Jeffrey J. Thomason

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.

Biomechanical and Mechanical Investigations of the Hoof-Track Interface in Racing Horses

The aim of this article is to review current knowledge of kinetic variables of the hoof-track interaction and track properties relevant to the objective of minimizing injuries to horses at racing tracks. In each phase of the stance--primary impact, secondary impact, support, and breakover, the hoof experiences different combinations of force and acceleration. The role of each combination, and of measured track properties, in causing catastrophic and chronic injuries to the limbs of racing horses is unknown. Limited data of this type have been provided in previous epidemiologic studies of risk factors for breakdown. Future epidemiological studies should include characterization of the track surfaces and a more complete description of the kinematics of the hoof and surface. Consideration of an appropriate range of physical properties is necessary in track design, testing, and maintenance.

Cranial dimensions and forces of biting in the domestic dog

The purpose of this paper is to analyse the effects of cranial size and shape in domestic dogs (Canis familiaris) on predicted forces of biting. In addition to continuous size‐shape analysis, nine size‐shape groups were developed based on three skull shape categories and three skull size categories. Bite forces were predicted from measurements made on dried skulls using two lever models of the skull, as well as simple models derived by regression analysis. Observed bite force values were not available for the database used in this study, so only comparisons between categories and models were undertaken. The effects of shape and size on scaled predicted bite forces were evaluated. Results show that bite force increases as size increases, and this effect was highly significant (P < 0.0001). The effect of skull shape on bite force was significant in medium and large dogs (P < 0.05). Significant differences were not evident in small dogs. Size × shape interactions were also significant (P < 0.05). Bite force predictions by the two lever models were relatively close to each other, whereas the regression models diverged slightly with some negative numbers for very small dogs. The lever models may thus be more robust across a wider range of skull size‐shapes. Results obtained here would be useful to the pet food industry for food product development, as well as to paleontologists interested in methods of estimating bite force from dry skulls.

Calibration of estimated biting forces in domestic canids: comparison of post-mortem and in vivo measurements

Estimates of biting forces are widely used in paleontological and comparative studies of feeding mechanics and performance, and are usually derived from lever models based on measurements made on the skull that are relevant to the mechanics of the masticatory system. Owing to assumptions and unmeasurable errors in their estimation, such values are used comparatively rather than as absolute estimates. The purpose of this paper was to provide calibration of post‐mortem calculated bite force estimates by comparing them to in vivo forces derived from a sample of 20 domestic dogs (Canis familiaris) during muscle stimulation under general anaesthesia. Two lever models previously described in the literature were used to estimate post‐mortem values, and regression analysis was also performed to derive best‐fit equations against a number of morphometric measurements on the skull. The ranges of observed forces in vivo were 147–946 N at the canine, and 524–3417 N at the second molar. The lever models substantially underestimated these forces, giving mean values between 39% and 61% of the observed means. Predictability was considerably improved by removing the linear bias and deviation of the regression slope from unity with an adjustment equation. Best‐fit statistical models developed on these animals performed considerably better (calculated means within 0.54% of observed means) and included easily measureable variables such as bodyweight, dimensions of the temporalis fossa and out‐lever from the jaw joint to the biting tooth. These data should lead to more accurate absolute, rather than relative, estimates of biting forces for other extant and fossil canids, and other carnivorans by extrapolation.

In vivo surface strain and stereology of the frontal and maxillary bones of sheep: implications for the structural design of the mammalian skull.

Does the skull of the sheep behave as a tube or as a complex of independent bones linked by sutures? Is the architecture within cranial bones optimized to local strain alignment? We attempted to answer these questions for the sheep by recording from rosette strain gauges on each frontal and maxillary bone and from single‐axis gauges on each dentary of five sheep while they fed on hay. Bone structure was assessed at each rosette gauge site by stereological analysis of high‐resolution radiographs. Structural and strain orientations were tested for statistical agreement. Ranges of strain magnitudes were ±1200 μϵ on the mandible, ±650 μϵ on the frontals, and ±400 μϵ on the maxillae. Each gauge site experienced one strain signal when on the working (chewing) side and a different one when on the balancing (nonchewing) side. The two signals differed in mode, magnitude, and orientation. For example, on the working side, maxillary gauges were under mean compressive strains of –132 μϵ (S.D., 73.3 μϵ), oriented rostroventrally at 25°–70° to the long axis of the skull. On the balancing side, the same gauges were under mean tensile strains of +319 μϵ (S.D., 193.9 μϵ), at greater than 65° to the cranial axis. Strain patterns on the frontals are consistent with torsion and bending of the whole skull, indicating some degree of tube‐like mechanical behavior. Frontal and maxillary strains also showed a degree of individual loading, resulting from modulation of strains across sutures and local effects of muscle activity. The sheep skull seems to behave as a tube made of a complex of independent bones. Structural orientation was in statistically significant agreement with the orientation of working‐side compressive principal strain ϵ2, even though principal tensile strains may be as large or larger. Cranial bone architecture in sheep is not optimized to both strain signals it experiences. Anat Rec 264:325–338, 2001.

Shape, Orientation and Spacing of the Primary Epidermal Laminae in the Hooves of Neonatal and Adult Horses (Equus caballus)

Circumferential and proximodistal variations in the morphology of the primary epidermal laminae of six neonatal and five adult equine feet were documented. Three parameters were quantified: interlaminar spacing, the orientation of the laminae with respect to the overlying wall, and any angulation within the laminae themselves (‘internal angle’). In adult feet, the laminae were most closely spaced at the dorsum, the spacing increasing gradually towards the heels. In foals there was a non-significant trend for the dorsal laminae to be more widely spaced than those in more caudal parts of the foot. In both age groups, the dorsal laminae were almost straight (mean divergence from linearity at all sites 2°), and were oriented at approximately 90° to the tangent to the overlying wall (mean orientation for all sites 91°). At the quarters, the laminae were in general oriented caudally relative to the tangential position from their epidermal to their dermal ends (mean orientation of >90° at 12 of 16 sampling sites, where an orientation of >90° defines a ‘caudally directed’ orientation) and, in general, had a bend within their length (mean absolute value of internal angle for all sites 9°). At the heels there was greater variability in the data for both laminar orientation and internal angle. Overall, the foal feet showed greater mediolateral symmetry and less proximodistal variation than did the adult feet. In both age groups, rapid spatial changes in laminar morphology were closely associated with the position of the margins of the third phalanx.

Analysis of strain and stress in the equine hoof capsule using finite element methods: comparison with principal strains recorded in vivo

Finite-element (FE) methods have great potential in equine biomechanics in evaluating mechanical stresses and strains in tissues deep within the hoof. In this study, we critically assessed that potential by comparing results of FE analyses of capsular strain with in vivo data. Nine FE models were developed, corresponding to the shape of hooves for which in vivo principal strain data are available. Each model had the wall, laminar junction, sole and distal phalanx (PIII). In a first loading condition (LC1), force is distributed uniformly to the bearing surface of the wall to determine reaction forces and moment on PIII. These reaction forces were subsequently applied to PIII in loading condition 2 (LC2) to simulate loading via the skeleton. Magnitude of the force resultant was equivalent to the vertical force on the hoof at midstance. Principal compressive strains epsilon2 were calculated at the locations of 5 rosette gauges on the real hooves and are compared with the in vivo strains at midstance. FE strains were from 16 to 221% of comparable in vivo values, averaging 104%. All models in this, and reports by other workers, show predominance of stress and strain at the toe to a greater extent than in the real hoof. The primary conclusion is that FE modelling of strain in the hoof capsule or deeper tissues of individual horses should not be attempted without corroborating experimental data.

Morphology of the Laminar Junction in Relation to the Shape of the Hoof Capsule and Distal Phalanx in Adult Horses (Equus caballus)

The purpose was to investigate whether differences in equine hoof shape, which are inferred to alter foot function, are accompanied by differences in morphology of the laminar junction. Ten fore feet from adult horses were segregated into normal and low-angle groups, depending on the dorsal angle of the hoof wall. Twenty measurements of external hoof shape and four of the enclosed distal phalanx were tested for differences between groups, and for intragroup correlations. Three measurements of laminar morphology (spacing, orientation and degree of bend) were recorded for samples of up to 50 primary epidermal laminae at each of 20 sample sites. Sites were distributed over the foot in 5 circumferential columns and 4 proximodistal rows. Intergroup differences were investigated, as were correlations among sample sites of the laminar variables with the shape measurements. Results show differences in hoof shape between groups (but not bone shape) and laminar morphology. Six shape measurements are significantly different between groups: dorsal angle, medial and lateral angles, lateral sole width, solar circumference, and dorsal length. In the normal group, shape measurements show patterns of correlation among regions of the hoof, and between hoof and bone measurements. In the low-angle group, shape correlations occur largely within one region of the hoof (the heels) and in the bone measurements. Laminar spacing tends to be nonsignificantly greater in the low-angle group, while variances for laminar spacing and orientation are significantly greater in this group. Laminar spacing correlates with bone width and coronary circumference (CC) of the hoof in the normal group, but only with CC in the low-angle group. When taken as a whole, and interpreted in light of a model of foot mechanical function, the results appear to indicate a deterioration in structural coherence of the foot in the low-angle group.

Changes in growth of the hoof wall and hoof morphology in response to regular periods of trotting exercise in Standardbreds

OBJECTIVE To quantitate changes in hoof wall growth and hoof morphology induced by mild exercise in Standardbreds. ANIMALS 18 Standardbreds. PROCEDURES Horses were exercised at approximately 6 m/s (4,200 to 5,600 m/d) on 4 d/wk for 17 weeks. Both exercise (n = 9) and nonexercise (control group; 9) groups were housed in a large paddock throughout the study. At the beginning and end of the study, right forelimb feet of all horses were digitally photographed and underwent magnetic resonance imaging. Hoof wall measurements were obtained from the images to evaluate hoof wall growth and morphometric variables. Data were compared between the groups and within each group via a quadratic model. Changes in each variable and pairwise correlations between variables were evaluated. RESULTS Morphometric variables did not significantly differ between the control and exercise groups. However, differences within each group between the start and the end of the study were significant for several variables; overall, values for hoof wall variables increased and those for solar variables decreased. Between the beginning and the end of the study, the amount of variation in values of hoof capsule variables in the exercise group decreased to a greater extent, compared with control group findings. Patterns of pairwise correlations for variables differed between the groups. CONCLUSIONS AND CLINICAL RELEVANCE In Standardbreds, mild exercise for 17 weeks caused no significant changes in hoof wall growth or morphometric variables. Subtle changes may develop in equine hooves in response to loading, and mild exercise may not be a strong adaptive stimulus.

Effects of hoof shape, body mass and velocity on surface strain in the wall of the unshod forehoof of Standardbreds trotting on a treadmill

The purpose of this work is to investigate the effects of body mass (BM), velocity ( V ), and hoof shape on compressive surface strains in the wall of the front hoof at the trot. Toe angle (TA), heel angle (HA), toe length (TL), medial and lateral wall length (MWL, LWL) and BM were measured for nine adult, unshod Standardbreds. Five rosette gauges were glued around the circumference of the left forehoof of each animal which was then trotted on a treadmill at a set range of velocities from 3.5 to 7.5 m s −1 . Analysis of variance (ANOVA) of principal compressive strains ɛ 2 at midstance identified that all primary variables (BM, V , TA, HA, etc.) had a significant effect as did the interactions of TA×HA and BM×TA. These significant variables explained over 96% of the variation in ɛ 2 . Multiple regression of ɛ 2 on these variables gave equations which accurately predicted ɛ 2 within 3%, but the individual coefficients did not accurately describe how each variable affected ɛ 2 . Further tests using bivariate regression gave equations that enabled ɛ 2 data to be standardized for BM and V at the gauge locations used here. Strain ɛ 2 increased linearly with mass and curvilinearly with velocity ( ɛ 2 ∝ V + V 2 ), and both caused redistribution of strain to the dorsum and lateral quarter. Variation in each shape variable caused redistribution rather than simple increase or decrease in strains. The primary conclusion with regard to hoof shape is that the effects of change in any one measurement on strain magnitudes are affected by the values of all other measurements. Resolving the interplay among measurements in their effects on ɛ 2 will need a considerably larger sample size than that used here.