Search

Synopsis Recent advances in non-invasive imaging methods have revitalized the field of comparative anatomy, and reproductive anatomy has been no exception. The reproductive systems of female amniotes present specific challenges, namely their often internal “hidden” anatomy. Quantifying female reproductive systems is crucial to recognizing reproductive pathologies, monitoring menstrual cycles, and understanding copulatory mechanics. Here, we conduct a review of the application of non-invasive imaging techniques to female amniote reproductive anatomy. We introduce the commonly used imaging modalities of computed tomography (CT) and magnetic resonance imaging (MRI), highlighting their advantages and limitations when applied to female reproductive tissues, and make suggestions for future advances. We also include a case study of micro CT and MRI, along with their associated staining protocols, applied to cadavers of female adult stoats (Mustela erminea). In doing so, we will progress the discussion surrounding the imaging of female reproductive anatomy, whilst also impacting the fields of sexual selection research and comparative anatomy more broadly.

This paper presents the results of SHREC’21 track: Quantifying Shape Complexity. Our goal is to investigate how good the submitted shape complexity measures are (i.e. with respect to ground truth) and investigate the relationships between these complexity measures (i.e. with respect to correlations). The dataset consists of three collections: 1800 perturbed cube and sphere models classified into 4 categories, 50 shapes inspired from the fields of architecture and design classified into 2 categories, and the data from the Princeton Segmentation Benchmark, which consists of 19 natural object categories. We evaluate the performances of the methods by computing Kendall rank correlation coefficients both between the orders produced by each complexity measure and the ground truth and between the pair of orders produced by each pair of complexity measures. Our work, being a quantitative and reproducible analysis with justified ground truths, presents an improved means and methodology for the evaluation of shape complexity.

Amblypygi is an arachnid order possessing a unique pair of spined pedipalps: appendages that perform in prey capture, courtship, and contest. Pedipalp length, hypothesized to be under sexual selection, varies markedly across amblypygid species, and pedipalp spination, thought to reflect selection for function in prey capture, also differs interspecifically. Differences in pedipalp shape between species may indicate that the relative strength of selection for prey capture and sexual selection vary across the group. However, interspecific differences in pedipalp shape have not been quantified, due to difficulties in identifying homologous features. For the first time, we quantify trends in amblypygid pedipalp shape complexity. We use elliptical Fourier analysis to quantify 2D complexity in pedipalp outlines across eleven species and six genera. We find that complexity significantly decreases as pedipalp length increases. This appears to be driven by relative spine length, suggesting that a trade‐off exists between pedipalp length and spination. Furthermore, significant female‐biased sexual dimorphism in shape complexity is present in the tibial segment of the amblypygid pedipalp. Our results provide novel insights into the drivers of amblypygid pedipalp evolution and suggest that a functional trade‐off between performance in prey capture and other functions under sexual selection exist in this enigmatic structure., Amblypygids are an arachnid order possessing a unique pair of spined pedipalps: appendages that perform in prey capture, courtship, and contest. We quantify trends in amblypygid pedipalp shape complexity using elliptical Fourier analysis applied to pedipalp outlines. Our results provide novel insights into the drivers of amblypygid pedipalp evolution and suggest that a functional trade‐off between performance in prey capture and other functions under sexual selection exist in this enigmatic structure.

Amblypygids are an arachnid order possessing a unique pair of spined pedipalps: appendages that perform in prey capture, courtship and contest. Pedipalp length, hypothesised to be under sexual selection, varies markedly across amblypygid species, and pedipalp spination, thought to reflect selection for function in prey capture, also differs interspecifically. Differences in pedipalp shape between species may indicate that the relative strength of selection for prey capture and sexual selection vary across the group. However, interspecific differences in pedipalp shape have not been quantified, due to difficulties in identifying homologous features. For the first time, we quantify trends in amblypygid pedipalp shape complexity. We use elliptical Fourier analysis to quantify 2D complexity in pedipalp outlines across eleven species and six genera. We find that complexity significantly decreases as pedipalp length increases. This appears to be driven by relative spine length, suggesting that a trade-off exists between pedipalp length and spination. Furthermore, significant female-biased sexual dimorphism in shape complexity is present in the tibial segment of the amblypygid pedipalp. Our results provide novel insights into the drivers of amblypygid pedipalp evolution, and suggest that a functional trade-off between performance in prey capture and other functions under sexual selection exist in this enigmatic structure.

The baculum is an enigmatic bone within the mammalian glans penis, and the driving forces behind its often bizarre shape have captivated evolutionary biologists for over a century. Hypotheses for the function of the baculum include aiding in intromission, stimulating females and assisting with prolonged mating. Previous attempts to test these hypotheses have focused on the gross size of the baculum and have failed to reach a consensus. We conducted three-dimensional imaging and apply a new method to quantify three-dimensional shape complexity in the carnivoran baculum. We show that socially monogamous species are evolving towards complex-shaped bacula, whereas group-living species are evolving towards simple bacula. Overall three-dimensional baculum shape complexity is not related to relative testes mass, but tip complexity is higher in induced ovulators and species engaging in prolonged copulation. Our study provides evidence of postcopulatory sexual selection pressures driving three-dimensional shape complexity in the carnivore baculum.

Fossil body mass estimation is a well established practice within the field of physical anthropology. Previous studies have relied upon traditional allometric approaches, in which the relationship between one/several skeletal dimensions and body mass in a range of modern taxa is used in a predictive capacity. The lack of relatively complete skeletons has thus far limited the potential application of alternative mass estimation techniques, such as volumetric reconstruction, to fossil hominins. Yet across vertebrate paleontology more broadly, novel volumetric approaches are resulting in predicted values for fossil body mass very different to those estimated by traditional allometry. Here we present a new digital reconstruction of Australopithecus afarensis (A.L. 288-1; 'Lucy') and a convex hull-based volumetric estimate of body mass. The technique relies upon identifying a predictable relationship between the 'shrink-wrapped' volume of the skeleton and known body mass in a range of modern taxa, and subsequent application to an articulated model of the fossil taxa of interest. Our calibration dataset comprises whole body computed tomography (CT) scans of 15 species of modern primate. The resulting predictive model is characterized by a high correlation coefficient (r2 = 0.988) and a percentage standard error of 20%, and performs well when applied to modern individuals of known body mass. Application of the convex hull technique to A. afarensis results in a relatively low body mass estimate of 20.4 kg (95% prediction interval 13.5-30.9 kg). A sensitivity analysis on the articulation of the chest region highlights the sensitivity of our approach to the reconstruction of the trunk, and the incomplete nature of the preserved ribcage may explain the low values for predicted body mass here. We suggest that the heaviest of previous estimates would require the thorax to be expanded to an unlikely extent, yet this can only be properly tested when more complete fossils are available.

Collagen peptide mass fingerprinting of archaeofaunal remains, or Zooarchaeology by Mass Spectrometry (ZooMS), has increasingly established itself as a valuable tool for improving our understanding of highly-fragmented faunal assemblages. Although there have been developments in sampling strategies that have attempted to reduce damage to precious archaeological specimens, these tend to yield spectra inferior in quality to those yielded by destructive approaches. Also, in an effort to mitigate the impacts on faunal assemblages, researchers are beginning to turn to microCT for the digital preservation of specimens, as this enhances their value for further studies. Here we combine ZooMS and microCT, in application to over one hundred scombrid remains from the Hanamiai site in the Marquesas, which spans the time period from initial human colonization by Polynesians at ca. AD 1250 until around 1900. Almost all of the hypurals that yielded collagen fingerprints (71 of 73) were confirmed as skipjack. The results suggest striking continuity over time in fishing practices with an inferred emphasis on fishing strategies involving sailing canoes being present from the time of initial settlement. The focus appears to be on daytime fishing for skipjack rather than on night fishing for yellowfin, kawakawa and dogtooth tuna as might otherwise have been expected.

Sexual dimorphism in the form of elaborate crests, horns and swellings can be a clear indicator of the differing evolutionary pressures to which males and females are subject. However, dimorphism can also be expressed in more subtle shape differences not outwardly obvious to the observer. Whip spiders (Amblypygi) possess a unique pair of spined pedipalps hypothesized to primarily function in prey capture, but also serving multiple other functions. Little is known regarding the intraspecific shape variation of these limbs and its potential causes. Because a role during courtship and male contest has also been hypothesized, sexual selection may contribute to shape differences. As such, we hypothesize that sexual dimorphism will be present in the size and shape of amblypygid pedipalps, with male contest selecting for longer and thicker pedipalps, and larger spines in males. This study aims to test this hypothesis, by quantifying the contribution of ontogeny and sexual dimorphism to shape within the raptorial pedipalps of Damon variegatus. Discriminant function analysis using GMM landmark data reveals statistically significant sexual shape dimorphism in both the tibia and femur of the pedipalp. Contrary to our hypothesis, males display a more gracile pedipalp morphology with reduced spination. Sex differences in the allometric slope and overall size were also found in a number of linear appendicular metrics using Type-II regression. Males have statistically longer pedipalp tibiae, whip femora, and leg two femora. We propose that males have evolved a longer pedipalps in the context of display contest rather than physical aggression. The elongation of structures used in display-based contest and courtship found herein further emphasizes the contribution of visual cues to the evolution of morphology more broadly.

Jordan’s petroglyphs are riddled with bullet holes from target practice and nearby battles. A multidisciplinary forensics team went to Wadi Rum to assess the damage.

The baculum (os penis) is a mineralized bone within the glans of the mammalian penis and is one of the most morphologically diverse structures in the mammal skeleton. Recent experimental work provides compelling evidence for sexual selection shaping the baculum, yet the functional mechanism by which this occurs remains unknown. Previous studies have tested biomechanical hypotheses for the role of the baculum based on simple metrics such as length and diameter, ignoring the wealth of additional shape complexity present. For the first time, to our knowledge, we apply a computational simulation approach (finite-element analysis; FEA) to quantify the three-dimensional biomechanical performance of carnivoran bacula (n= 74) based upon high-resolution micro-computed tomography scans. We find a marginally significant positive correlation between sexual size dimorphism and baculum stress under compressive loading, counter to the ‘vaginal friction’ hypothesis of bacula becoming more robust to overcome resistance during initial intromission. However, a highly significant negative relationship exists between intromission duration and baculum stress under dorsoventral bending. Furthermore, additional FEA simulations confirm that the presence of a ventral groove would reduce deformation of the urethra. We take this as evidence in support of the ‘prolonged intromission’ hypothesis, suggesting the carnivoran baculum has evolved in response to pressures on the duration of copulation and protection of the urethra.

Where rare species are concerned, including those with restricted range, their use for traditional medicine can have disastrous impacts on local populations already under pressure. Difficulty in monitoring such illegal activity has been illustrated by enforcement raids across India since June 2017, with authorities seizing supposedly rare Himalayan plant roots referred to as hatha jodi.

Sexual differences in size and shape are common across the animal kingdom. The study of sexual dimorphism can provide insight into the sexual- and natural-selection pressures experienced by males and females in different species. Arachnids are diverse, comprising over 100,000 species, and exhibit some of the more extreme forms of sexual dimorphism in the animal kingdom, with the males and females of some species differing dramatically in body shape and/or size. Despite this, research on arachnid sexual dimorphism has primarily focused on specific clades as opposed to observing traits across arachnid orders, the smallest of which have received comparatively little attention. This review provides an overview of the research to date on the trends and potential evolutionary drivers for sexual dimorphism and sexual size dimorphism (SSD) in individual arachnid orders, and across arachnids as a whole. The most common trends across Arachnida are female-biased SSD in total body size, male-biased SSD in relative leg length, and sexual dimorphism in pedipalp length and shape. However, the evolution of sexually dimorphic traits within the group is difficult to elucidate due to uncertainty in arachnid phylogenetic relationships. Based on the dataset we gathered here we highlight gaps in our current understanding and suggest areas for future research.

The moment arm of a muscle defines its leverage around a given joint. In a clinical setting, the quantification of muscle moment arms is an important means of establishing the ‘healthy’ functioning of a muscle and in identifying and treating musculoskeletal abnormalities. Elsewhere in modern animal taxa, moment arm studies aim to illuminate adaptions of the musculoskeletal system towards particular locomotor or feeding behaviors. In the absence of kinematic data, paleontologists have likewise relied upon estimated muscle moment arms as a means of reconstructing musculoskeletal function and biomechanical performance in fossil species. With the application of ‘virtual paleontological’ techniques, it is possible to generate increasingly detailed musculoskeletal models of extinct taxa. However, the steps taken to derive such models of complex systems are seldom reported in detail. Here we present a case study for calculating three-dimensional muscle moment arms using Stegosaurus stenops Marsh, 1887 to highlight both the potential and the limitations of this approach in vertebrate paleontology. We find the technique to be mostly insensitive to choices in muscle modeling parameters (particularly relative to other sources of uncertainty in paleontological studies), although exceptions do exist. Of more concern is the current lack of consensus on what functional signals, if any, are contained within moment arm data derived from extant species. Until a correlation between muscle moment arm and function can be broadly identified across a range of modern taxa, the interpretation of moment arms calculated for extinct taxa should be approached with caution.

Body mass is a key parameter for understanding the physiology, biomechanics, and ecology of an organism. Within paleontology, body mass is a fundamental prerequisite for many studies considering body-size evolution, survivorship patterns, and the occurrence of dwarfism and gigantism. The conventional method for estimating fossil body mass relies on allometric scaling relationships derived from skeletal metrics of extant taxa, but the recent application of three-dimensional imaging techniques to paleontology (e.g., surface laser scanning, computed tomography, and photogrammetry) has allowed for the rapid digitization of fossil specimens. Volumetric body-mass estimation methods based on whole articulated skeletons are therefore becoming increasingly popular. Volume-based approaches offer several advantages, including the ability to reconstruct body-mass distribution around the body, and their relative insensitivity to particularly robust or gracile elements, i.e., the so-called ‘one bone effect.’ Yet their application to the fossil record will always be limited by the paucity of well-preserved specimens. Furthermore, uncertainties with regards to skeletal articulation, body density, and soft-tissue distribution must be acknowledged and their effects quantified. Future work should focus on extant taxa to improve our understanding of body composition and increase confidence in volumetric model input parameters.

The limb bones of an elephant are considered to experience similar peak locomotory stresses as a shrew. "Safety factors" are maintained across the entire range of body masses through a combination of robusticity of long bones, postural variation, and modification of gait. The relative contributions of these variables remain uncertain. To test the role of shape change, we undertook X-ray tomographic scans of the leg bones of 60 species of mammals and birds, and extracted geometric properties. The maximum resistible forces the bones could withstand before yield under compressive, bending, and torsional loads were calculated using standard engineering equations incorporating curvature. Positive allometric scaling of cross-sectional properties with body mass was insufficient to prevent negative allometry of bending (F(b) ) and torsional maximum force (F(t) ) (and hence decreasing safety factors) in mammalian (femur F(b) ∞M(b) (0.76) , F(t) ∞M(b) (0.80) ; tibia F(b) ∞M(b) (0.80) , F(t) ∞M(b) (0.76) ) and avian hindlimbs (tibiotarsus F(b) ∞M(b) (0.88) , F(t) ∞M(b) (0.89) ) with the exception of avian femoral F(b) and F(t) . The minimum angle from horizontal a bone must be held while maintaining a given safety factor under combined compressive and bending loads increases with M(b) , with the exception of the avian femur. Postural erectness is shown as an effective means of achieving stress similarity in mammals. The scaling behavior of the avian femur is discussed in light of unusual posture and kinematics.

People suffering from locomotor impairment find turning manoeuvres more challenging than straightahead walking. Turning manoeuvres are estimated to comprise a substantial proportion of steps taken daily, yet research has predominantly focused on straight-line walking, meaning that the basic kinetic, kinematic and foot pressure adaptations required for turning are not as well understood. We investigated how healthy subjects adapt their locomotion patterns to accommodate walking along a gently curved trajectory (radius 2.75 m). Twenty healthy adult participants performed walking tasks at self-selected speeds along straight and curved pathways. For the first time for this mode of turning, plantar pressures were recorded using insole foot pressure sensors while participants’ movements were simultaneously tracked using marker-based 3D motion capture. During the steady-state strides at the apex of the turn, the mean operating point of the inside ankle shifted by 1 degree towards dorsiflexion and that for the outside ankle shifted towards plantarflexion. The largest change in relative joint angle range was an increase in hip rotation in the inside leg (>60%). In addition, the inside foot was subject to a prolonged stance phase and a 10% increase in vertical force in the posteromedial section of the foot compared to straight-line walking. Most of the mechanical change required was therefore generated by the inside leg with hip rotation being a major driver of the gentle turn. This study provides new insight into healthy gait during gentle turns and may help us to understand the mechanics behind some forms of impairment.

Convergent evolution, the acquisition of morphologically similar traits in unrelated taxa due to similar functional demands or environmental factors, is a common phenomenon in the animal kingdom. Consequently, the occurrence of similar form is used routinely to address fundamental questions in morphofunctional research and to infer function in fossils. However, such qualitative assessments can be misleading and it is essential to test form/function relationships quantitatively. The parallel occurrence of a suite of morphologically convergent craniodental characteristics in three herbivorous, phylogenetically disparate dinosaur clades (Sauropodomorpha, Ornithischia, Theropoda) provides an ideal test case. A combination of computational biomechanical models (Finite Element Analysis, Multibody Dynamics Analysis) demonstrate that despite a high degree of morphological similarity between representative taxa (Plateosaurus engelhardti, Stegosaurus stenops, Erlikosaurus andrewsi) from these clades, their biomechanical behaviours are notably different and difficult to predict on the basis of form alone. These functional differences likely reflect dietary specialisations, demonstrating the value of quantitative biomechanical approaches when evaluating form/function relationships in extinct taxa.

Projectile damage to building stone is a widespread phenomenon. Sites damaged 100 years ago during the First World War still see daily use, while in a more contemporary setting numerous reports show the damage to buildings in Babylon, Mosul and Palmyra. While research has been carried out on the long-term effects of conflict such as fire damage, little is known about the protracted damage sustained through the impact of bullets, shrapnel and other metal projectiles outside of the field of engineering focused on ceramics and metals. To investigate alterations to mineral structure caused by projectile damage, impacts were created in medium-grained, well-compacted, mesoporous sandstone samples using 0.22 calibre lead bullets shot at a distance of 20 m. Half these samples were treated with a surface consolidant (Wacker OH 100), to mimic natural cementation of the rock surface. These samples were then tested for changes to surface hardness and moisture movement during temperature cycles of 15–65°C. Petrographic thin section analysis was carried out to investigate the micro-scale deformation associated with high-speed impact. The results surprisingly show that stress build-up behind pre-existing cementation of the surface, as found in heritage sites that have been exposed to moisture and temperature fluctuations for longer periods of time, can be alleviated with a bullet impact. However, fracture networks and alteration of the mineral matrices still form a weak point within the structure, even at a relatively low impact calibre. This initial study illustrates the need for geomorphologists, geologists, engineers and heritage specialists to work collectively to gain further insights into the long-term impact of higher calibre armed warfare on heritage deterioration.

Body mass is a critical parameter used to constrain biomechanical and physiological traits of organisms. Volumetric methods are becoming more common as techniques for estimating the body masses of fossil vertebrates. However, they are often accused of excessive subjective input when estimating the thickness of missing soft tissue. Here, we demonstrate an alternative approach where a minimum convex hull is derived mathematically from the point cloud generated by laser-scanning mounted skeletons. This has the advantage of requiring minimal user intervention and is thus more objective and far quicker. We test this method on 14 relatively large-bodied mammalian skeletons and demonstrate that it consistently underestimates body mass by 21 per cent with minimal scatter around the regression line. We therefore suggest that it is a robust method of estimating body mass where a mounted skeletal reconstruction is available and demonstrate its usage to predict the body mass of one of the largest, relatively complete sauropod dinosaurs: Giraffatitan brancai (previously Brachiosaurus ) as 23200 kg.

Although Stegosaurus is one of the most iconic dinosaurs, well-preserved fossils are rare and as a consequence there is still much that remains unknown about the taxon. A new, exceptionally complete individual affords the opportunity to describe the anatomy of Stegosaurus in detail for the first time in over a century, and enables additional comparisons with other stegosaurian dinosaurs. The new specimen is from the Red Canyon Ranch Quarry, near Shell Wyoming, and appears to have been so well preserved because it was buried rapidly in a pond or body of standing water immediately after death. The quarry is probably located in the middle part of the Morrison Formation, which is believed to be Tithonian in age in this area. The specimen is referable to Stegosaurus stenops based on the possession of an edentulous anterior portion of the dentary and elevated postzygapophyses on the cervical vertebrae. New information provided by the specimen concerns the morphology of the vertebrae, the iliosacral block and dermal armor. Several aspects of its morphology indicate the individual was not fully skeletally mature at the time of death, corroborating a previous histological study.

The external appearance of the dodo (Raphus cucullatus,Linnaeus, 1758) has been a source of considerable intrigue, as contemporaneous accounts or depictions are rare. The body mass of the dodo has been particularly contentious, with the flightless pigeon alternatively reconstructed as slim or fat depending upon the skeletal metric used as the basis for mass prediction. Resolving this dichotomy and obtaining a reliable estimate for mass is essential before future analyses regarding dodo life history, physiology or biomechanics can be conducted. Previous mass estimates of the dodo have relied upon predictive equations based upon hind limb dimensions of extant pigeons. Yet the hind limb proportions of dodo have been found to differ considerably from those of their modern relatives, particularly with regards to midshaft diameter. Therefore, application of predictive equations to unusually robust fossil skeletal elements may bias mass estimates. We present a whole-body computed tomography (CT) -based mass estimation technique for application to the dodo. We generate 3D volumetric renders of the articulated skeletons of 20 species of extant pigeons, and wrap minimum-fit ‘convex hulls’ around their bony extremities. Convex hull volume is subsequently regressed against mass to generate predictive models based upon whole skeletons. Our best-performing predictive model is characterized by high correlation coefficients and low mean squared error (a= − 2.31,b= 0.90,r2= 0.97, MSE = 0.0046). When applied to articulated composite skeletons of the dodo (National Museums Scotland, NMS.Z.1993.13; Natural History Museum, NHMUK A.9040 and S/1988.50.1), we estimate eviscerated body masses of 8–10.8 kg. When accounting for missing soft tissues, this may equate to live masses of 10.6–14.3 kg. Mass predictions presented here overlap at the lower end of those previously published, and support recent suggestions of a relatively slim dodo. CT-based reconstructions provide a means of objectively estimating mass and body segment properties of extinct species using whole articulated skeletons.

Body mass is a key biological variable, but difficult to assess from fossils. Various techniques exist for estimating body mass from skeletal parameters, but few studies have compared outputs from different methods. Here, we apply several mass estimation methods to an exceptionally complete skeleton of the dinosaur Stegosaurus . Applying a volumetric convex-hulling technique to a digital model of Stegosaurus , we estimate a mass of 1560 kg (95% prediction interval 1082–2256 kg) for this individual. By contrast, bivariate equations based on limb dimensions predict values between 2355 and 3751 kg and require implausible amounts of soft tissue and/or high body densities. When corrected for ontogenetic scaling, however, volumetric and linear equations are brought into close agreement. Our results raise concerns regarding the application of predictive equations to extinct taxa with no living analogues in terms of overall morphology and highlight the sensitivity of bivariate predictive equations to the ontogenetic status of the specimen. We emphasize the significance of rare, complete fossil skeletons in validating widely applied mass estimation equations based on incomplete skeletal material and stress the importance of accurately determining specimen age prior to further analyses.

Classic beam theory is frequently used in biomechanics to model the stress behaviour of vertebrate long bones, particularly when creating intraspecific scaling models. Although methodologically straightforward, classic beam theory requires complex irregular bones to be approximated as slender beams, and the errors associated with simplifying complex organic structures to such an extent are unknown. Alternative approaches, such as finite element analysis (FEA), while much more time-consuming to perform, require no such assumptions. This study compares the results obtained using classic beam theory with those from FEA to quantify the beam theory errors and to provide recommendations about when a full FEA is essential for reasonable biomechanical predictions. High-resolution computed tomographic scans of eight vertebrate long bones were used to calculate diaphyseal stress owing to various loading regimes. Under compression, FEA values of minimum principal stress (σmin) were on average 142 per cent (±28% s.e.) larger than those predicted by beam theory, with deviation between the two models correlated to shaft curvature (two-tailedp= 0.03,r2= 0.56). Under bending, FEA values of maximum principal stress (σmax) and beam theory values differed on average by 12 per cent (±4% s.e.), with deviation between the models significantly correlated to cross-sectional asymmetry at midshaft (two-tailedp= 0.02,r2= 0.62). In torsion, assuming maximum stress values occurred at the location of minimum cortical thickness brought beam theory and FEA values closest in line, and in this case FEA values ofτtorsionwere on average 14 per cent (±5% s.e.) higher than beam theory. Therefore, FEA is the preferred modelling solution when estimates of absolute diaphyseal stress are required, although values calculated by beam theory for bending may be acceptable in some situations.

The extinct moa of New Zealand included three families (Megalapterygidae; Dinornithidae; Emeidae) of flightless palaeognath bird, ranging in mass from 200 kg. They are perceived to have evolved extremely robust leg bones, yet current estimates of body mass have very wide confidence intervals. Without reliable estimators of mass, the extent to which dinornithid and emeid hindlimbs were more robust than modern species remains unclear. Using the convex hull volumetric-based method on CT-scanned skeletons, we estimate the mass of a female Dinornis robustus (Dinornithidae) at 196 kg (range 155–245 kg) and of a female Pachyornis australis (Emeidae) as 50 kg (range 33–68 kg). Finite element analysis of CT-scanned femora and tibiotarsi of two moa and six species of modern palaeognath showed that P. australis experienced the lowest values for stress under all loading conditions, confirming it to be highly robust. In contrast, stress values in the femur of D. robustus were similar to those of modern flightless birds, whereas the tibiotarsus experienced the highest level of stress of any palaeognath. We consider that these two families of Dinornithiformes diverged in their biomechanical responses to selection for robustness and mobility, and exaggerated hindlimb strength was not the only successful evolutionary pathway.

The limb bones of an elephant are considered to experience similar peak locomotory stresses as a shrew. "Safety factors" are maintained across the entire range of body masses through a combination of robusticity of long bones, postural variation, and modification of gait. The relative contributions of these variables remain uncertain. To test the role of shape change, we undertook X-ray tomographic scans of the leg bones of 60 species of mammals and birds, and extracted geometric properties. The maximum resistible forces the bones could withstand before yield under compressive, bending, and torsional loads were calculated using standard engineering equations incorporating curvature. Positive allometric scaling of cross-sectional properties with body mass was insufficient to prevent negative allometry of bending (F(b) ) and torsional maximum force (F(t) ) (and hence decreasing safety factors) in mammalian (femur F(b) ∞M(b) (0.76) , F(t) ∞M(b) (0.80) ; tibia F(b) ∞M(b) (0.80) , F(t) ∞M(b) (0.76) ) and avian hindlimbs (tibiotarsus F(b) ∞M(b) (0.88) , F(t) ∞M(b) (0.89) ) with the exception of avian femoral F(b) and F(t) . The minimum angle from horizontal a bone must be held while maintaining a given safety factor under combined compressive and bending loads increases with M(b) , with the exception of the avian femur. Postural erectness is shown as an effective means of achieving stress similarity in mammals. The scaling behavior of the avian femur is discussed in light of unusual posture and kinematics.

Body mass is a fundamental physical property of an individual and has enormous bearing upon ecology and physiology. Generating reliable estimates for body mass is therefore a necessary step in many palaeontological studies. Whilst early reconstructions of mass in extinct species relied upon isolated skeletal elements, volumetric techniques are increasingly applied to fossils when skeletal completeness allows. We apply a new ‘alpha shapes’ (α-shapes) algorithm to volumetric mass estimation in quadrupedal mammals.α-shapes are defined by: (i) the underlying skeletal structure to which they are fitted; and (ii) the valueα, determining the refinement of fit. For a given skeleton, a range ofα-shapes may be fitted around the individual, spanning from very coarse to very fine. We fitα-shapes to three-dimensional models of extant mammals and calculate volumes, which are regressed against mass to generate predictive equations. Our optimal model is characterized by a high correlation coefficient and mean square error (r2=0.975, m.s.e.=0.025). When applied to the woolly mammoth (Mammuthus primigenius) and giant ground sloth (Megatherium americanum), we reconstruct masses of 3635 and 3706 kg, respectively. We considerα-shapes an improvement upon previous techniques as resulting volumes are less sensitive to uncertainties in skeletal reconstructions, and do not require manual separation of body segments from skeletons.

The volumetric method of ‘convex hulling’ has recently been put forward as a mass prediction technique for fossil vertebrates. Convex hulling involves the calculation of minimum convex hull volumes (vol CH) from the complete mounted skeletons of modern museum specimens, which are subsequently regressed against body mass (M b) to derive predictive equations for extinct species. The convex hulling technique has recently been applied to estimate body mass in giant sauropods and fossil ratites, however the biomechanical signal contained within vol CH has remained unclear. Specifically, when vol CH scaling departs from isometry in a group of vertebrates, how might this be interpreted? Here we derive predictive equations for primates, non-primate mammals and birds and compare the scaling behaviour of M b to vol CH between groups. We find predictive equations to be characterised by extremely high correlation coefficients (r 2 = 0.97–0.99) and low mean percentage prediction error (11–20%). Results suggest non-primate mammals scale body mass to vol CH isometrically (b = 0.92, 95%CI = 0.85–1.00, p = 0.08). Birds scale body mass to vol CH with negative allometry (b = 0.81, 95%CI = 0.70–0.91, p = 0.011) and apparent density (vol CH/M b) therefore decreases with mass (r 2 = 0.36, p