a dry skull model and assessed the relationship of root surface area to skull size, maximal bite force, food properties, and prey size. We found that postcanine tooth root surface areas corrected for skull size serve as a proxy for bite force potentials and, by extension, dietary specialization in carnivores. Irrespective of taxonomic affinity, species that feed on hard food objects have larger tooth roots than those that eat soft or tough foods. Moreover, carnivores that prey on large animals have larger tooth root surface areas. Our results show that tooth root morphology is a useful indicator of bite force production and allows inferences to be made about dietary ecology in both extant and extinct mammals. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105, 456–471.     

Tooth Root Morphology in the Early Pliocene African Bear Agriotherium africanum (Mammalia, Carnivora, Ursidae) and its Implications for Feeding Ecology

Tooth root surface areas serve as proxies for bite force potentials, and by extension, dietary specialization in extant carnivorans. Here, we investigate the feeding ecology of the extinct large-bodied ursid Agriotherium africanum, by comparing its root surface areas (reconstructed with the aid of computed tomography and three-dimensional image processing) and bite force estimates, with those of extant carnivorans. Results show that in absolute terms, canine and carnassial bite forces, as well as root surface areas were highest in A. africanum. However, when adjusted for skull size, A. africanum’s canine roots were smaller than those of extant solitary predators. With teeth being the limiting factor in the masticatory system, low canine root surface areas suggest that A. africanum would have struggled to bring down large vertebrate prey. Its adjusted carnassial root sizes were found to be smaller than those of extant hard object feeders and the most carnivorous tough object feeders, but larger than those of extant omnivorous ursids and Ursus maritimus. This and the fact that it displayed its highest postcanine root surface areas in the carnassial region (rather than the most distal tooth in the tooth row) suggest that A. africanum consumed more vertebrate tissue than extant omnivorous ursids. With an apparent inability to routinely bring down large prey or to consume mechanically demanding skeletal elements, its focus was most likely on tough tissue, which it acquired by actively scavenging the carcasses of freshly dead/freshly killed animals. Mechanically less demanding skeletal elements would have been a secondary food source, ingested and processed mainly in association with muscle and connective tissue.     

Variation in Craniomandibular Morphology and Sexual Dimorphism in Pantherines and the Sabercat Smilodon fatalis

Sexual dimorphism is widespread among carnivorans, and has been an important evolutionary factor in social ecology. However, its presence in sabertoothed felids remains contentious. Here we present a comprehensive analysis of extant Panthera and the sabertoothed felid Smilodon fatalis. S. fatalis has been reported to show little or no sexual dimorphism but to have been intraspecifically variable in skull morphology. We found that large and small specimens of S. fatalis could be assigned to male and female sexes with similar degrees of confidence as Panthera based on craniomandibular shape. P. uncia is much less craniomandibularly variable and has low levels of sexual size-dimorphism. Shape variation in S. fatalis probably reflects sexual differences. Craniomandibular size-dimorphism is lower in S. fatalis than in Panthera except P. uncia. Sexual dimorphism in felids is related to more than overall size, and S. fatalis and the four large Panthera species show marked and similar craniomandibular and dental morphometric sexual dimorphism, whereas morphometric dimorphism in P. uncia is less. Many morphometric-sexually dimorphic characters in Panthera and Smilodon are related to bite strength and presumably to killing ecology. This suggests that morphometric sexual dimorphism is an evolutionary adaptation to intraspecific resource partitioning, since large males with thicker upper canines and stronger bite forces would be able to hunt larger prey than females, which is corroborated by feeding ecology in P. leo. Sexual dimorphism indicates that S. fatalis could have been social, but it is unlikely that it lived in fusion-fission units dominated by one or a few males, as in sub-Saharan populations of P. leo. Instead, S. fatalis could have been solitary and polygynous, as most extant felids, or it may have lived in unisexual groups, as is common in P. leo persica.     

The geography of diet variation in Neotropical Carnivora

1. Mammalian carnivores (order Carnivora) perform important regulatory functions in terrestrial food webs. Building a comprehensive knowledge of the dietary patterns of carnivorans and the factors determining such patterns is essential for improving our understanding of the role of carnivorans in ecosystem functioning.
2. In the Neotropics, there are 64 extant species of terrestrial Carnivora, but information on their trophic ecology is diffuse. We compiled and analysed the available quantitative dietary data for Neotropical carnivorans, aiming to detect patterns of intraspecific and interspecific dietary variation at a large geographical scale.
3. The resulting database encompasses information on trophic interactions of 37 native carnivoran species from six families across 14 countries. There are clear geographical biases towards southern Brazil, Chile, and Argentina, and a noticeable knowledge gap within the Amazon. Also, most studies are focused on canids and felids, especially Puma concolor, Panthera onca, Cerdocyon thous, Leopardus pardalis, and Chrysocyon brachyurus, whereas for 27 native species, we found no quantitative dietary information.
4. Neotropical carnivorans consume species from at least 651 genera of vertebrates, invertebrates, and plants. We found clear species-specific dietary patterns and marked differences between Neotropical felids and canids. Although predators generally exhibit high levels of consistency in their diets regarding prey body mass, we detected significant intraspecific variation for all species analysed across study sites.
5. Body mass imposes strong constraints on prey use, but biogeographical differences in prey availability and human influence may drive the geographical variation we found. Overall, observed patterns show not only similarities with resource-use patterns found for carnivorans in other continents, such as nestedness driven by body mass, but also differences, such as high levels of frugivory and consumption of invertebrates by canids. Assessing resource-use patterns is the first step towards a better understanding of processes underlying the organisation of trophic interactions, and is imperative for addressing impacts of defaunation on ecosystems and for informing conservation efforts.

     

Feeding mechanics in Triassic stem-group sauropterygians: the anatomy of a successful invasion of Mesozoic seas

The jaw adductor musculature in Triassic stem-group sauropterygians is reconstructed on the basis of a paradigmatic model of muscle architecture (functional equivalence of sarcomeres) and using invariant traits of the anatomy of the trigeminal jaw adductor muscles in extant reptiles. The reconstructed jaw adductor musculature predicts trophic specializations in stem-group sauropterygians. Suction feeding is a component in prey capture for some benthic feeding, as well as for some pelagic feeding taxa. The differentiation of 'pincer' jaws is correlated with the potential for rapid, snapping bites. There is some evidence for habitat partitioning among Triassic stem-group sauropterygians with respect to trophic specialization. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 135 , 33–63.     

Convergence and divergence in prey of sympatric canids and felids: opportunism or phylogenetic constraint?

Since the canids and felids diverged in the mid‐Eocene or earlier, each family has developed a suite of morphological and behavioural adaptations for obtaining and consuming prey. We here distinguish between prey taxa captured and eaten as a result of these phylogenetic adaptations, and those because they are fortuitously encountered, and argue that such supplementary prey, often opportunistically caught, create a buffer between sympatric, and potentially competitive, canids and felids and thus enhance coexistence. We base our analysis on dietary data derived from the stomach contents of four sympatric canid and felid species in the Free State Province, South Africa (canids: Cape fox Vulpes chama and black‐backed jackal Canis mesomelas; felids: African wild cat Felis silvestris lybica and caracal Caracal caracal), and from results of studies on these species elsewhere in southern Africa. The two canid species preyed heavily on invertebrates, and thus opportunistically, while the felids (especially the caracal) concentrated on mammals, prey they are phylogenetically adapted to capture. Only three species of mammalian prey are shared by the four species. The ratio of opportunistically‐to‐phylogenetically mediated prey taxa used (the O/P ratio) differ between the species, with the black‐backed jackal having the most opportunistically caught taxa in its diet, and the caracal the least. As predicted, a comparison of this data with those from dietary studies of the same species carried out elsewhere indicates that the number of opportunistically obtained prey taxa varies more than those resulting from phylogenetic adaptations. The largest canid had the widest food spectrum (35 prey taxa) while the smallest felid had the most restricted one (11 prey taxa). We argue that using the O/P distinction allows a better understanding of changes in food niche breadth of particular species, especially in xeric areas, and gives a better indication of possible exploitative competition for food by sympatric carnivores than when regarding all prey taxa as actively pursued. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83 , 527–538.     

Allometry and performance: the evolution of skull form and function in felids

Allometric patterns of skull‐shape variation can have significant impacts on cranial mechanics and feeding performance, but have received little attention in previous studies. Here, we examine the impacts of allometric skull‐shape variation on feeding capabilities in the cat family (Felidae) with linear morphometrics and finite element analysis. Our results reveal that relative bite force diminishes slightly with increasing skull size, and that the skulls of the smallest species undergo the least strain during biting. However, larger felids are able to produce greater gapes for a given angle of jaw opening, and they have overall stronger skulls. The two large felids in this study achieved increased cranial strength by increasing skull bone volume relative to surface area. Allometry of skull geometry in large felids reflects a trade‐off between the need to increase gape to access larger prey while maintaining the ability to resist unpredictable loading when taking large, struggling prey.     

Morphological investigation into functions of the jaw symphysis in carnivorans

The hemimandibles in carnivorans may be united in various ways at the symphysis menti. The symphysis may contain a readily flexible joint that permits a moderate amount of independent movement of the hemimandibles. This type of symphyseal union is primitive for and widely distributed in extant carnivorans. In other carnivorans, the symphysis is patent but allows slight or essentially no independent movement of the hemimandibles. Finally, the hemimandibles may be rigidly united by synostosis of the symphysis. The morphology, movement and, insofar as possible, function of these types of symphyses are described.     

Ecomorphology of the African felid ensemble: The role of the skull and postcranium in determining species segregation and assembling history

Morphology of extant felids is regarded as highly conservative. Most previous studies have focussed on skull morphology, so a vacuum exists about morphofunctional variation in postcranium and its role in structuring ensembles of felids in different continents. The African felid ensemble is particularly rich in ecologically specialized felids. We studied the ecomorphology of this ensemble using 31 cranial and 93 postcranial morphometric variables measured in 49 specimens of all 10 African species. We took a multivariate approach controlling for phylogeny, with and without body size correction. Postcranial and skull + postcranial analyses (but not skull‐only analyses) allowed for a complete segregation of species in morphospace. Morphofunctional factors segregating species included body size, bite force, zeugopodial lengths and osteological features related to parasagittal leg movement. A general gradient of bodily proportions was recovered: lightly built, long‐legged felids with small heads and weak bite forces vs. the opposite. Three loose groups were recognized: small terrestrial felids, mid‐to‐large sized scansorial felids and specialized Acinonyx jubatus and Leptailurus serval. As predicted from a previous study, the assembling of the African felid ensemble during the Plio‐Pleistocene occurred by the arrival of distinct felid lineages that occupied then vacant areas of morphospace, later diversifying in the continent.