A new species of Prosansanosmilus: implications for the systematic relationships of the family Barbourofelidae new rank (Carnivora, Mammalia)

A new barbourofelid species, Prosansanosmilus eggeri , is described from the Middle Miocene (MN 5) locality of Sandelzhausen, Germany. It differs from all other European barbourofelid species in being smaller and showing a more plesiomorphic morphology, especially in the relatively less developed sabretooth adaptations, low accessory cusps on the premolars, and the remnant of a very small talonid on the carnassial. The species is, however, stratigraphically later than the more apomorphic P. peregrinus, which is known from MN 4 of Germany and France. A phylogenetic analysis based on dental characters of early nimravids, barbourofelids and felids supports previous results on skull morphology of Barbourofelis that Barbourofelinae is not closely related to the Late Eocene and Oligocene Nimravinae. Instead, both subfamilies should be treated as separate families, with the Barbourofelidae closely related to the Felidae. The Barbourofelidae differ from the Felidae as well as from the Nimravidae s.s ., particularly in the unique morphology of their basicranium. They presumably originated in Africa; P. eggeri sp. nov. is interpreted as part of a Miocene immigration of African faunal elements into Europe that took place at the beginning of MN 5. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 43−61.     

Feeding behaviour and bite force of sabretoothed predators

The feeding behaviour of extinct sabretoothed predators (machaeroidines, nimravids, barbourofelids, machairodonts and thylacosmilines) is investigated using beam theory. Because bite force applied along the mandible should be proportional to the external dimension of the mandibular corpus, patterns of variation in these dimensions at interdental gaps will reflect the adaptation of the jaw to specific loads, related to killing methods. Comparison of the mandibular force profiles of sabretooths to those of extant conical‐toothed carnivorans of known feeding behaviour reveals that sabretooths had a powerful bite, as strong or stronger than extant felids of similar mandibular length. Loads exerted at the lower canine were better constrained in the sagittal plane than in extant conical‐toothed carnivorans, indicating that prey was efficiently restrained when the sabre bite was delivered. The mandibular symphysis is generally better buttressed dorsoventrally in dirk‐toothed sabretooths than in scimitar‐toothed sabretooths, implying different killing strategies for the two ecomorphs: dirktooths delivered powerful sabre bites on prey they restrained with their forelimbs, while scimitartooths delivered slashing sabre bites and may have used their incisor battery to subdue their prey. The mandibular symphysis of Smilodon fatalis is less buttressed dorsoventrally than that of other dirk‐toothed sabretooths, possibly as a consequence of the greater torsional stresses induced while feeding rapidly on carcasses in response to intense competition. The mandibular symphysis of Thylacosmilus atrox is better buttressed dorsoventrally in juveniles than in adults, suggesting that young marsupial sabretooths underwent an extended period of parental care as typically observed in modern felids and inferred for eutherian sabretooths. Finally, machaeroidines and the nimravid Nimravus brachyops are exceptional in exhibiting a degree of dorsoventral buttressing of the mandibular symphysis that is intermediate between advanced sabretooths and conical‐toothed felids but similar to the extant Neofelis nebulosa, suggesting that the latter taxon may be close to the ancestral condition of a new sabretooth radiation. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 145 , 393–426.     

A primitive nimravine skull from the Quercy fissures, France: implications for the origin and evolution of Nimravidae (Carnivora)

A carnivore skull from the Phosphorites of Quercy, France, is described herein. Its well-preserved basicranial structure, cat-like morphology, and dental reduction support the assignment of this specimen to the Nimravidae. This is the most primitive nimravid described and its overall morphology is the closest yet to the hypothetical ancestor of the Nimravinae as previously conceived. In addition, the monophyly of the Nimravinae is reinforced by the similar basicranium of the specimen described here compared to other nimravid genera. It suggests that this typical basicranial structure appeared well ahead the sabretooth features in this group. The early evolution and diversification of the Nimravinae occurred during the Late Eocene in the northern hemisphere but the precise geographic area is still unknown. A rapid evolution during the initial radiation of the group or an incomplete fossil record in Eurasia could both equally explain the rapid and worldwide distribution of the Nimravinae.     

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.     

Functional anatomy of the cervical region in the late Miocene amphicyonid Magericyon anceps (Carnivora,Amphicyonidae): implications for its feeding behaviour

We describe the skull and neck morphology of the late Miocene amphicyonid Magericyon anceps, focusing on aspects related to functional anatomy. This species, recorded only from the Vallesian sites of Batallones‐1 and Batallones‐3 (Madrid, Spain), is the last known amphicyonid in the fossil record of Western Europe, with the Batallones populations being one of the best‐known of the family. The morphology of its skull and cervical vertebrae allows us to infer aspects of its associated musculature, such as muscle strength and range of movement. Magericyon anceps had well‐developed neck muscles, suited for providing the head with a high capacity for lateral and rotatory movements, as well as for playing an important role in the extension and stabilization of the head and neck, improving its efficiency in killing and consuming prey. Magericyon anceps shared its habitat with other large carnivorans, which would have strongly influenced its behaviour. Rapid killing and processing of prey would have been an advantage for avoiding kleptoparasitism by other large predators, as well as reducing consumption time, during which M. anceps would have been more vulnerable to attack from competitors.     

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.     

Ecomorphology of the giant bear-dogs Amphicyon and Ischyrocyon

Giant bear-dogs of the genera Amphicyon and Ischyrocyon (Carnivora, Amphicyonidae, Amphicyoninae) were the largest carnivorans in North America during middle and late Miocene (17.5–8.8 Mya) with a dental and skeletal morphology that combined features found in living Ursidae, Canidae, and Felidae. This study tests previously proposed models of diet and hunting behaviour of these extinct carnivorans. Relative grinding area (RGA) of lower molars and wear pattern on upper molars suggest that bear-dogs were carnivorous. Amphicyon and Ischyrocyon possessed skeletal features of both ambush (short distal limb segments) and pursuit (caudally bent olecranon process of ulna) living predators. Therefore, bear-dogs probably pursued their prey (mediportal ungulates) for a longer distance but at a slower speed than do living ambush predators. Upon catching up to its prey a bear-dog probably seized it with powerfully muscled forelimbs and killed it by tearing into its ribcage or neck with canines set in a narrow rostrum.     

Ontogenetic shift in susceptibility to predators in juvenile northern abalone, Haliotis kamtschatkana

Predation has been suggested as a major cause of juvenile mortality in benthic marine invertebrates. However, the extent to which juveniles are susceptible to predators is unknown for most species, and it remains unclear to what extent ontogenetic shifts in susceptibility to predators are common among marine invertebrates. This study examined the northern abalone Haliotis kamtschatkana, a species listed as threatened in British Columbia, Canada. Our goals were to characterize the diversity and abundance of species that prey on juvenile abalone and determine if abalone experience an ontogenetic shift in susceptibility to predators. Juvenile H. kamtschatkana were found to be susceptible to a broad variety of predators: 14 of the 37 potential predator species to which we offered juvenile abalone (≤ 28 mm shell length (SL)) consumed at least one juvenile abalone. Four of those species (three crabs and one seastar) consumed ≥ 10% of the juvenile abalone that were offered in the laboratory. These species were present at field sites where abalone are found, indicating that they have the potential to be significant predators of juvenile H. kamtschatkana in the wild. The most abundant predators were small crabs, especially Lophopanopeus bellus (black-clawed crabs) and Scyra acutifrons (sharp-nosed crabs). Juvenile H. kamtschatkana also experienced a pronounced ontogenetic shift in susceptibility to predators. The risk of predation for juvenile H. kamtschatkana decreased rapidly with increasing body size, especially over the 12–13 mm SL size range. Susceptibility remained low beyond 13 mm SL, indicating relatively low and unchanging levels of predation risk once the individual reaches this size. Although abalone are susceptible to several species during the first 1–2 years of life, predator effects on juvenile abalone abundance and microhabitat use may largely be attributable to the influence of only 1 or 2 predator species that can only kill abalone < 13 mm SL.     

A juvenile skull of Dysalotosaurus lettowvorbecki (Ornithischia: Iguanodontia), and implications for cranial ontogeny,phylogeny, and taxonomy in ornithopod dinosaurs

A juvenile skull of Dysalotosaurus lettowvorbecki, a basal iguanodontian from the Middle Dinosaur Member (Kimmeridgian) of the Tendaguru Beds (Tanzania), is described and reconstructed in detail. Further preparation and computed tomography scans have uncovered additional and formerly unknown skull elements, especially of the lower jaw. The reconstruction of the skull reveals ontogenetic differences to earlier reconstructions, which were based on specimens of relatively older individuals. The most notable ontogenetic changes in Dysalotosaurus are a relative lengthening of the muzzle, a decrease in the relative size of the orbit, and a straightening of the posterior skull roof. Additionally, ontogenetic variations were found in many single elements of the skull, all of which reflect the three main tendencies described above. Furthermore, there might have been an ontogenetic change in the diet, from omnivorous juveniles to fully herbivorous adults. The results of this study will help to evaluate ontogenetic stages in other ornithopods, and will shed light on a crucial stage of ornithopod evolution that culminated in the highly specialized and diverse hadrosaurs of the Cretaceous. Many of the evolutionary changes seen in this lineage can be attributed to peramorphism, and probably reflect a perfection of the adaptation towards an obligatory herbivorous diet. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 366–396.     

Aspects of the functional morphology in the cranial and cervical skeleton of the sabre-toothed cat Paramachairodus ogygia (Kaup, 1832) (Felidae, Machairodontinae) from the Late Miocene of Spain: implications for the origins of the machairodont killing bite

The skull and cervical anatomy of the sabre-toothed felid Paramachairodus ogygia (Kaup, 1832) is described in this paper, with special attention paid to its functional morphology. Because of the scarcity of fossil remains, the anatomy of this felid has been very poorly known. However, the recently discovered Miocene carnivore trap of Batallones-1, near Madrid, Spain, has yielded almost complete skeletons of this animal, which is now one of the best known machairodontines. Consequently, the machairodont adaptations of this primitive sabre-toothed felid can be assessed for the first time. Some characters, such as the morphology of the mastoid area, reveal an intermediate state between that of felines and machairodontines, while others, such as the flattened upper canines and verticalized mandibular symphysis, show clear machairodont affinities.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 363−377.     

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.     

Intraspecific versus interspecific variation in Miocene Great Basin mylagaulids: implications for systematics and evolutionary history

The Mylagaulidae are a family of burrowing rodents abundant in Miocene faunas from western North America. Recent taxonomic revisions of mylagaulids from the Great Plains suggest that their systematics may be best understood on a regional basis. Previous studies addressed the taxonomy and evolutionary history of mylagaulids from the Great Basin, but recent discoveries of specimens, new phylogenetic data, and more detailed stratigraphical information necessitate a thorough reanalysis of their relationships and occurrences. We present a revision of the systematics of the mylagaulids from the Great Basin. In addition to rare large mylagaulids of uncertain taxonomic affinity, we recognize four species of mylagaulids distributed throughout Oregon and Nevada from the late Hemingfordian through to the early late Hemphillian: Alphagaulus vetus, Hesperogaulus gazini, Hesperogaulus wilsoni, and a new species from the genus Hesperogaulus. All species are known from large sample sizes of isolated premolars, allowing consideration of ontogenetic variation in determining the key morphological differences that allow recognition of different species. Although the number of enamel lakes varies within a given taxon, the presence of some of these lakes is taxonomically significant. This result emphasizes the importance of understanding ontogeny in describing species of fossil hypsodont mammals. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 427–450.     

Influence of body mass on the shape of forelimb in musteloid carnivorans

In the majority of mammals, the limbs are positioned under the body and play an important role in gravitational support, allowing the transfer of the load and providing stability to the animal. For this reason, an animal's body mass likely has a significant effect on the shape of its limb bones. In the present study, we investigate the influence of body mass variation on the shape of the three long bones of the forelimb in a group of closely‐related species of mammals: the musteloid carnivorans. We use geometric morphometric techniques to quantify forelimb shape; then estimate phylogenetic signal in the shape of each long bone; and, finally, we apply an independent contrasts approach to assess evolutionary associations between forelimb shape and body mass. The results obtained show that body mass evolution is tightly coordinated with the evolution of forelimb shape, although not equally in all elements. In particular, the humeral and radial shapes of heavier species appear better suited for load bearing and load transmission than the ulna. Nevertheless, our results also show that body mass influences only part of forelimb long bone shape and that other factors, such as locomotor ecology, must be considered to fully understand forelimb evolution. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 91–103.     

Ontogenetic Evidence for Dental Homologies and Premolar Replacement in Fossil and Extant Caenolestids (Marsupialia)

It is generally accepted that the South American marsupial family Caenolestidae is characterized in part by the absence or noneruption of the third deciduous premolar (dP3) in both jaws, although juvenile stages have rarely been identified in extant or fossil representatives of the family. Published illustrations of the dentary of the Miocene caenolestid Stilotherium suggested to us, however, that P3 erupted relatively late during ontogeny, after the eruption of M4. In extant marsupials, this eruption sequence appears to represent the plesiomorphic state and this pattern is generally associated with the eruption of dP3 earlier in ontogeny, and its subsequent replacement by the erupting P3. Therefore, we suspected that a dP3 erupted in earlier ontogenetic stages of caenolestids; to test this hypothesis we searched the mammalogy collections of three museums for evidence of dP3 in juvenile specimens of caenolestids. Examination of more than 180 specimens of the three extant genera of caenolestid marsupials resulted in the identification of only nine juvenile or subadult stages of dental eruption. Seven specimens of Caenolestes and Rhyncholestes corroborated our hypotheses of late eruption of P3 in Caenolestidae. In addition, the two youngest specimens of Caenolestes possessed a tiny, rudimentary dP3, measuring about 0.4 to 0.5 mm in greatest length, associated with a mesiolingual eruption pit containing the apex of the larger P3 in very early phases of eruption above the alveolar margins. The tiny dP3 is clearly nonfunctional in occlusion, and it is questionable whether it erupted above the gun margins in life. Comparison of the dentaries of subadult caenolestids with four dentaries of the Miocene genus Stilotherium corroborated our initial impression that the fossil genus exhibited evidence of a late-erupting P3, comparable to the condition in extant caenolestids. We suggest that examination of other specimens of juvenile dentitions, skulls, and skeletons in museum collections can provide additional insight into the developmental and evolutionary biology of mammals.     

Does intraspecific niche partitioning in a native predator influence its response to an invasion by a toxic prey species?

Abstract The introduced and highly toxic cane toad (Bufo marinus) is rapidly spreading across northern Australia where it may affect populations of large terrestrial vertebrate predators. The ecological impact of cane toads will depend upon the diets, foraging modes and habitat use of native predators, and their feeding responses to cane toads. However, intraspecific niche partitioning may influence the degree of vulnerability of predators to toxic prey, as well as the time course of the impact of alien invaders on native species. We studied the diet of the northern death adder Acanthophis praelongus and their feeding responses to cane toads. In the laboratory, death adders from all size classes and sexes readily consumed frogs and cane toads. Diets of free ranging A. praelongus from the Adelaide River floodplain were more heterogeneous. Juvenile snakes ate mainly frogs (39% of prey items) and small scincid lizards (43%). Both sexes displayed an ontogenetic dietary shift from lizards to mammals, but adult males fed on frogs (49%) and mammals (39%) whereas adult females (which grew larger than males) fed mainly on mammals (91%) and occasionally, frogs (9%). Feeding rates and body condition of adult snakes varied temporally and tracked fluctuations in prey availability. These results suggest that cane toads may negatively affect populations of northern death adders in the Darwin region. However, we predict that different size and sex classes of A. praelongus will experience differential mortality rates over different timescales. The initial invasion of large toads may affect adult males, but juveniles may be unaffected until juvenile toads appear the following year, and major affects on adult female death adders may be delayed until annual rainfall fluctuations reduce the availability of alternative (rodent) prey.     

Patterns of growth of the mandibular corpus in spotted hyenas (Crocuta crocuta) and cougars (Puma concolor)

Differences in jaw morphology among adult carnivorans are well established, but the ontogenetic mechanisms by which these differences arise are largely unexplored. Mandibular ontogeny in Crocuta crocuta and Puma concolor is analysed biomechanically using principles of beam theory. In each species, the development of cross-sectional properties of the mandibular corpus associated with rigidity under loading follows a biphasic pattern of growth. In early postnatal growth, deposition of cortical bone appears to be constrained by the overall weaker tissue with which juvenile skeletons are constructed and by the need to volumelrically accommodate the developing teeth within their bony crypts. Thus, this stage of growth is characterized by a net periosteal deposition of bone and a swelling of the medullary cavity. In late postnatal growth, the constraints on endosteal deposition of bone are relieved as the permanent teeth erupt; thus, cortical thicknesses increase sharply by periosteal expansion as well as medullary contraction. Finally, it is noted that basic differences in jaw construction between Crocuta and Puma appear to develop prenatally as they are largely in place at birth. Hence, postnatal development enhances, but does not soley contribute to, the biomechanical differences in the jaws of these species.     

Petrosal and inner ear anatomy and allometry amongst specimens referred to Litopterna (Placentalia)

New isolated petrosals from the Itaboraí beds of Brazil (late Palaeocene or early Eocene) are here described and referred to the early diverging litoptern Miguelsoria parayirunhor, based on phylogenetic, size, and abundance arguments. Both the external and internal anatomy of these specimens were investigated, which for the first time document many details of the auditory region of a Palaeogene litoptern. Our cladistic analysis, which included our new observations, failed to recover a monophyletic Litopterna but did not exclude it. A constrained analysis for the monophyly of this order showed that several features such as a (sub)quadrangular and anteroposteriorly elongated tensor tympani fossa and a large notch in the vicinity of the external opening of the cochlear canaliculus may constitute synapomorphies for Litopterna. The evolution of several other auditory characters amongst Litopterna is discussed and the relative dimensions of the inner ear and surrounding petrosal in the group were also investigated. This allowed detection of negative allometry of the bony labyrinth within the petrosal, which was confirmed by measurements and regression analysis across a larger sample of placental mammals. This scaling effect probably has an important influence on several characters of the bony labyrinth and petrosal, amongst which are the length of the vestibular aqueduct and cochlear canaliculus. It demonstrates that many aspects of the morphological variation of the bony labyrinth need to be thoroughly investigated before being incorporated into phylogenetic analyses. © 2015 The Linnean Society of London     

The anatomy of the basal ornithischian dinosaur Eocursor parvus from the lower Elliot Formation (Late Triassic) of South Africa

Ornithischia is a morphologically and taxonomically diverse clade of dinosaurs that originated during the Late Triassic and were the dominant large‐bodied herbivores in many Cretaceous ecosystems. The early evolution of ornithischian dinosaurs is poorly understood, as a result in part of a paucity of fossil specimens, particularly during the Triassic. The most complete Triassic ornithischian dinosaur yet discovered is Eocursor parvus from the lower Elliot Formation (Late Triassic: Norian–Rhaetian) of Free State, South Africa, represented by a partial skull and relatively complete postcranial skeleton. Here, the anatomy of Eocursor is described in detail for the first time, and detailed comparisons are provided to other basal ornithischian taxa. Eocursor is a small‐bodied taxon (approximately 1 m in length) that possesses a plesiomorphic dentition consisting of unworn leaf‐shaped crowns, a proportionally large manus with similarities to heterodontosaurids, a pelvis that contains an intriguing mix of plesiomorphic and derived character states, and elongate distal hindlimbs suggesting well‐developed cursorial ability. The ontogenetic status of the holotype material is uncertain. Eocursor may represent the sister taxon to Genasauria, the clade that includes most of ornithischian diversity, although this phylogenetic position is partially dependent upon the uncertain phylogenetic position of the enigmatic and controversial clade Heterodontosauridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 648–684.     

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.