Robust Femur Research Articles

Blood flow rate to the femur of extinct kangaroos implies a higher locomotor intensity compared to living hopping macropods

The stocky skeletons and post-cranial anatomy of many extinct kangaroos indicate that they might have engaged in varied locomotor behaviors, rather than bipedal hopping, as their primary mode of locomotion. This study investigates support for this idea by estimating femoral bone perfusion, which is a correlate of locomotor intensity, in extinct kangaroos compared to living hopping species. Femur blood flow rates can be estimated from the sizes of nutrient foramina on the femur shaft of living and extinct species, without preservation of soft tissue. Estimated femur blood flow rates among the extinct Macropus, Protemnodon and Sthenurinae (Sthenurus, Simosthenurus and Procoptodon) are not significantly different from one another but are significantly greater than in living hopping macropods after accounting for the effect of body mass, consistent with their purportedly different locomotor style. The giant sthenurines have more robust femora than extrapolated from data of living hopping macropods, possibly due to the larger sthenurines requiring relatively stronger leg bones to support their heavier body weights, especially if loaded onto a single limb during bipedal striding.

The Missing Late Pleistocene Ostrich Femur from Zhoukoudian (China): New Information Provided by a Rediscovered Old Cast

A complete ostrich femur from the Late Pleistocene deposits of the Upper Cave at Zhoukoudian (China) was referred by Shaw to Struthio anderssoni in the 1930s, but its present whereabouts are unknown. A good quality plaster cast of the missing specimen has been found in the collections of the Natural History Museum (London). This cast provides interesting information about the morphology of this large ostrich femur, which had previously been only summarily described and not illustrated. Although smaller than the femora of the Early Pleistocene giant ostrich Pachystruthio, the robust femur from Zhoukoudian shows morphological similarities with them, and it is suggested that ‘Struthio’ anderssoni should be placed in the genus Pachystruthio. The importance of old palaeontological casts is emphasized, as well as the need to preserve and curate them properly.

Body mass estimates from postcranial skeletons and implication for positional behavior in Nacholapithecus kerioi: Evolutionary scenarios of modern apes.

This study reported the body mass (BM) estimates of the Middle Miocene fossil hominoid Nacholapithecus kerioi from Africa. The average BM estimates from all forelimb and hindlimb skeletal elements was 22.7 kg, which is slightly higher than the previously reported estimate of ~22 kg. This study revealed that Nacholapithecus has a unique body proportion with an enlarged forelimb relative to a smaller hindlimb, suggesting an antipronograde posture/locomotion, which may be related to the long clavicle, robust ribs, and some hominoid-like vertebral morphology. Because the BM of Nacholapithecus in this study was estimated to be below 30 kg, Nacholapithecus probably did not have relatively shorter and robust femora, which may result from other mechanical constraints, as seen in extant African hominoids. The BM estimate of Nacholapithecus suggests that full substantial modifications of the trunk and forelimb anatomy for risk avoidance and foraging efficiency, as seen in extant great apes, would not be expected in Nacholapithecus. Because larger monkeys are less arboreal (e.g., Mandrillus sphinx or Papio spp.), and the maximum BM among extant constant arboreal cercopithecoids is ~24 kg (male Nasalis larvatus), Nacholapithecus would be a constant arboreal primate. Although caution should be applied because of targeting only males in this study, arboreal quadrupedalism with upright posture and occasional antipronograde locomotion (e.g., climbing, chambering, descending, arm-swing, and sway) using the powerful grasping capacity of the hand and foot may be assumed for positional behavior of Nacholapithecus.

The first discovery of in situ Pelophylax pueyoi (Amphibia: Anura) from the Late Miocene of Libros Konservat-Lagerstätte (Teruel, Spain)

ABSTRACT For the first time, a specimen of Pelophylax pueyoi Navás, 1922a has been recovered in situ, about 600 meters from the main entrance inside the Libros II mine (Teruel, Spain). The fossil comes from the Libros Gypsum Unit (Late Miocene), and represents a nearly complete articulated adult frog exposed in ventral view, with partial preservation of soft tissues. The only apomorphic character diagnosing the family Ranidae is the presence of cylindrical sacral diapophyses. Other characteristics congruent with that of the family Ranidae can be observed: sphenethmoid fused medially, large and oval orbital fossa, well-developed pterygoids without alar expansion and the inner ramus clearly shorter than the posterolateral ramus, transverse processes of presacral vertebrae and sacral diapophyses of subequal length, sacrum unfused with the urostyle, probable bicondylar sacro-urostylar articulation, transverse processes of the urostyle lacking, absence of ribs, and elongated hind limb. An apomorphic combination of features of the genus Pelophylax can also be recognized, such as the more open sacral diapophysis, and less elongated and more robust femora than usually observed in the genus Rana. Finally, some discrete characteristics permit establishing a close relationship with P. pueyoi, such as the large size of the fossil, the wide proportion of the skull, and the somewhat trapezoidal distal contour of the V4 transverse processes. The carpus of P. pueyoi is described here for the first time, suggesting a similar condition of the adult carpal morphology found in most Neobatrachian species. Presence or absence of a praepollex in P. pueyoi still cannot be resolved.

Fast and robust femur segmentation from computed tomography images for patient-specific hip fracture risk screening

ABSTRACT Osteoporosis is a common bone disease that increases the risk of bone fracture. Hip-fracture risk screening methods based on flnite element analysis depend on segmented computed tomography (CT) images; however, current femur segmentation methods require manual delineations of large data sets. Here we propose a deep neural network for fully automated, accurate, and fast segmentation of the proximal femur from CT. Evaluation on a set of 1147 proximal femurs with ground truth segmentations demonstrates that our method is apt for hip-fracture risk screening, bringing us one step closer to a clinically viable option for screening at-risk patients for hip-fracture susceptibility.

Femoral specializations to locomotor habits in early archosauriforms.

The evolutionary history of archosaurs and their closest relatives is characterized by a wide diversity of locomotor modes, which has even been suggested as a pivotal aspect underlying the evolutionary success of dinosaurs vs. pseudosuchians across the Triassic–Jurassic transition. This locomotor diversity (e.g., more sprawling/erect; crouched/upright; quadrupedal/bipedal) led to several morphofunctional specializations of archosauriform limb bones that have been studied qualitatively as well as quantitatively through various linear morphometric studies. However, differences in locomotor habits have never been studied across the Triassic–Jurassic transition using 3D geometric morphometrics, which can relate how morphological features vary according to biological factors such as locomotor habit and body mass. Herein, we investigate morphological variation across a dataset of 72 femora from 36 different species of archosauriforms. First, we identify femoral head rotation, distal slope of the fourth trochanter, femoral curvature, and the angle between the lateral condyle and crista tibiofibularis as the main features varying between bipedal and quadrupedal taxa, all of these traits having a stronger locomotor signal than the lesser trochanter's proximal extent. We show a significant association between locomotor mode and phylogeny, but with the locomotor signal being stronger than the phylogenetic signal. This enables us to predict locomotor modes of some of the more ambiguous early archosauriforms without relying on the relationships between hindlimb and forelimb linear bone dimensions as in prior studies. Second, we highlight that the most important morphological variation is linked to the increase of body size, which impacts the width of the epiphyses and the roundness and proximodistal position of the fourth trochanter. Furthermore, we show that bipedal and quadrupedal archosauriforms have different allometric trajectories along the morphological variation in relation to body size. Finally, we demonstrate a covariation between locomotor mode and body size, with variations in femoral bowing (anteroposterior curvature) being more distinct among robust femora than gracile ones. We also identify a decoupling in fourth trochanter variation between locomotor mode (symmetrical to semi‐pendant) and body size (sharp to rounded). Our results indicate a similar level of morphological disparity linked to a clear convergence in femoral robusticity between the two clades of archosauriforms (Pseudosuchia and Avemetatarsalia), emphasizing the importance of accounting for body size when studying their evolutionary history, as well as when studying the functional morphology of appendicular features. Determining how early archosauriform skeletal features were impacted by locomotor habits and body size also enables us to discuss the potential homoplasy of some phylogenetic characters used previously in cladistic analyses as well as when bipedalism evolved in the avemetatarsalian lineage. This study illuminates how the evolution of femoral morphology in early archosauriforms was functionally constrained by locomotor habit and body size, which should aid ongoing discussions about the early evolution of dinosaurs and the nature of their evolutionary “success” over pseudosuchians.

Convergent morphological responses to loss of flight in rails (Aves: Rallidae).

The physiological demands of flight exert strong selection pressure on avian morphology and so it is to be expected that the evolutionary loss of flight capacity would involve profound changes in traits. Here, we investigate morphological consequences of flightlessness in a bird family where the condition has evolved repeatedly. The Rallidae include more than 130 recognized species of which over 30 are flightless. Morphological and molecular phylogenetic data were used here to compare species with and without the ability to fly in order to determine major phenotypic effects of the transition from flighted to flightless. We find statistical support for similar morphological response among unrelated flightless lineages, characterized by a shift in energy allocation from the forelimbs to the hindlimbs. Indeed, flightless birds exhibit smaller sterna and wings than flighted taxa in the same family along with wider pelves and more robust femora. Phylogenetic signal tests demonstrate that those differences are independent of phylogeny and instead demonstrate convergent morphological adaptation associated with a walking ecology. We found too that morphological variation was greater among flightless rails than flighted ones, suggesting that relaxation of physiological demands during the transition to flightlessness frees morphological traits to evolve in response to more varied ecological opportunities.

Comparative morphology and scaling of the femur in yangochiropteran bats.

Better known by their remarkable forelimb morphology, bats are also unique among mammals with respect to their hindlimbs. Their legs are rotated through 180°, generally reduced in size, and in some extant taxa particular bones (e.g. fibula) can even be absent. The femur is the main leg bone, but to date few bat studies have considered its morphology in detail, none in a wide-scale comparative study. Yangochiroptera is the largest bat taxon, spans nearly three orders of magnitude in body mass, and is highly diverse both in ecology and behavior, representing a good model for comparative analyses. Here, we describe the anatomy of the femur in a large sample of yangochiropteran bats (125 species, 70 genera, and 12 families), and explore major trends of morphological variation and scaling patterns in this bone. We used 13 categorical characters in the anatomical description and five linear dimensions in the quantitative analyses. Based on the categorical data, each family studied here was diagnosed, and those from the Neotropical region were included in an identification key. From the phylogenetic principal component analysis (pPCA) we showed that, in addition to size, major axes of variation in bat femur are related to robusticity and head morphology, features that are clearly distinct among some families. We also generated a phylomorphospace based on pPCA scores, highlighting convergences in femur shape. Molossidae, Mystacinidae, and Desmodontinae were grouped based on their greater robusticity, a pattern that was also recovered from categorical data. In these families, we found anatomical features (e.g. presence of tubercles and posterior ridges on the greater trochanter, long or medially/distally displaced lateral ridges on the shaft) that are well-known from their functional link with quadrupedal locomotion. Using phylogenetic regressions, we found out that compared with body mass, femur length scaled with negative allometry, as expected, but that femur width scaled isometrically, counter to expectations. As a result, robusticity index (the ratio of width to length), scaled with positive allometry - larger bats tended to have more robust hindlimbs. At species level, our most remarkable finding was related to Myotis simus, which presented the most robust femur (for its size) among yangochiropterans. Our results reinforce the informative potential of the chiropteran femur from both taxonomic and functional perspectives. Furthermore, the allometric trends seen in this bone may help understand the strategies adopted by flying vertebrates to deal with the high energetic cost of flight and, at the same time, evolve diversified foraging behaviors.

A new transitional therizinosaurian theropod from the Early Cretaceous Jehol Biota of China

Therizinosaurian theropods evolved many highly specialized osteological features in association with their bulky proportions, which were unusual in the context of the generally gracile Theropoda. Here we report a new therizinosaur, Lingyuanosaurus sihedangensis gen. et sp. nov., based on a specimen recovered from the Lower Cretaceous Jehol Group of Lingyuan, Liaoning Province, China, which displays a combination of plesiomorphic and derived features. Most notably, the specimen is characterized by posterior dorsal vertebrae with a complex and unusual laminar structure; an ilium with a highly dorsoventrally expanded preacetabular process showing only slight lateral flaring of the ventral margin, a strongly anterodorsally inclined iliac blade, a small postacetabular process with a strongly concave dorsal margin, and a relatively robust pubic peduncle with a posteroventrally facing distal articular surface; a straight and robust femur with a small lesser trochanter; and a tibia that is longer than the femur. Phylogenetic analysis places Lingyuanosaurus in an intermediate position within Therizinosauria, i.e., between the early-branching therizinosaurs such as Falcarius, Jianchangosaurus, and Beipiaosaurus and the late-branching ones such as Alxasaurus and Therizinosaurus. This new therizinosaur sheds additional light on the evolution of major therizinosaurian characteristics, including particularly the distinctive pelvic girdle and hindlimb morphology seen in this group.

Getting into Shape: Limb Bone Strength in Perinatal Lemur catta and Propithecus coquereli.

Functional studies of skeletal anatomy are predicated on the fundamental assumption that form will follow function. For instance, previous studies have shown that the femora of specialized leaping primates are more robust than those of more generalized primate quadrupeds. Are such differences solely a plastic response to differential loading patterns during postnatal life, or might they also reflect more canalized developmental mechanisms present at birth? Here, we show that perinatal Lemur catta, an arboreal/terrestrial quadruped, have less robust femora than perinatal Propithecus coquereli, a closely related species specialized for vertical clinging and leaping (a highly unusual locomotor mode in which the hindlimbs are used to launch the animal between vertical tree trunks). These results suggest that functional differences in long bone cross-sectional dimensions are manifest at birth, belying simple interpretations of adult postcranial form as a direct record of loading patterns during postnatal life. Despite these significant differences in bone robusticity, we find that hindlimb bone mineralization, material properties, and measures of whole-bone strength generally overlap in perinatal L. catta and P. coquereli, indicating little differentiation in postcranial maturity at birth despite known differences in the pace of craniodental development between the species. In a broader perspective, our results likely reflect evolution acting during prenatal ontogeny. Even though primates are notable for relatively prolonged gestation and postnatal parental care, neonates are not buffered from selection, perhaps especially in the unpredictable and volatile environment of Madagascar. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 303:250-264, 2020. © 2018 American Association for Anatomy.

Global deletion of Panx3 produces multiple phenotypic effects in mouse humeri and femora.

Pannexins form single-membrane channels that allow passage of small molecules between the intracellular and extracellular compartments. Of the three pannexin family members, Pannexin3 (Panx3) is the least studied but is highly expressed in skeletal tissues and is thought to play a role in the regulation of chondrocyte and osteoblast proliferation and differentiation. The purpose of our study is to closely examine the invivo effects of Panx3 ablation on long bone morphology using micro-computed tomography. Using Panx3 knockout (KO) and wildtype (WT) adult mice, we measured and compared aspects of phenotypic shape, bone mineral density (BMD), cross-sectional geometric properties of right femora and humeri, and lean mass. We found that KO mice have absolutely and relatively shorter diaphyseal shafts compared with WT mice, and relatively larger areas of muscle attachment sites. No differences in BMD or lean mass were found between WT and KO mice. Interestingly, KO mice had more robust femora and humeri compared with WT mice when assessed in cross-section at the midshaft. Our results clearly show that Panx3 ablation produces phenotypic effects in mouse femora and humeri, and support the premise that Panx3 has a role in regulating long bone growth and development.

The interindividual variation in femoral neck width is associated with the acquisition of predictable sets of morphological and tissue-quality traits and differential bone loss patterns

A better understanding of femoral neck structure and age-related bone loss will benefit research aimed at reducing fracture risk. We used the natural variation in robustness (bone width relative to length) to analyze how adaptive processes covary traits in association with robustness, and whether the variation in robustness affects age-related bone loss patterns. Femoral necks from 49 female cadavers (29-93 years of age) were evaluated for morphological and tissue-level traits using radiography, peripheral quantitative computed tomography, micro-computed tomography, and ash-content analysis. Femoral neck robustness was normally distributed and varied widely with a coefficient of variation of 14.9%. Age-adjusted partial regression analysis revealed significant negative correlations (p < 0.05) between robustness and relative cortical area, cortical tissue-mineral density (Ct.TMD), and trabecular bone mineral density (Ma.BMD). Path analysis confirmed these results showing that a one standard deviation (SD) increase in robustness was associated with a 0.70 SD decrease in RCA, 0.47 SD decrease in Ct.TMD, and 0.43 SD decrease in Ma.BMD. Significantly different bone loss patterns were observed when comparing the most slender and most robust tertiles. Robust femora showed significant negative correlations with age for cortical area (R(2) = 0.29, p < 0.03), Ma.BMD (R(2) = 0.34, p < 0.01), and Ct.TMD (R(2) = 0.4, p < 0.003). However, slender femora did not show these age-related changes (R(2) < 0.09, p > 0.2). The results indicated that slender femora were constructed with a different set of traits compared to robust femora, and that the natural variation in robustness was a determinant of age-related bone loss patterns. Clinical diagnoses and treatments may benefit from a better understanding of these robustness-specific structural and aging patterns.

Robust femur condyle disambiguation on biplanar X-rays

Three-dimensional (3D) reconstruction of the skeleton from biplanar X-rays relies on scarce information digitalised by an operator on both frontal and lateral radiographs. In clinical routine, difficulties occur for non-skilled operators to discriminate the medial from the lateral femur condyle on the lateral view. Our study proposes an algorithm able to detect automatically a possible inversion of the two condyles by the operator at an early stage of the reconstruction process. It relies on the computation of two 3D femur surfaces, one directly from the operator digitalisation and the other from the same digitalisation with medial and lateral condyles automatically swapped. Pairs of virtual biplanar X-rays are computed for both reconstructions and the closest pair to the original X-rays is selected on the basis of similarity measures, pointing the correct 3D surface. The algorithm shows a success rate higher than 85% for both asymptomatic and pathological femurs whatever the initial condyle digitalisation of the operator, bringing automatically non-skilled operators acting in clinical routine to the level of skilled operators. This study validates moreover the proof-of-concept of automatic shape adjustments of a 3D surface on the basis of similarity measures in the process of 3D reconstruction from biplanar X-rays.

Allometric and Group Differences in the Xenarthran Femur

The aim of this study is to analyze shape variation in the xenarthran femur to gain insights into their behavior and locomotion. Specimens of both Cingulata (armadillos and glyptodonts) and Pilosa (anteaters and sloths) were studied and within each group body mass varies by several orders of magnitude. The main focus of the analysis was allometric variation in femoral shape in the three groups studied, armadillos, glyptodonts, and pilosans. Three dimensional coordinates were recorded for 40 homologous landmarks on each of 51 xenarthran femurs. The data were analyzed by geometric morphometric methods, and form space analysis was used to identify the allometric variation in each of the three groups. Across all groups, larger specimens tended to have larger articular surfaces, more robust femora generally, and the shape of the femoral condyles was more suited to extended postures. In addition, in larger specimens the medial condyle was much larger than the lateral condyle and the third trochanter was located more distally. The larger armadillo femora had a greater trochanter located considerably proximal to the femoral head and this is thought to improve femoral extension, but in glyptodonts and pilosans the larger specimens had a greater trochanter that was far lateral to the femoral head and this is interpreted as enhancing femoral rotation.

Does Obesity Really Make the Femur Stronger? BMD, Geometry, and Fracture Incidence in the Women's Health Initiative-Observational Study

Heavier individuals have higher hip BMD and more robust femur geometry, but it is unclear whether values vary in proportion with body weight in obesity. We studied the variation of hip BMD and geometry across categories of body mass index (BMI) in a subset of postmenopausal non-Hispanic whites (NHWs) from the Women's Health Initiative Observational Cohort (WHI-OS). The implications on fracture incidence were studied among NHWs in the entire WHI-OS. Baseline DXA scans of hip and total body from 4642 NHW women were divided into BMI (kg/m(2)) categories: underweight (<18.5), healthy weight (18.5-24.9), overweight (25-29.9), and mild (30-34.9), moderate (35-39.9), and extreme obesity (>40). Femur BMD and indices of bone axial (cross-sectional area [CSA]) and bending strength (section modulus [SM]) were extracted from DXA scans using the hip structure analysis (HSA) method and compared among BMI categories after adjustment for height, age, hormone use, diabetes, activity level, femur neck-shaft angle, and neck length. The association between BMI and incident fracture was studied in 78,013 NHWs from the entire WHI-OS over 8.5 +/- 2.6 (SD) yr of follow-up. Fracture incidence (cases/1000 person-years) was compared among BMI categories for hip alone, central body (hip, pelvis, spine, ribs, and shoulder girdle), upper extremity (humerus and distal), and lower extremity (femur shaft and distal but not hip). Femur BMD, CSA, and SM were larger in women with higher BMI, but values scaled in proportion to lean and not to fat or total body mass. Women with highest BMI reported more falls in the 12 mo before enrollment, more prevalent fractures, and had lower measures of physical activity and function. Incidence of hip fractures and all central body fractures declined with BMI. Lower extremity fractures distal to the hip trended upward, and upper extremity incidence was independent of BMI. BMD, CSA, and SM vary in proportion to total body lean mass, supporting the view that bones adapt to prevalent muscle loads. Because lean mass is a progressively smaller fraction of total mass in obesity, femur BMD, CSA, and SM decline relative to body weight in higher BMI categories. Traumatic forces increase with body weight, but fracture rates at the hip and central body were less frequent with increasing BMI, possibly because of greater soft tissue padding. There was no evident protective effect in fracture rates at less padded distal extremity sites. Upper extremity fractures showed no variation with BMI, and lower extremity fracture rates were higher only in the overweight (BMI = 25-29.9 kg/m(2)).

A new giant ground bird from the Upper Cretaceous of southern France

New discoveries afford a much better knowledge of the large birds previously reported from the Upper Cretaceous of southern France on the basis of a single synsacrum fragment. An incomplete pelvis consisting of the synsacrum with part of the ilia is used to define a new taxon of bird, Gargantuavis philoinos , to which a femur is also referred. This bird is characterized by its very large size (comparable to that of an ostrich), a broad pelvis with an anteriorly placed acetabulum, and a short robust femur. The occurrence of this very large, flightless, bird in the Late Cretaceous shows that, contrary to widespread opinion, the evolution of large terrestrial birds was not simply the result of adaptation during the Cenozoic to ecological niches left empty by the extinction of the dinosaurs. It also strongly suggests that some of the large fossil eggs from the Upper Cretaceous of southern France were laid by large ground birds rather than dinosaurs.

Long bone cross-sectional dimensions, locomotor adaptations and body size in prosimian primates

The cross-sectional geometry of primate long bones appears to be closely related to body size and locomotor behavior. This study investigates this aspect of skeletal design in the humerus and femur of living prosimian primates, a group characterized by a diversity of locomotor modes and a wide range of body sizes. The geometrical variables were collected from biplanar radiographs using formulae for hollow ellipses to approximate cortical area, wall thickness, second and polar moments of area, and section modulus. Indices of strength in static compression and bending were also derived from these data, limb length and body mass. For prosimians as a group, all cross-sectional geometrical properties scale in a positive allometric fashion with body mass (although some of the humeral values have confidence limits that include isometry). Despite positive allometry, measures of strength in static loading decrease as a function of body size and imply that alterations in body posture and locomotion are necessary in order to maintain stress/strain similarity. Within prosimians, leapers tend to have more robust femora (with elongated anteroposterior axes) than non-saltatory species of the same size; no such distinctions exist for the humerus (but Daubentonia exhibits an unusually strong humerus). Slow climbing lorisines, although exposed to only moderate locomotor forces, are not at the low end of absolute and relative bone strength variation. Prosimians less than 1·2 kg exhibit considerable variation in cross-sectional geometry, a finding that could complicate attempts to use such data to reconstruct body size in small-bodied fossil prosimians. Measures of wall thickness (K-values) appear to be of limited utility in the functional analysis of skeletal design in extant prosimians. Compound indices of relative strength—although useful in functional allometric analyses—do not reliably differentiate among either locomotor or taxonomic groups. Functional differences among taxa are more obvious in geometrical properties when body size is used as a covariate.

Hind limb scaling in birds and other theropods: Implications for terrestrial locomotion.

An analysis of hind limb skeletal elements of non-avian theropods and ground-dwelling birds was performed to reveal patterns of change in shape and proportion with size. When femora of equal length are compared, birds exhibit a significantly larger midshaft diameter than non-avian theropods. As total limb length increases, avian femora become relatively shorter (negative allometry), while those of non-avian theropods become relatively longer (positive allometry). Avian femoral/tibiotarsal ratios are all below 0.8 and decrease with limb size, whereas ratios of non-avian theropods are well above 0.8 and tend to increase with limb size. In addition, avian femora exhibit a unique diameter/length relationship not seen in other theropod hind limb bones. Several studies have shown that within the avian limb, the short, robust femur resists bending to a far greater degree than the relatively longer, slimmer tibiotarsus. This is to be expected, as analyses of running birds show that the femur is oriented relatively perpendicular to the ground reaction force throughout the stride, which would subject it to high bending moments. When compared to birds, non-avian theropods have relatively long, slender femora that do not seem to be built to withstand the forces associated with such an orientation. Reconstructing all non-avian theropods in avian-like poses (subhorizontal femur, knee well flexed) with avian locomotor kinematics (relatively little hip extension at most speeds) ignores major differences in scaling between these groups of organisms.