Intra-guild competition and its implications for one of the biggest terrestrial predators, Tyrannosaurus rex

Identifying tradeoffs between hunting and scavenging in an ecological context is important for understanding predatory guilds. In the past century, the feeding strategy of one of the largest and best-known terrestrial carnivores, Tyrannosaurus rex, has been the subject of much debate: was it an active predator or an obligate scavenger? Here we look at the feasibility of an adult T. rex being an obligate scavenger in the environmental conditions of Late Cretaceous North America, given the size distributions of sympatric herbivorous dinosaurs and likely competition with more abundant small-bodied theropods. We predict that nearly 50 per cent of herbivores would have been within a 55–85 kg range, and calculate based on expected encounter rates that carcasses from these individuals would have been quickly consumed by smaller theropods. Larger carcasses would have been very rare and heavily competed for, making them an unreliable food source. The potential carcass search rates of smaller theropods are predicted to be 14–60 times that of an adult T. rex. Our results suggest that T. rex and other extremely large carnivorous dinosaurs would have been unable to compete as obligate scavengers and would have primarily hunted large vertebrate prey, similar to many large mammalian carnivores in modern-day ecosystems.     

Vital statistics,absolute abundance and preservation rate of Tyrannosaurus rex

I present a simulation model on vital statistics, absolute abundance (N, total number of individuals that ever lived) and preservation rate (p, minimum number of fossils known divided by N) of Tyrannosaurus rex. It is based on a published age-structured population model that assumes a reptile or bird-like reproduction for T. rex to estimate its age-specific survival rates. My model applies input variables and equations from a recently published model on N and p. This model yielded 2.5 billion T. rex individuals (N) and one fossil per 80 million individuals (p). The average N values calculated by my model were at minimum 27.6% and p values at maximum 361.5% that of a previous model and uncertainties in all output variables were always larger in my model. The equation on output variable ‘population density’ introduced the largest uncertainty to N and p. The output variable ‘generation time’ differed the most between models, but for N and p, the huge size of the input area modelled and geological longevity minimized this difference. Unlike my model, the generation time as well as life expectancies, gross reproduction rates, and reproductive values of individuals calculated from the previous model all strongly contradicted our current understanding of the biology of T. rex and of other theropods. Their values also disagreed with those of large extant reptiles, birds and mammals. All of these shortcomings of the previous model favour the assessment of individual and population characteristics of T. rex and of other extinct species using my model.     

A 3D interactive method for estimating body segmental parameters in animals: application to the turning and running performance of Tyrannosaurus rex

We developed a method based on interactive B-spline solids for estimating and visualizing biomechanically important parameters for animal body segments. Although the method is most useful for assessing the importance of unknowns in extinct animals, such as body contours, muscle bulk, or inertial parameters, it is also useful for non-invasive measurement of segmental dimensions in extant animals. Points measured directly from bodies or skeletons are digitized and visualized on a computer, and then a B-spline solid is fitted to enclose these points, allowing quantification of segment dimensions. The method is computationally fast enough so that software implementations can interactively deform the shape of body segments (by warping the solid) or adjust the shape quantitatively (e.g., expanding the solid boundary by some percentage or a specific distance beyond measured skeletal coordinates). As the shape changes, the resulting changes in segment mass, center of mass (CM), and moments of inertia can be recomputed immediately. Volumes of reduced or increased density can be embedded to represent lungs, bones, or other structures within the body. The method was validated by reconstructing an ostrich body from a fleshed and defleshed carcass and comparing the estimated dimensions to empirically measured values from the original carcass. We then used the method to calculate the segmental masses, centers of mass, and moments of inertia for an adult Tyrannosaurus rex, with measurements taken directly from a complete skeleton. We compare these results to other estimates, using the model to compute the sensitivities of unknown parameter values based upon 30 different combinations of trunk, lung and air sac, and hindlimb dimensions. The conclusion that T. rex was not an exceptionally fast runner remains strongly supported by our models-the main area of ambiguity for estimating running ability seems to be estimating fascicle lengths, not body dimensions. Additionally, the craniad position of the CM in all of our models reinforces the notion that T. rex did not stand or move with extremely columnar, elephantine limbs. It required some flexion in the limbs to stand still, but how much flexion depends directly on where its CM is assumed to lie. Finally we used our model to test an unsolved problem in dinosaur biomechanics: how fast a huge biped like T. rex could turn. Depending on the assumptions, our whole body model integrated with a musculoskeletal model estimates that turning 45 degrees on one leg could be achieved slowly, in about 1-2s.     

Understanding hind limb weight support in chimpanzees with implications for the evolution of primate locomotion

Most quadrupedal mammals support a larger amount of body weight on their forelimbs compared with their hind limbs during locomotion, whereas most primates support more of their body weight on their hind limbs. Increased hind limb weight support is generally interpreted as an adaptation that reduces stress on primates' highly mobile forelimb joints. Thus, increased hind limb weight support was likely vital for the evolution of primate arboreality. Despite its evolutionary importance, the mechanism used by primates to achieve this important kinetic pattern remains unclear. Here, we examine weight support patterns in a sample of chimpanzees (Pan troglodytes) to test the hypothesis that limb position, combined with whole body center of mass position (COM), explains increased hind limb weight support in this taxon. Chimpanzees have a COM midway between their shoulders and hips and walk with a relatively protracted hind limb and a relatively vertical forelimb, averaged over a step. Thus, the limb kinematics of chimpanzees brings their feet closer to the COM than their hands, generating greater hind limb weight support. Comparative data suggest that these same factors likely explain weight support patterns for a broader sample of primates. It remains unclear whether primates use these limb kinematics to increase hind limb weight support, or whether they are byproducts of other gait characteristics. The latter hypothesis raises the intriguing possibility that primate weight support patterns actually evolved as byproducts of other traits, or spandrels, rather than as adaptations to increase forelimb mobility. Am J Phys Anthropol, 2009. © 2008 Wiley‐Liss, Inc.     

A computational model of ameboid deformation and locomotion

Traditional continuum models of ameboid deformation and locomotion are limited by the computational difficulties intrinsic in free boundary conditions. A new model using the immersed boundary method overcomes these difficulties by representing the cell as a force field immersed in fluid domain. The forces can be derived from a direct mechanical interpretation of such cell components as the cell membrane, the actin cortex, and the transmembrane adhesions between the cytoskeleton and the substratum. The numerical cytoskeleton, modeled as a dynamic network of immersed springs, is able to qualitatively model the passive mechanical behavior of a shear-thinning viscoelastic fluid (Bottino 1997). The same network is used to generate active protrusive and contractile forces. When coordinated with the attachment-detachment cycle of the cell's adhesions to the substratum, these forces produce directed locomotion of the model ameba. With this model it is possible to study the effects of altering the numerical parameters upon the motility of the model cell in a manner suggestive of genetic deletion experiments. In the context of this ameboid cell model and its numerical implementation, simulations involving multicellular interaction, detailed internal signaling, and complex substrate geometries are tractable. Received: 5 January 1998 / Revised version: 23 March 1998 / Accepted: 26 March 1998     

The ontogeny of cranial base angulation in humans and chimpanzees and its implications for reconstructing pharyngeal dimensions

This paper examines differences in the processes by which the cranial base flexes in humans and extends in chimpanzees. In addition, we test the extent to which one can use comparisons of cranial base angles in humans and non-human primates to predict vocal tract dimensions. Four internal cranial base angles and one external cranial base angle were measured in a longitudinal sample of Homo sapiens and a cross-sectional sample of Pan troglodytes. These data show that the processes of cranial base angulation differ substantially in these species. While the human cranial base flexes postnatally in a rapid growth trajectory that is complete by two years, the cranial base in P. troglodytes extends postnatally in a more prolonged skeletal growth trajectory. These comparisons also demonstrate that the rate of cranial base angulation is comparable for different measures, but that angles which incorporate different anterior cranial base measurements correlate poorly. We also examined ontogenetic relationships between internal and external cranial base angles and vocal tract growth in humans to test the hypothesis that cranial base angulation influences pharyngeal dimensions and can, therefore, be used to estimate vocal tract proportions in fossil hominids. Our results indicate that internal and external cranial base angles are independent of the horizontal and vertical dimensions of the vocal tract. Instead, a combination of mandibular and palatal landmarks can be used to predict dimensions of the vocal tract in H. sapiens. The developmental contrasts in cranial base angulation between humans and non-human primates may have important implications for testing hypotheses about the relationship between cranial base flexion and other craniofacial dimensions in hominid evolution.     

A computational model of visually guided locomotion in lamprey

This study addresses mechanisms for the generation and selection of visual behaviors in anamniotes. To demonstrate the function of these mechanisms, we have constructed an experimental platform where a simulated animal swims around in a virtual environment containing visually detectable objects. The simulated animal moves as a result of simulated mechanical forces between the water and its body. The undulations of the body are generated by contraction of simulated muscles attached to realistic body components. Muscles are driven by simulated motoneurons within networks of central pattern generators. Reticulospinal neurons, which drive the spinal pattern generators, are in turn driven directly and indirectly by visuomotor centers in the brainstem. The neural networks representing visuomotor centers receive sensory input from a simplified retina. The model also includes major components of the basal ganglia, as these are hypothesized to be key components in behavior selection. We have hypothesized that sensorimotor transformation in tectum and pretectum transforms the place-coded retinal information into rate-coded turning commands in the reticulospinal neurons via a recruitment network mimicking the layered structure of tectal areas. Via engagement of the basal ganglia, the system proves to be capable of selecting among several possible responses, even if exposed to conflicting stimuli. The anatomically based structure of the control system makes it possible to disconnect different neural components, yielding concrete predictions of how animals with corresponding lesions would behave. The model confirms that the neural networks identified in the lamprey are capable of responding appropriately to simple, multiple, and conflicting stimuli.     

Kinematics and ontogeny of locomotion in monkeys and human babies

Early ontogenetic stages are often assumed to reflect or to be similar to past phylogenetic stages within the evolution of man. Therefore, as a first step, the quadrupedal crawling locomotion of human children was analysed and compared to the quadrupedal walk of Macaca fascicularis. The movements of the human child were not only more irregular, they differed from the walk of the monkey mainly through extraordinarily short swing phases, and also through strong scoliotic movements of the spine. There is a compulsory synchronisation in the hip and knee joint movements of the human crawling baby. We conclude that human crawling may be a behavioural recapitulation of a quadrupedal evolutionary stage. However, with reference to kinematics, man is not only characterised by his unique, habitually bipedal, upright gait but also by a second, equally unique locomotion, namely crawling, which he assumes for a short phase during his first year of life.--The walking movements of the limbs in toddling infants were mainly characterised by i) rather stiff, abducted arms, which were moved mostly by spine torsions (similar to those of bipedally walking Gorilla) and not as a suspensory pendulum. However, they rather work as levers for the elastic torsion pendulum of the spine. ii) They are also characterised by frequently lacking the minor knee flexion, which occurs at about the heel strike within each stride of the adult human. Besides many other details of the results, foot movements differed from adult ones mainly in that the whole plantar surface was placed flat on the ground within a few milliseconds.     

Postnatal temporal bone ontogeny in Pan,Gorilla, and Homo,and the implications for temporal bone ontogeny in Australopithecus afarensis

Assessments of temporal bone morphology have played an important role in taxonomic and phylogenetic evaluations of fossil taxa, and recent three‐dimensional analyses of this region have supported the utility of the temporal bone for testing taxonomic and phylogenetic hypotheses. But while clinical analyses have examined aspects of temporal bone ontogeny in humans, the ontogeny of the temporal bone in non‐human taxa is less well documented. This study examines ontogenetic allometry of the temporal bone in order to address several research questions related to the pattern and trajectory of temporal bone shape change during ontogeny in the African apes and humans. We further apply these data to a preliminary analysis of temporal bone ontogeny in Australopithecus afarensis. Three‐dimensional landmarks were digitized on an ontogenetic series of specimens of Homo sapiens, Pan troglodytes, Pan paniscus, and Gorilla gorilla. Data were analyzed using geometric morphometric methods, and shape changes throughout ontogeny in relation to size were compared. Results of these analyses indicate that, despite broadly similar patterns, African apes and humans show marked differences in development of the mandibular fossa and tympanic portions of the temporal bone. These findings indicate divergent, rather than parallel, postnatal ontogenetic allometric trajectories for temporal bone shape in these taxa. The pattern of temporal bone shape change with size exhibited by A. afarensis showed some affinities to that of humans, but was most similar to extant African apes, particularly Gorilla. Am J Phys Anthropol 151:630–642, 2013. © 2013 Wiley Periodicals, Inc.     

The effect of lower limb length on the energetic cost of locomotion: implications for fossil hominins

The consequences of the relatively short lower limbs characteristic of AL 288-1 have been widely discussed, as have the causes and consequences of the short limbs of Neanderthals. Previous studies of the effect of limb length on the energetic cost of locomotion have reported no relationship; however, limb length could have accounted for as much as 19% of the variation in cost and gone undetected (Steudel and Beattie, 1995; Steudel, 1994, 1996). Kramer (1999) and Kramer and Eck (2000) have recently used a theoretical model to predict the effect of the shorter limbs of early hominids, concluding that the shorter limbs may actually have been energetically advantageous. Here, we took an experimental approach. Twenty-one human subjects, of varying limb lengths, walked on a treadmill at 2.6, 2.8, 3.0 and 3.2m.p.h., while their expired gases were analyzed. The subjects walked for 12 minutes at each speed and their rates of oxygen consumption (VO2) over four minutes were averaged to estimate VO2. We also measured each subject's height, weight and lower limb length. Lean body mass and % fat were determined using dual-energy x-ray absorptiometry. ANCOVA with total VO2 at either speed as the dependent variable and total lean mass, % fat and lower limb length as covariates resulted in all three covariates having a significant positive effect on VO2 at p<0.01. Subjects with relatively longer lower limbs had lower locomotor costs. Thus the short lower limbs characteristic of some hominid taxa would have resulted in more costly locomotion, barring some physiological anomaly. The magnitude of this effect is substantial; Neanderthals are estimated to have had locomotor costs 30% greater than those of contemporary anatomically modern humans. By contrast the increase in lower limb length seen in H. erectus would have mitigated the increase in locomotor costs produced by the increase in body size.     

Relationship between dental development and skeletal growth in modern humans and its implications for interpreting ontogeny in fossil hominins

Dental development and skeletal growth are central aspects used by anthropologists when investigating the ontogeny of a population or species. The interrelatedness of the two phenomena is often assumed to be high, but the nature of their relationship is obscured by the fact that they are both highly dependent upon chronological age. The exact relationship between the tempo of dental development and skeletal growth is unclear even in modern humans, which limits the ability to extrapolate to archaeological or fossil forms. It is clear that the influence of chronological age on these two aspects of ontogeny must be accounted for before examining their relationship to one another. This study tests whether dental development and skeletal growth are conditionally independent given age using known‐age modern human skeletal samples and proportional odds logistic regression. The results suggest that dental development and skeletal growth are moderately correlated and thus not conditionally independent given age. That is, individuals that are dentally advanced relative to their peers also tend to be skeletally advanced. However, this relationship is moderate at best, so dental development does not appear to be a highly reliable proxy for skeletal growth, or vice versa, in modern humans. These findings have implications for the reconstruction of ontogeny and life history of fossil hominin taxa, since the pace of dental development is often used as a life history proxy. Implications of this study suggest that the proposed accelerated dental development in Pleistocene hominins was not necessarily accompanied by faster skeletal growth. Am J Phys Anthropol, 2013. © 2012 Wiley Periodicals, Inc.     

Postcranial functional morphology of Morotopithecus bishopi, with implications for the evolution of modern ape locomotion

The large-bodied hominoid from Moroto, Uganda has until recently been known only from proconsulid like craniodental remains and some vertebrae with modern ape like features. The discovery of two partial femora and the glenoid portion of a scapula demonstrates that the functional anatomy of Morotopithecus differed markedly from other early and middle Miocene hominoids. Previous studies have consistently associated the vertebral remains with a short, stiff back and with orthograde postures. Although the proximal femur more closely resembles the femora of monkeys than of apes and suggests a moderate degree of hip abduction, the distal femur resembles those of extant large bodied apes and suggests a varied loading regime and an arboreal repertoire that may have included substantial vertical climbing. The femoral shaft displays uniformly thick cortical bone, beyond the range of thickness seen in extant primates, and signifies higher axial loading than is typical of most extant primates. The glenoid fossa is broad and uniformly curved as in extant suspensory primates. Overall, Morotopithecus is reconstructed as an arboreal species that probably relied on forelimb-dominated, deliberate and vertical climbing, suspension and quadrupedalism. Morotopithecus thus marks the first appearance of certain aspects of the modern hominoid body plan by at least 20 Ma. If the suspensory and orthograde adaptations linking Morotopithecus to extant apes are synapomorphies, Morotopithecus may be the only well-documented African Miocene hominoid with a close relationship to living apes and humans.     

Bipedal locomotion: effects of speed, size and limb posture in birds and humans

Seven species of ground-dwelling birds (body mass range: 0.045-90 kg) were filmed while walking and running on a treadmill. High-speed light films were also taken of humans to compare kinematic patterns of avian with human bipedalism. Consistent patterns of stride frequency, stride length, step length, duty factor and limb excursion were observed in all species, with most of the variation among species being due to differences in body size. In general, smaller bipeds have higher stride frequencies (α M ^−0.18), shorter stride lengths (α M ^0.38) and more limited ranges of speed within each gait than large bipeds. After normalizing for size (based on Froude number, after Alexander, 1977), remaining kinematic variation is largely due to interspecific differences in posture and relative limb segment lengths. For their size, smaller bipeds have greater step lengths, limb excursion angles and duty factors than large bipeds because of their more crouched posture and greater effective limb length. The most notable differences in limb kinematics between birds and humans occur at the walk-run transition and are maintained as running speed increases. Change of gait is smooth and difficult to discern in birds, but distinct in humans, involving abrupt decreases in step length and duty factor (time of contact) and a corresponding increase in limb swing time. These differences appear to reflect a spring-like run that is stiff in humans (favouring elastic energy recovery) but more compliant in birds (increasing time of ground contact). Differences between birds and humans in balance of the body's centre of mass not only affect femoral orientation and motion, but also affect pattern of limb excursion with speed.