Colin R. McHenry

Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa

We provide the first predictions of bite force (BS) in a wide sample of living and fossil mammalian predators. To compare between taxa, we calculated an estimated bite force quotient (BFQ) as the residual of BS regressed on body mass. Estimated BS adjusted for body mass was higher for marsupials than placentals and the Tasmanian devil (Sarcophilus harrisii) had the highest relative BS among extant taxa. The highest overall BS was in two extinct marsupial lions. BFQ in hyaenas were similar to those of related, non-osteophagous taxa challenging the common assumption that osteophagy necessitates extreme jaw muscle forces. High BFQ in living carnivores was associated with greater maximal prey size and hypercarnivory. For fossil taxa anatomically similar to living relatives, BFQ can be directly compared, and high values in the dire wolf (Canis dirus) and thylacine (Thylacinus cynocephalus) suggest that they took relatively large prey. Direct inference may not be appropriate where morphologies depart widely from biomechanical models evident in living predators and must be considered together with evidence from other morphological indicators. Relatively low BFQ values in two extinct carnivores with morphologies not represented among extant species, the sabrecat, Smilodon fatalis, and marsupial sabretooth, Thylacosmilus atrox, support arguments that their killing techniques also differed from extant species and are consistent with ‘canine-shear bite’ and ‘stabbing’ models, respectively. Extremely high BFQ in the marsupial lion, Thylacoleo carnifex, indicates that it filled a large-prey hunting niche.

The production of anatomical teaching resources using three‐dimensional (3D) printing technology

The teaching of anatomy has consistently been the subject of societal controversy, especially in the context of employing cadaveric materials in professional medical and allied health professional training. The reduction in dissection‐based teaching in medical and allied health professional training programs has been in part due to the financial considerations involved in maintaining bequest programs, accessing human cadavers and concerns with health and safety considerations for students and staff exposed to formalin‐containing embalming fluids. This report details how additive manufacturing or three‐dimensional (3D) printing allows the creation of reproductions of prosected human cadaver and other anatomical specimens that obviates many of the above issues. These 3D prints are high resolution, accurate color reproductions of prosections based on data acquired by surface scanning or CT imaging. The application of 3D printing to produce models of negative spaces, contrast CT radiographic data using segmentation software is illustrated. The accuracy of printed specimens is compared with original specimens. This alternative approach to producing anatomically accurate reproductions offers many advantages over plastination as it allows rapid production of multiple copies of any dissected specimen, at any size scale and should be suitable for any teaching facility in any country, thereby avoiding some of the cultural and ethical issues associated with cadaver specimens either in an embalmed or plastinated form. Anat Sci Educ 7: 479–486.

Supermodeled sabercat, predatory behavior in Smilodon fatalis revealed by high-resolution 3D computer simulation

The American sabercat Smilodon fatalis is among the most charismatic of fossil carnivores. Despite broad agreement that its extraordinary anatomy reflects unique hunting techniques, after >150 years of study, many questions remain concerning its predatory behavior. Were the “sabers” used to take down large prey? Were prey killed with an eviscerating bite to the abdomen? Was its bite powerful or weak compared with that of modern big cats? Here we quantitatively assess the sabercats biomechanical performance using the most detailed computer reconstructions yet developed for the vertebrate skull. Our results demonstrate that bite force driven by jaw muscles was relatively weak in S. fatalis, one-third that of a lion (Panthera leo) of comparable size, and its skull was poorly optimized to resist the extrinsic loadings generated by struggling prey. Its skull is better optimized for bites on restrained prey where the bite is augmented by force from the cervical musculature. We conclude that prey were brought to ground and restrained before a killing bite, driven in large part by powerful cervical musculature. Because large prey is easier to restrain if its head is secured, the killing bite was most likely directed to the neck. We suggest that the more powerful jaw muscles of P. leo may be required for extended, asphyxiating bites and that the relatively low bite forces in S. fatalis might reflect its ability to kill large prey more quickly, avoiding the need for prolonged bites.

The craniomandibular mechanics of being human

Diminished bite force has been considered a defining feature of modern Homo sapiens, an interpretation inferred from the application of two-dimensional lever mechanics and the relative gracility of the human masticatory musculature and skull. This conclusion has various implications with regard to the evolution of human feeding behaviour. However, human dental anatomy suggests a capacity to withstand high loads and two-dimensional lever models greatly simplify muscle architecture, yielding less accurate results than three-dimensional modelling using multiple lines of action. Here, to our knowledge, in the most comprehensive three-dimensional finite element analysis performed to date for any taxon, we ask whether the traditional view that the bite of H. sapiens is weak and the skull too gracile to sustain high bite forces is supported. We further introduce a new method for reconstructing incomplete fossil material. Our findings show that the human masticatory apparatus is highly efficient, capable of producing a relatively powerful bite using low muscle forces. Thus, relative to other members of the superfamily Hominoidea, humans can achieve relatively high bite forces, while overall stresses are reduced. Our findings resolve apparently discordant lines of evidence, i.e. the presence of teeth well adapted to sustain high loads within a lightweight cranium and mandible.

Computer simulation of feeding behaviour in the thylacine and dingo as a novel test for convergence and niche overlap

The extinct marsupial thylacine (Thylacinus cynocephalus) and placental grey wolf (Canis lupus) are commonly presented as an iconic example of convergence. However, various analyses suggest distinctly different behaviours and specialization towards either relatively small or large prey in the thylacine, bringing the degree of apparent convergence into question. Here we apply a powerful engineering tool, three-dimensional finite element analysis incorporating multiple material properties for bone, to examine mechanical similarity and niche overlap in the thylacine and the wolf subspecies implicated in its extinction from mainland Australia, Canis lupus dingo. Comparisons of stress distributions not only reveal considerable similarity, but also informative differences. The thylacines mandible performs relatively poorly where only the actions of the jaw muscles are considered, although this must be considered in the light of relatively high bite forces. Stresses are high in the posterior of the thylacines cranium under loads that simulate struggling prey. We conclude that relative prey size may have been comparable where both species acted as solitary predators, but that the dingo is better adapted to withstand the high extrinsic loads likely to accompany social hunting of relatively large prey. It is probable that there was considerable ecological overlap. As a large mammalian hypercarnivore adapted to taking small-medium sized prey, the thylacine may have been particularly vulnerable to disturbance.

High‐Resolution Three‐Dimensional Computer Simulation of Hominid Cranial Mechanics

In vivo data demonstrates that strain is not distributed uniformly on the surface of the primate skull during feeding. However, in vivo studies are unable to identify or track changes in stress and strain throughout the whole structure. Finite element (FE) analysis, a powerful engineering tool long used to predict the performance of man‐made devices, has the capacity to track stress/strain in three dimensions (3‐D) and, despite the time‐consuming nature of model generation, FE has become an increasingly popular analytical device among biomechanists. Here, we apply the finite element method using sophisticated computer models to examine whether 3‐D stress and strain distributions are nonuniform throughout the primate skull, as has been strongly suggested by 2‐D in vivo strain analyses. Our simulations document steep internal stress/strain gradients, using models comprising up to three million tetrahedral finite elements and 3‐D reconstructions of jaw adducting musculature with both cranium and mandible in correct anatomical position. Results are in broad concurrence with the suggestion that few regions of the hominid cranium are clearly optimized for routine feeding and also show that external stress/strain does not necessarily reflect internal distributions. Findings further suggest that the complex heterogeneity of bone in the skull may act to dissipate stress, but that consequently higher strain must be offset by additional strain energy. We hypothesize that, despite energetic costs, this system may lend adaptive advantage through enhancing the organisms ability to modify its behavior before reaching catastrophic failure in bony or dental structures. Anat Rec, 290:1248–1255, 2007.

A central role for venom in predation by Varanus komodoensis (Komodo Dragon) and the extinct giant Varanus (Megalania) priscus

The predatory ecology of Varanus komodoensis (Komodo Dragon) has been a subject of long-standing interest and considerable conjecture. Here, we investigate the roles and potential interplay between cranial mechanics, toxic bacteria, and venom. Our analyses point to the presence of a sophisticated combined-arsenal killing apparatus. We find that the lightweight skull is relatively poorly adapted to generate high bite forces but better adapted to resist high pulling loads. We reject the popular notion regarding toxic bacteria utilization. Instead, we demonstrate that the effects of deep wounds inflicted are potentiated through venom with toxic activities including anticoagulation and shock induction. Anatomical comparisons of V. komodoensis with V. (Megalania) priscus fossils suggest that the closely related extinct giant was the largest venomous animal to have ever lived.

Toward integration of geometric morphometrics and computational biomechanics: New methods for 3D virtual reconstruction and quantitative analysis of Finite Element Models

The ability to warp three-dimensional (3D) meshes from known biological morphology to fit other known, predicted or hypothetical morphologies has a range of potential applications in functional morphology and biomechanics. One of the most challenging of these applications is Finite Element Analysis (FEA), a potentially powerful non-destructive tool in the prediction of mechanical behaviour. Geometric morphometrics is another typically computer-based approach commonly applied in morphological studies that allows for shape differences between specimens to be quantified and analysed. There has been some integration of these two fields in recent years. Although a number of shape warping approaches have been developed previously, none are easily accessible. Here we present an easily accessed method for warping meshes based on freely available software and test the effectiveness of the approach in FEA using the varanoid lizard mandible as a model. We further present new statistical approaches, strain frequency plots and landmark point strains, to analyse FEA results quantitatively and further integrate FEA with geometric morphometrics. Using strain frequency plots, strain field, bending displacements and landmark point strain data we demonstrate that the mechanical behaviour of warped specimens reproduces that of targets without significant error. The influence of including internal cavity morphology in FEA models was also examined and shown to increase bending displacements and strain magnitudes in FE models. The warping approaches presented here will be useful in a range of applications including the generation and analysis of virtual reconstructions, generic models that approximate species means, hypothetical morphologies and evolutionary intermediaries.

Cranial performance in the Komodo dragon (Varanus komodoensis) as revealed by high-resolution 3-D finite element analysis

The Komodo dragon (Varanus komodoensis) displays a unique hold and pull‐feeding technique. Its delicate ‘space‐frame’ skull morphology differs greatly from that apparent in most living large prey specialists and is suggestive of a high degree of optimization, wherein use of materials is minimized. Here, using high‐resolution finite element modelling based on dissection and in vivo bite and pull data, we present results detailing the mechanical performance of the giant lizards skull. Unlike most modern predators, V. komodoensis applies minimal input from the jaw muscles when butchering prey. Instead it uses series of actions controlled by postcranial muscles. A particularly interesting feature of the performance of the skull is that it reveals considerably lower overall stress when these additional extrinsic forces are added to those of the jaw adductors. This remarkable reduction in stress in response to additional force is facilitated by both internal and external bone anatomy. Functional correlations obtained from these analyses also provide a solid basis for the interpretation of feeding ecology in extinct species, including dinosaurs and sabre‐tooth cats, with which V. komodoensis shares various cranial and dental characteristics.

Effects of Gape and Tooth Position on Bite Force and Skull Stress in the Dingo (Canis lupus dingo) Using a 3-Dimensional Finite Element Approach

Models of the mammalian jaw have predicted that bite force is intimately linked to jaw gape and to tooth position. Despite widespread use, few empirical studies have provided evidence to validate these models in non-human mammals and none have considered the influence of gape angle on the distribution of stress. Here using a multi-property finite element (FE) model of Canis lupus dingo, we examined the influence of gape angle and bite point on both bite force and cranial stress. Bite force data in relation to jaw gape and along the tooth row, are in broad agreement with previously reported results. However stress data showed that the skull of C. l. dingo is mechanically suited to withstand stresses at wide gapes; a result that agreed well with previously held views regarding carnivoran evolution. Stress data, combined with bite force information, suggested that there is an optimal bite angle of between 25 degrees and 35 degrees in C. l. dingo. The function of these rather small bite angles remains unclear.