Using your head - cranial steering in pterosaurs.

The vast majority of pterosaurs are characterized by relatively large, elongate heads that are often adorned with large, elaborate crests. Projecting out in front of the body, these large heads and any crests must have had an aerodynamic effect. The working hypothesis of the present study is that these oversized heads were used to control the left-right motions of the body during flight. Using digital models of eight non-pterodactyloids ("rhamphorhyncoids") and ten pterodactyloids, the turning moments associated with the head + neck show a close and consistent correspondence with the rotational inertia of the whole body about a vertical axis in both groups, supporting the idea of a functional relationship. Turning moments come from calculating the lateral area of the head (plus any crests) and determining the associated lift (aerodynamic force) as a function of flight speed, with flight speeds being based on body mass. Rotational inertias were calculated from the three-dimensional mass distribution of the axial body, the limbs, and the flight membranes. The close correlation between turning moment and rotational inertia was used to revise the life restorations of two pterosaurs and to infer relatively lower flight speeds in another two.

Diving dinosaurs? Caveats on the use of bone compactness and pFDA for inferring lifestyle.

The lifestyle of spinosaurid dinosaurs has been a topic of lively debate ever since the unveiling of important new skeletal parts for Spinosaurus aegyptiacus in 2014 and 2020. Disparate lifestyles for this taxon have been proposed in the literature; some have argued that it was semiaquatic to varying degrees, hunting fish from the margins of water bodies, or perhaps while wading or swimming on the surface; others suggest that it was a fully aquatic underwater pursuit predator. The various proposals are based on equally disparate lines of evidence. A recent study by Fabbri and coworkers sought to resolve this matter by applying the statistical method of phylogenetic flexible discriminant analysis to femur and rib bone diameters and a bone microanatomy metric called global bone compactness. From their statistical analyses of datasets based on a wide range of extant and extinct taxa, they concluded that two spinosaurid dinosaurs (S. aegyptiacus, Baryonyx walkeri) were fully submerged "subaqueous foragers," whereas a third spinosaurid (Suchomimus tenerensis) remained a terrestrial predator. We performed a thorough reexamination of the datasets, analyses, and methodological assumptions on which those conclusions were based, which reveals substantial problems in each of these areas. In the datasets of exemplar taxa, we found unsupported categorization of taxon lifestyle, inconsistent inclusion and exclusion of taxa, and inappropriate choice of taxa and independent variables. We also explored the effects of uncontrolled sources of variation in estimates of bone compactness that arise from biological factors and measurement error. We found that the ability to draw quantitative conclusions is limited when taxa are represented by single data points with potentially large intrinsic variability. The results of our analysis of the statistical method show that it has low accuracy when applied to these datasets and that the data distributions do not meet fundamental assumptions of the method. These findings not only invalidate the conclusions of the particular analysis of Fabbri et al. but also have important implications for future quantitative uses of bone compactness and discriminant analysis in paleontology.

Growth constraints set an upper limit to theropod dinosaur body size.

Despite nearly 200 years of scientific collecting and study, none of the extinct, bipedal, predatory, theropod dinosaurs have been reliably shown to exceed 12 m in length. Using digital 3D models of theropods with lengths spanning 80 cm to 12 m, their body masses were found to scale to the 3.5 power of body lengths. The lateral area of the pelvis and the cross-sectional area of the tail base of these animals corresponds to the cross-sectional areas of key muscle groups important for balance and locomotion, and both scale to the 2.4 power of body length. Body accelerations in the lateral and forward directions are, using F = ma, given by dividing muscle area (force proxy) by body mass. Plotting these acceleration estimates against body length shows them to decrease exponentially. The largest theropods with body lengths of 10-12 m have less than 10% of the acceleration capacity of the smaller forms. The distinct lack of fossil remains of theropods demonstrably longer than 12 m suggests that the theropod body plan had an upper size limit based on a minimum acceleration threshold. Rotational inertia of the theropod body was found to be proportional to body length raised to the 5.5 power, and with increasing length, the capacity for agility would rapidly diminish. The tight relationship between theropod pelvic area and body length allows for the estimation of body lengths of specimens lacking complete axial skeletons, and this is done for four, large, well-preserved pelves.

Spinosaurus is not an aquatic dinosaur.

A predominantly fish-eating diet was envisioned for the sail-backed theropod dinosaur Spinosaurus aegyptiacus when its elongate jaws with subconical teeth were unearthed a century ago in Egypt. Recent discovery of the high-spined tail of that skeleton, however, led to a bolder conjecture that S. aegyptiacus was the first fully aquatic dinosaur. The 'aquatic hypothesis' posits that S. aegyptiacus was a slow quadruped on land but a capable pursuit predator in coastal waters, powered by an expanded tail. We test these functional claims with skeletal and flesh models of S. aegyptiacus. We assembled a CT-based skeletal reconstruction based on the fossils, to which we added internal air and muscle to create a posable flesh model. That model shows that on land S. aegyptiacus was bipedal and in deep water was an unstable, slow-surface swimmer (<1 m/s) too buoyant to dive. Living reptiles with similar spine-supported sails over trunk and tail are used for display rather than aquatic propulsion, and nearly all extant secondary swimmers have reduced limbs and fleshy tail flukes. New fossils also show that Spinosaurus ranged far inland. Two stages are clarified in the evolution of Spinosaurus, which is best understood as a semiaquatic bipedal ambush piscivore that frequented the margins of coastal and inland waterways.

Fluid dynamics, scaling laws and plesiosaur locomotion.

The evolutionary success of plesiosaurs has led to much attention regarding the dynamics of their locomotion. They exhibit identical tandem flippers, which is unique among all living and extinct species. However, these tandem flippers have been a source of debate regarding plesiosaurs' locomotion and behavior. Here we propose a new approach to studying plesiosaur locomotion based on universal scaling laws in fluid dynamics, which were used to estimate reduced frequency to characterize unsteadiness of an airfoil. It was found that, while the reduced frequency of plesiosaurs with high-aspect ratio flippers is similar to that of sea turtles, the most commonly used living analog, lower aspect ratio plesiosaurs were more similar in reduced frequency to penguins. This implies that plesiosaurs may have had large variations in agility among themselves, depending in particular on the specimen's flipper aspect ratio. While our results are consistent with the previous literature indicating a relationship between plesiosaur neck length and agility, our work supports broad and diverse analogies to living animals. Moreover, based on our results, cruising reduced frequency has some predictive value into manoeuvring behavior, rather than simply cruising behavior.

Dental microwear texture analysis along reptile tooth rows: complex variation with non-dietary variables.

Dental microwear texture analysis (DMTA) is a powerful technique for reconstructing the diets of extant and extinct taxa. Few studies have investigated intraspecific microwear differences along with tooth rows and the influence of endogenous non-dietary variables on texture characteristics. Sampling teeth that are minimally affected by non-dietary variables is vital for robust dietary reconstructions, especially for taxa with non-occlusal (non-chewing) dentitions as no standardized sampling strategies currently exist. Here, we apply DMTA to 13 species of extant reptile (crocodilians and monitor lizards) to investigate intraspecific microwear differences along with tooth rows and to explore the influence of three non-dietary variables on exhibited differences: (i) tooth position, (ii) mechanical advantage, and (iii) tooth aspect ratio. Five species exhibited intraspecific microwear differences. In several crocodilians, the distally positioned teeth exhibited the 'roughest' textures, and texture characteristics correlated with all non-dietary variables. By contrast, the mesial teeth of the roughneck monitor (Varanus rudicollis) exhibited the 'roughest' textures, and texture characteristics did not correlate with aspect ratio. These results are somewhat consistent with how reptiles preferentially use their teeth during feeding. We argue that DMTA has the potential to track mechanical and behavioural differences in tooth use which should be taken into consideration in future dietary reconstructions.

Maniraptoran pelvic musculature highlights evolutionary patterns in theropod locomotion on the line to birds.

Locomotion is a fundamental aspect of palaeobiology and often investigated by comparing osteological structures and proportions. Previous studies document a stepwise accumulation of avian-like features in theropod dinosaurs that accelerates in the clade Maniraptora. However, the soft tissues that influenced the skeleton offer another perspective on locomotory adaptations. Examination of the pelvis for osteological correlates of hind limb and tail musculature allowed reconstruction of primary locomotory muscles across theropods and their closest extant relatives. Additionally, the areas of pelvic muscle origins were quantified to measure relative differences within and between taxa, to compare morphological features associated with cursoriality, and offer insight into the evolution of locomotor modules. Locomotory inferences based on myology often corroborate those based on osteology, although they occasionally conflict and indicate greater complexity than previously appreciated. Maniraptoran pelvic musculature underscores previous studies noting the multifaceted nature of cursoriality and suggests that a more punctuated step in caudal decoupling occurred at or near the base of Maniraptora.

Dietary palaeoecology of an Early Cretaceous armoured dinosaur (Ornithischia; Nodosauridae) based on floral analysis of stomach contents.

The exceptionally well-preserved holotype of the armoured dinosaur Borealopelta markmitchelli (Ornithischia; Nodosauridae) from the Early Cretaceous (Clearwater Formation) of northern Alberta preserves a distinct mass within the abdominal cavity. Fourteen independent criteria (including: co-allochthony, anatomical position, gastroliths) support the interpretation of this mass as ingested stomach contents-a cololite. Palynomorphs in the cololite are a subset of the more diverse external sample. Analysis of the cololite documents well-preserved plant material dominated by leaf tissue (88%), including intact sporangia, leaf cross-sections and cuticle, but also including stems, wood and charcoal. The leaf fraction is dominated (85%) by leptosporangiate ferns (subclass Polypodiidae), with low cycad-cycadophyte (3%) and trace conifer foliage. These data represent the most well-supported and detailed direct evidence of diet in an herbivorous dinosaur. Details of the dietary palaeoecology of this nodosaur are revealed, including: selective feeding on ferns; preferential ingestion of leptosporangiate ferns to the exclusion of Osmundaceae and eusporangiate ferns such as Marattiaceae; and incidental consumption of cycad-cycadophyte and conifer leaves. The presence of significant (6%) charcoal may represent the dietary use of recently burned conifer forest undergoing fern succession, early evidence of a fire succession ecology, as is associated with many modern large herbivores.

Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods.

SYNOPSIS: Tyrannosaurid dinosaurs had large preserved leg muscle attachments and low rotational inertia relative to their body mass, indicating that they could turn more quickly than other large theropods. METHODS: To compare turning capability in theropods, we regressed agility estimates against body mass, incorporating superellipse-based modeled mass, centers of mass, and rotational inertia (mass moment of inertia). Muscle force relative to body mass is a direct correlate of agility in humans, and torque gives potential angular acceleration. Agility scores therefore include rotational inertia values divided by proxies for (1) muscle force (ilium area and estimates of m. caudofemoralis longus cross-section), and (2) musculoskeletal torque. Phylogenetic ANCOVA (phylANCOVA) allow assessment of differences in agility between tyrannosaurids and non-tyrannosaurid theropods (accounting for both ontogeny and phylogeny). We applied conditional error probabilities a(p) to stringently test the null hypothesis of equal agility. RESULTS: Tyrannosaurids consistently have agility index magnitudes twice those of allosauroids and some other theropods of equivalent mass, turning the body with both legs planted or pivoting over a stance leg. PhylANCOVA demonstrates definitively greater agilities in tyrannosaurids, and phylogeny explains nearly all covariance. Mass property results are consistent with those of other studies based on skeletal mounts, and between different figure-based methods (our main mathematical slicing procedures, lofted 3D computer models, and simplified graphical double integration). IMPLICATIONS: The capacity for relatively rapid turns in tyrannosaurids is ecologically intriguing in light of their monopolization of large (>400 kg), toothed dinosaurian predator niches in their habitats.

A method for deducing neck mobility in plesiosaurs, using the exceptionally preserved Nichollssaura borealis.

The elongate-necked aquatic plesiosaurs existed for 135 Myr during the Mesozoic. The function of this elongate neck is a point of debate. Using computed tomography and three-dimensional (3D) modelling, the range of motion (ROM) of the plesiosaur Nichollssaura borealis neck was assessed. To quantify the ROM, the intervertebral mobility was measured along the cervical vertebral column. This was done by manipulating the 3D models in the lateral and dorsoventral directions during two trials. The first assessed the mean intervertebral ROM between pairs of cervical vertebrae along the entire column, and the second assessed ROM with reduced intervertebral spaces. The results suggest that there may be preference for lateral neck movements in N. borealis, which could correspond to an ecological function related to prey capture. This study demonstrates that 3D modelling is an effective tool for assessing function morphology for structures where no good modern analogue exists.

A buoyancy, balance and stability challenge to the hypothesis of a semi-aquatic Spinosaurus Stromer, 1915 (Dinosauria: Theropoda).

A recent interpretation of the fossil remains of the enigmatic, large predatory dinosaur Spinosaurus aegyptiacus Stromer 1915 proposed that it was specially adapted for a semi-aquatic mode of life-a first for any predatory dinosaur. To test some aspects of this suggestion, a three-dimensional, digital model of the animal that incorporates regional density variations, lungs and air sacs was generated, and the flotation potential of the model was investigated using specially written software. It was found that Spinosaurus would have been able to float with its head clear of the water surface, although it was laterally unstable and would tend to roll onto its side. Similarly detailed models of another spinosaurid Baryonyx (Suchomimus) tenerensis Sereno et al. 1998, along with models of the more distantly related Tyrannosaurus rex Osborn 1905, Allosaurus fragilis Marsh 1877, Struthiomimus altus Lambe 1902, and Coelophysis bauri Cope 1887 were also able to float in positions that enabled the animals to breathe freely, showing that there is nothing exceptional about a floating Spinosaurus. Validation of the modelling methods was done with floated models of an alligator and an emperor penguin. The software also showed that the center of mass of Spinosaurus was much closer to the hips than previously estimated, similar to that observed in other theropods, implying that this dinosaur would still have been a competent walker on land. With its pneumatised skeleton and a system of air sacs (modelled after birds), the Spinosaurus model was found to be unsinkable, even with its lungs deflated by 75%, and this would greatly hinder a semi-aquatic, pursuit predator. The conclusion is that Spinosaurus may have been specialized for a shoreline or shallow water mode of life, but would still have been a competent terrestrial animal.

Pterosaur dietary hypotheses: a review of ideas and approaches.

Pterosaurs are an extinct group of Mesozoic flying reptiles, whose fossil record extends from approximately 210 to 66 million years ago. They were integral components of continental and marginal marine ecosystems, yet their diets remain poorly constrained. Numerous dietary hypotheses have been proposed for different pterosaur groups, including insectivory, piscivory, carnivory, durophagy, herbivory/frugivory, filter-feeding and generalism. These hypotheses, and subsequent interpretations of pterosaur diet, are supported by qualitative (content fossils, associations, ichnology, comparative anatomy) and/or quantitative (functional morphology, stable isotope analysis) evidence. Pterosaur dietary interpretations are scattered throughout the literature with little attention paid to the supporting evidence. Reaching a robustly supported consensus on pterosaur diets is important for understanding their dietary evolution, and their roles in Mesozoic ecosystems. A comprehensive examination of the pterosaur literature identified 314 dietary interpretations (dietary statement plus supporting evidence) from 126 published studies. Multiple alternative diets have been hypothesised for most principal taxonomic pterosaur groups. Some groups exhibit a high degree of consensus, supported by multiple lines of evidence, while others exhibit less consensus. Qualitative evidence supports 87.3% of dietary interpretations, with comparative anatomy most common (62.1% of total). More speciose groups of pterosaur tend to have a greater range of hypothesised diets. Consideration of dietary interpretations within alternative phylogenetic contexts reveals high levels of consensus between equivalent monofenestratan groups, and lower levels of consensus between equivalent non-monofenestratan groups. Evaluating the possible non-biological controls on apparent patterns of dietary diversity reveals that numbers of dietary interpretations through time exhibit no correlation with patterns of publication (number of peer-reviewed publications through time). 73.8% of dietary interpretations were published in the 21st century. Overall, consensus interpretations of pterosaur diets are better accounted for by non-biological signals, such as the impact of the respective quality of the fossil record of different pterosaur groups on research levels. That many interpretations are based on qualitative, often untestable lines of evidence adds significant noise to the data. More experiment-led pterosaur dietary research, with greater consideration of pterosaurs as organisms with independent evolutionary histories, will lead to more robust conclusions drawn from repeatable results. This will allow greater understanding of pterosaur dietary diversity, disparity and evolution and facilitate reconstructions of Mesozoic ecosystems.

An Exceptionally Preserved Three-Dimensional Armored Dinosaur Reveals Insights into Coloration and Cretaceous Predator-Prey Dynamics.

Predator-prey dynamics are an important evolutionary driver of escalating predation mode and efficiency, and commensurate responses of prey [1-3]. Among these strategies, camouflage is important for visual concealment, with countershading the most universally observed [4-6]. Extant terrestrial herbivores free of significant predation pressure, due to large size or isolation, do not exhibit countershading. Modern predator-prey dynamics may not be directly applicable to those of the Mesozoic due to the dominance of very large, visually oriented theropod dinosaurs [7]. Despite thyreophoran dinosaurs' possessing extensive dermal armor, some of the most extreme examples of anti-predator structures [8, 9], little direct evidence of predation on these and other dinosaur megaherbivores has been documented. Here we describe a new, exquisitely three-dimensionally preserved nodosaurid ankylosaur, Borealopelta markmitchelli gen. et sp. nov., from the Early Cretaceous of Alberta, which preserves integumentary structures as organic layers, including continuous fields of epidermal scales and intact horn sheaths capping the body armor. We identify melanin in the organic residues through mass spectroscopic analyses and observe lighter pigmentation of the large parascapular spines, consistent with display, and a pattern of countershading across the body. With an estimated body mass exceeding 1,300 kg, B. markmitchelli was much larger than modern terrestrial mammals that either are countershaded or experience significant predation pressure as adults. Presence of countershading suggests predation pressure strong enough to select for concealment in this megaherbivore despite possession of massive dorsal and lateral armor, illustrating a significant dichotomy between Mesozoic predator-prey dynamics and those of modern terrestrial systems.

A specimen of Rhamphorhynchus with soft tissue preservation, stomach contents and a putative coprolite.

Despite being known for nearly two centuries, new specimens of the derived non-pterodactyloid pterosaur Rhamphorhynchus continue to be discovered and reveal new information about their anatomy and palaeobiology. Here we describe a specimen held in the collections of the Royal Tyrrell Museum of Palaeontology, Alberta, Canada that shows both preservation and impressions of soft tissues, and also preserves material interpreted as stomach contents of vertebrate remains and, uniquely, a putative coprolite. The specimen also preserves additional evidence for fibers in the uropatagium.

A new horned dinosaur reveals convergent evolution in cranial ornamentation in Ceratopsidae.

Ceratopsid (horned) dinosaurs are an iconic group of large-bodied, quadrupedal, herbivorous dinosaurs that evolved in the Late Cretaceous and were largely restricted to western North America [1-5]. Ceratopsids are easily recognized by their cranial ornamentation in the form of nasal and postorbital horns and frill (capped by epiossifications); these structures show high morphological disparity and also represent the largest cranial display structures known to have evolved [2, 4]. Despite their restricted occurrence in time and space, this group has one of the best fossil records within Dinosauria, showing a rapid diversification in horn and frill morphology [1]. Here a new genus and species of chasmosaurine ceratopsid is described based on a nearly complete and three-dimensionally preserved cranium recovered from the uppermost St. Mary River Formation (Maastrichtian) of southwestern Alberta. Regaliceratops peterhewsi gen. et sp. nov. exhibits many unique characters of the frill and is characterized by a large nasal horncore, small postorbital horncores, and massive parietal epiossifications. Cranial morphology, particularly the epiossifications, suggests close affinity with the late Campanian/early Maastrichian taxon Anchiceratops, as well as with the late Maastrichtian taxon Triceratops. A median epiparietal necessitates a reassessment of epiossification homology and results in a more resolved phylogeny. Most surprisingly, Regaliceratops exhibits a suite of cranial ornamentations that are superficially similar to Campanian centrosaurines, indicating both exploration of novel display morphospace in Chasmosaurinae, especially Maastrichtian forms, and convergent evolution in horn morphology with the recently extinct Centrosaurinae. This marks the first time that evolutionary convergence in horn-like display structures has been demonstrated between dinosaur clades, similar to those seen in fossil and extant mammals [6].

Balance and Strength - Estimating the Maximum Prey-Lifting Potential of the Large Predatory Dinosaur Carcharodontosaurus saharicus.

Motivated by the work of palaeo-art "Double Death (2011)," a biomechanical analysis using three-dimensional digital models was conducted to assess the potential of a pair of the large, Late Cretaceous theropod dinosaur Carcharodontosaurus saharicus to successfully lift a medium-sized sauropod and not lose balance. Limaysaurus tessonei from the Late Cretaceous of South America was chosen as the sauropod as it is more completely known, but closely related to the rebbachisaurid sauropods found in the same deposits with C. saharicus. The body models incorporate the details of the low-density regions associated with lungs, systems of air sacs, and pneumatized axial skeletal regions. These details, along with the surface meshes of the models, were used to estimate the body masses and centers of mass of the two animals. It was found that a 6 t C. saharicus could successfully lift a mass of 2.5 t and not lose balance as the combined center of mass of the body and the load in the jaws would still be over the feet. However, the neck muscles were found to only be capable of producing enough force to hold up the head with an added mass of 424 kg held at the midpoint of the maxillary tooth row. The jaw adductor muscles were more powerful, and could have held a load of 512 kg. The more limiting neck constraint leads to the conclusion that two, adult C. saharicus could successfully lift a L. tessonei with a maximum body mass of 850 kg and a body length of 8.3 m.

What drove reversions to quadrupedality in ornithischian dinosaurs? Testing hypotheses using centre of mass modelling.

The exceptionally rare transition to quadrupedalism from bipedal ancestors occurred on three independent occasions in ornithischian dinosaurs. The possible driving forces behind these transitions remain elusive, but several hypotheses-including the development of dermal armour and the expansion of head size and cranial ornamentation-have been proposed to account for this major shift in stance. We modelled the position of the centre of mass (CoM) in several exemplar ornithischian taxa and demonstrate that the anterior shifts in CoM position associated with the development of an enlarged skull ornamented with horns and frills for display/defence may have been one of the drivers promoting ceratopsian quadrupedality. A posterior shift in CoM position coincident with the development of extensive dermal armour in thyreophorans demonstrates this cannot have been a primary causative mechanism for quadrupedality in this clade. Quadrupedalism developed in response to different selective pressures in each ornithischian lineage, indicating different evolutionary pathways to convergent quadrupedal morphology.

Sauropod necks: are they really for heat loss?

Three-dimensional digital models of 16 different sauropods were used to examine the scaling relationship between metabolism and surface areas of the whole body, the neck, and the tail in an attempt to see if the necks could have functioned as radiators for the elimination of excess body heat. The sauropod taxa sample ranged in body mass from a 639 kg juvenile Camarasaurus to a 25 t adult Brachiosaurus. Metabolism was assumed to be directly proportional to body mass raised to the ¾ power, and estimates of body mass accounted for the presence of lungs and systems of air sacs in the trunk and neck. Surface areas were determined by decomposing the model surfaces into triangles and their areas being computed by vector methods. It was found that total body surface area was almost isometric with body mass, and that it showed negative allometry when plotted against metabolic rate. In contrast, neck area showed positive allometry when plotted against metabolic rate. Tail area show negative allometry with respect to metabolic rate. The many uncertainties about the biology of sauropods, and the variety of environmental conditions that different species experienced during the groups 150 million years of existence, make it difficult to be absolutely certain about the function of the neck as a radiator. However, the functional combination of the allometric increase of neck area, the systems of air sacs in the neck and trunk, the active control of blood flow between the core and surface of the body, changing skin color, and strategic orientation of the neck with respect to wind, make it plausible that the neck could have functioned as a radiator to avoid over-heating.

Blood flow to long bones indicates activity metabolism in mammals, reptiles and dinosaurs.

The cross-sectional area of a nutrient foramen of a long bone is related to blood flow requirements of the internal bone cells that are essential for dynamic bone remodelling. Foramen area increases with body size in parallel among living mammals and non-varanid reptiles, but is significantly larger in mammals. An index of blood flow rate through the foramina is about 10 times higher in mammals than in reptiles, and even higher if differences in blood pressure are considered. The scaling of foramen size correlates well with maximum whole-body metabolic rate during exercise in mammals and reptiles, but less well with resting metabolic rate. This relates to the role of blood flow associated with bone remodelling during and following activity. Mammals and varanid lizards have much higher aerobic metabolic rates and exercise-induced bone remodelling than non-varanid reptiles. Foramen areas of 10 species of dinosaur from five taxonomic groups are generally larger than from mammals, indicating a routinely highly active and aerobic lifestyle. The simple measurement holds possibilities offers the possibility of assessing other groups of extinct and living vertebrates in relation to body size, behaviour and habitat.

Predicting the buoyancy, equilibrium and potential swimming ability of giraffes by computational analysis.

Giraffes (Giraffa camelopardalis) are often stated to be unable to swim, and while few observations supporting this have ever been offered, we sought to test the hypothesis that giraffes exhibited a body shape or density unsuited for locomotion in water. We assessed the floating capability of giraffes by simulating their buoyancy with a three-dimensional mathematical/computational model. A similar model of a horse (Equus caballus) was used as a control, and its floating behaviour replicates the observed orientations of immersed horses. The floating giraffe model has its neck sub-horizontal, and the animal would struggle to keep its head clear of the water surface. Using an isometrically scaled-down giraffe model with a total mass equal to that of the horse, the giraffe's proportionally larger limbs have much higher rotational inertias than do those of horses, and their wetted surface areas are 13.5% greater relative to that of the horse, thus making rapid swimming motions more strenuous. The mean density of the giraffe model (960 gm/l) is also higher than that of the horse (930 gm/l), and closer to that causing negative buoyancy (1000 gm/l). A swimming giraffe - forced into a posture where the neck is sub-horizontal and with a thorax that is pulled downwards by the large fore limbs - would not be able to move the neck and limbs synchronously as giraffes do when moving on land, possibly further hampering the animal's ability to move its limbs effectively underwater. We found that a full-sized, adult giraffe will become buoyant in water deeper than 2.8m. While it is not impossible for giraffes to swim, we speculate that they would perform poorly compared to other mammals and are hence likely to avoid swimming if possible.