The respiratory system influences flight mechanics in soaring birds.

The subpectoral diverticulum (SPD) is an extension of the respiratory system in birds that is located between the primary muscles responsible for flapping the wing1,2. Here we survey the pulmonary apparatus in 68 avian species, and show that the SPD was present in virtually all of the soaring taxa investigated but absent in non-soarers. We find that this structure evolved independently with soaring flight at least seven times, which indicates that the diverticulum might have a functional and adaptive relationship with this flight style. Using the soaring hawks Buteo jamaicensis and Buteo swainsoni as models, we show that the SPD is not integral for ventilation, that an inflated SPD can increase the moment arm of cranial parts of the pectoralis, and that pectoralis muscle fascicles are significantly shorter in soaring hawks than in non-soaring birds. This coupling of an SPD-mediated increase in pectoralis leverage with force-specialized muscle architecture produces a pneumatic system that is adapted for the isometric contractile conditions expected in soaring flight. The discovery of a mechanical role for the respiratory system in avian locomotion underscores the functional complexity and heterogeneity of this organ system, and suggests that pulmonary diverticula are likely to have other undiscovered secondary functions. These data provide a mechanistic explanation for the repeated appearance of the SPD in soaring lineages and show that the respiratory system can be co-opted to provide biomechanical solutions to the challenges of flight and thereby influence the evolution of avian volancy.

Evaluation of a cadaveric wrist motion simulator using marker-based X-ray reconstruction of moving morphology.

Surgical intervention is a common option for the treatment of wrist joint arthritis and traumatic wrist injury. Whether this surgery is arthrodesis or a motion preserving procedure such as arthroplasty, wrist joint biomechanics are inevitably altered. To evaluate effects of surgery on parameters such as range of motion, efficiency and carpal kinematics, repeatable and controlled motion of cadaveric specimens is required. This study describes the development of a device that enables cadaveric wrist motion to be simulated before and after motion preserving surgery in a highly controlled manner. The simulator achieves joint motion through the application of predetermined displacements to the five major tendons of the wrist, and records tendon forces. A pilot experiment using six wrists aimed to evaluate its accuracy and reproducibility. Biplanar X-ray videoradiography (BPVR) and X-Ray Reconstruction of Moving Morphology (XROMM) were used to measure overall wrist angles before and after total wrist arthroplasty. The simulator was able to produce flexion, extension, radioulnar deviation, dart thrower's motion and circumduction within previously reported functional ranges of motion. Pre- and post-surgical wrist angles did not significantly differ. Intra-specimen motion trials were repeatable; root mean square errors between individual trials and average wrist angle and tendon force profiles were below 1° and 2 N respectively. Inter-specimen variation was higher, likely due to anatomical variation and lack of wrist position feedback. In conclusion, combining repeatable intra-specimen cadaveric motion simulation with BPVR and XROMM can be used to determine potential effects of motion preserving surgeries on wrist range of motion and biomechanics.

Regional variation of the cortical and trabecular bone material properties in the rabbit skull.

The material properties of some bones are known to vary with anatomical location, orientation and position within the bone (e.g., cortical and trabecular bone). Details of the heterogeneity and anisotropy of bone is an important consideration for biomechanical studies that apply techniques such as finite element analysis, as the outcomes will be influenced by the choice of material properties used. Datasets detailing the regional variation of material properties in the bones of the skull are sparse, leaving many finite element analyses of skulls no choice but to employ homogeneous, isotropic material properties, often using data from a different species to the one under investigation. Due to the growing significance of investigating the cranial biomechanics of the rabbit in basic science and clinical research, this study used nanoindentation to measure the elastic modulus of cortical and trabecular bone throughout the skull. The elastic moduli of cortical bone measured in the mediolateral and ventrodorsal direction were found to decrease posteriorly through the skull, while it was evenly distributed when measured in the anteroposterior direction. Furthermore, statistical tests showed that the variation of elastic moduli between separate regions (anterior, middle and posterior) of the skull were significantly different in cortical bone, but was not in trabecular bone. Elastic moduli measured in different orthotropic planes were also significantly different, with the moduli measured in the mediolateral direction consistently lower than that measured in either the anteroposterior or ventrodorsal direction. These findings demonstrate the significance of regional and directional variation in cortical bone elastic modulus, and therefore material properties in finite element models of the skull, particularly those of the rabbit, should consider the heterogeneous and orthotropic properties of skull bone when possible.

GenPup-M: A novel validated owner-reported clinical metrology instrument for detecting early mobility changes in dogs.

OBJECTIVE: To use a previously validated veterinary clinical examination sheet, Liverpool Osteoarthritis in Dogs (LOAD) questionnaire, combined with kinetic and kinematic gait analysis in dogs with/without mobility problems to demonstrate the capacity of a novel clinical metrology instrument ("GenPup-M") to detect canine mobility impairments. DESIGN: Quantitative study. ANIMALS: 62 dogs (31 with mobility impairments and 31 without mobility impairments). PROCEDURE: The dogs' clinical history was obtained from owners and all dogs underwent a validated orthopaedic clinical examination. Mobility impairments were diagnosed in the mobility impaired group based on clinical history and orthopaedic examination. Owners were asked to complete GenPup-M along with a previously validated mobility questionnaire (Liverpool Osteoarthritis in Dogs (LOAD)) to identify construct validity. As a test of criterion validity, the correlation between instrument scores and the overall clinical examination scores, along with force-platform obtained peak vertical forces (PVF) were calculated. GenPup-M underwent internal consistency and factor analysis. Spatiotemporal parameters were calculated for dogs with/without mobility impairments to define the gait differences between these two groups. RESULTS: Principal Component Analysis identified GenPup-M had two components with Eigenvalues >1 ("stiffness/ease of movement" and "willingness to be active/exercise"). Cronbach's α was used to test internal consistency of GenPup-M and was found to be "good" (0.87). There was a strong, positive correlation between GenPup-M and LOAD responses (r2 = 0.69, p<0.001) highlighting construct validity. Criterion validity was also shown when comparing GenPup-M to clinical examination scores (r2 = 0.74, p<0.001) and PVF (r2 = 0.43, p<0.001). Quantitative canine gait analysis showed that there were statistically significant differences between peak vertical forces (PVF) of mobility impaired and non-mobility impaired dogs (p<0.05). Analyses of PVF showed that non-mobility impaired dogs more evenly distributed their weight across all thoracic and pelvic limbs when compared to mobility impaired dogs. There were also consistent findings that mobility impaired dogs moved slower than non-mobility impaired dogs. CONCLUSION AND CLINICAL RELEVANCE: GenPup-M is a clinical metrology instrument (CMI) that can be completed by dog owners to detect all mobility impairments, including those that are early in onset, indicating the versatility of GenPup-M to assess dogs with and without mobility impairments. Results of the study found that GenPup-M positively correlated with all three objective measures of canine mobility and consequently showed criterion and construct validity. Owner-reported CMIs such as GenPup-M allow non-invasive scoring systems which veterinary surgeons and owners can use to allow communication and longitudinal assessment of a dog's mobility. It is anticipated that GenPup-M will be used by owners at yearly vaccinations/health checks, allowing identification of any subtle mobility changes, and enabling early intervention.

The Functional and Anatomical Impacts of Healthy Muscle Ageing.

Even "healthy" muscle ageing is often associated with substantial changes in muscle form and function and can lead to increased injury risks and significant negative impacts on quality of life. However, the impacts of healthy muscle ageing on the fibre architecture and microstructure of different muscles and muscle groups throughout the lower limb, and how these are related to their functional capabilities, are not fully understood. Here, a previously established framework of magnetic resonance and diffusion tensor imaging was used to measure the muscle volumes, intramuscular fat, fibre lengths and physiological cross-sectional areas of 12 lower limb muscles in a cohort of healthily aged individuals, which were compared to the same data from a young population. Maximum muscle forces were also measured from an isokinetic dynamometer. The more substantial interpopulation differences in architecture and functional performance were located within the knee extensor muscles, while the aged muscles were also more heterogeneous in muscle fibre type and atrophy. The relationships between architecture and muscle strength were also more significant in the knee extensors compared to other functional groups. These data highlight the importance of the knee extensors as a potential focus for interventions to negate the impacts of muscle ageing.

Decoupling body shape and mass distribution in birds and their dinosaurian ancestors.

It is accepted that non-avian theropod dinosaurs, with their long muscular tails and small forelimbs, had a centre-of-mass close to the hip, while extant birds, with their reduced tails and enlarged wings have their mass centred more cranially. Transition between these states is considered crucial to two key innovations in the avian locomotor system: crouched bipedalism and powered flight. Here we use image-based models to challenge this dichotomy. Rather than a phylogenetic distinction between 'dinosaurian' and 'avian' conditions, we find terrestrial versus volant taxa occupy distinct regions of centre-of-mass morphospace consistent with the disparate demands of terrestrial bipedalism and flight. We track this decoupled evolution of body shape and mass distribution through bird evolution, including the origin of centre-of-mass positions more advantageous for flight and major reversions coincident with terrestriality. We recover modularity in the evolution of limb proportions and centre-of-mass that suggests fully crouched bipedalism evolved after powered flight.

Convergent evolution of quadrupedality in ornithischian dinosaurs was achieved through disparate forelimb muscle mechanics.

The secondary evolution of quadrupedality from bipedal ancestry is a rare evolutionary transition in tetrapods yet occurred convergently at least three times within ornithischian dinosaurs. Despite convergently evolving quadrupedal gait, ornithischians exhibited variable anatomy, particularly in the forelimbs, which underwent a major functional change from assisting in foraging and feeding in bipeds to becoming principal weight-bearing components of the locomotor system in quadrupeds. Here, we use three-dimensional multi-body dynamics models to demonstrate quantitatively that different quadrupedal ornithischian clades evolved distinct forelimb musculature, particularly around the shoulder. We find that major differences in glenohumeral abduction-adduction and long axis rotation muscle leverages were key drivers of mechanical disparity, thereby refuting previous hypotheses about functional convergence in major clades. Elbow muscle leverages were also disparate across the major ornithischian lineages, although high elbow extension muscle leverages were convergent between most quadrupeds. Unlike in ornithischian hind limbs, where differences are more closely tied to functional similarity than phylogenetic relatedness, mechanical disparity in ornithischian forelimbs appears to have been shaped primarily by phylogenetic constraints. Differences in ancestral bipedal taxa within each clade may have resulted in disparate ecomorphological constraints on the evolutionary pathways driving divergence in their quadrupedal descendants.

The association between muscle architecture and muscle spindle abundance.

Across the human body, skeletal muscles have a broad range of biomechanical roles that employ complex proprioceptive control strategies to successfully execute a desired movement. This information is derived from peripherally located sensory apparatus, the muscle spindle and Golgi tendon organs. The abundance of these sensory organs, particularly muscle spindles, is known to differ considerably across individual muscles. Here we present a comprehensive data set of 119 muscles across the human body including architectural properties (muscle fibre length, mass, pennation angle and physiological cross-sectional area) and statistically test their relationships with absolute spindle number and relative spindle abundance (the residual value of the linear regression of the log-transformed spindle number and muscle mass). These data highlight a significant positive relationship between muscle spindle number and fibre length, emphasising the importance of fibre length as an input into the central nervous system. However, there appears to be no relationship between muscles architecturally optimised to function as displacement specialists and their provision of muscle spindles. Additionally, while there appears to be regional differences in muscle spindle abundance, independent of muscle mass and fibre length, our data provide no support for the hypothesis that muscle spindle abundance is related to anatomical specialisation.

Why does the metabolic cost of walking increase on compliant substrates?

Walking on compliant substrates requires more energy than walking on hard substrates but the biomechanical factors that contribute to this increase are debated. Previous studies suggest various causative mechanical factors, including disruption to pendular energy recovery, increased muscle work, decreased muscle efficiency and increased gait variability. We test each of these hypotheses simultaneously by collecting a large kinematic and kinetic dataset of human walking on foams of differing thickness. This allowed us to systematically characterize changes in gait with substrate compliance, and, by combining data with mechanical substrate testing, drive the very first subject-specific computer simulations of human locomotion on compliant substrates to estimate the internal kinetic demands on the musculoskeletal system. Negative changes to pendular energy exchange or ankle mechanics are not supported by our analyses. Instead we find that the mechanistic causes of increased energetic costs on compliant substrates are more complex than captured by any single previous hypothesis. We present a model in which elevated activity and mechanical work by muscles crossing the hip and knee are required to support the changes in joint (greater excursion and maximum flexion) and spatio-temporal kinematics (longer stride lengths, stride times and stance times, and duty factors) on compliant substrates.

Ligament mechanics of ageing and osteoarthritic human knees.

Knee joint ligaments provide stability to the joint by preventing excessive movement. There has been no systematic effort to study the effect of OA and ageing on the mechanical properties of the four major human knee ligaments. This study aims to collate data on the material properties of the anterior (ACL) and posterior (PCL) cruciate ligaments, medial (MCL) and lateral (LCL) collateral ligaments. Bone-ligament-bone specimens from twelve cadaveric human knee joints were extracted for this study. The cadaveric knee joints were previously collected to study ageing and OA on bone and cartilage material properties; therefore, combining our previous bone and cartilage data with the new ligament data from this study will facilitate subject-specific whole-joint modelling studies. The bone-ligament-bone specimens were tested under tensile loading to failure, determining material parameters including yield and ultimate (failure) stress and strain, secant modulus, tangent modulus, and stiffness. There were significant negative correlations between age and ACL yield stress (p = 0.03), ACL failure stress (p = 0.02), PCL secant (p = 0.02) and tangent (p = 0.02) modulus, and LCL stiffness (p = 0.046). Significant negative correlations were also found between OA grades and ACL yield stress (p = 0.02) and strain (p = 0.03), and LCL failure stress (p = 0.048). However, changes in age or OA grade did not show a statistically significant correlation with the MCL tensile parameters. Due to the small sample size, the combined effect of age and the presence of OA could not be statistically derived. This research is the first to report tensile properties of the four major human knee ligaments from a diverse demographic. When combined with our previous findings on bone and cartilage for the same twelve knee cadavers, the current ligament study supports the conceptualisation of OA as a whole-joint disease that impairs the integrity of many peri-articular tissues within the knee. The subject-specific data pool consisting of the material properties of the four major knee ligaments, subchondral and trabecular bones and articular cartilage will advance knee joint finite element models.

Body size, shape and ecology in tetrapods.

Body size and shape play fundamental roles in organismal function and it is expected that animals may possess body proportions that are well-suited to their ecological niche. Tetrapods exhibit a diverse array of body shapes, but to date this diversity in body proportions and its relationship to ecology have not been systematically quantified. Using whole-body skeletal models of 410 extinct and extant tetrapods, we show that allometric relationships vary across individual body segments thereby yielding changes in overall body shape as size increases. However, we also find statistical support for quadratic relationships indicative of differential scaling in small-medium versus large animals. Comparisons of locomotor and dietary groups highlight key differences in body proportions that may mechanistically underlie occupation of major ecological niches. Our results emphasise the pivotal role of body proportions in the broad-scale ecological diversity of tetrapods.

Skeletal muscle function underpins muscle spindle abundance.

Muscle spindle abundance is highly variable within and across species, but we currently lack any clear picture of the mechanistic causes or consequences of this variation. Previous use of spindle abundance as a correlate for muscle function implies a mechanical underpinning to this variation, but these ideas have not been tested. Herein, we use integrated medical imaging and subject-specific musculoskeletal models to investigate the relationship between spindle abundance, muscle architecture and in vivo muscle behaviour in the human locomotor system. These analyses indicate that muscle spindle number is tightly correlated with muscle fascicle length, absolute fascicle length change, velocity of fibre lengthening and active muscle forces during walking. Novel correlations between functional indices and spindle abundance are also recovered, where muscles with a high abundance predominantly function as springs, compared to those with a lower abundance mostly functioning as brakes during walking. These data demonstrate that muscle fibre length, lengthening velocity and fibre force are key physiological signals to the central nervous system and its modulation of locomotion, and that muscle spindle abundance may be tightly correlated to how a muscle generates work. These insights may be combined with neuromechanics and robotic studies of motor control to help further tease apart the functional drivers of muscle spindle composition.

From fibre to function: are we accurately representing muscle architecture and performance?

The size and arrangement of fibres play a determinate role in the kinetic and energetic performance of muscles. Extrapolations between fibre architecture and performance underpin our understanding of how muscles function and how they are adapted to power specific motions within and across species. Here we provide a synopsis of how this 'fibre to function' paradigm has been applied to understand muscle design, performance and adaptation in animals. Our review highlights the widespread application of the fibre to function paradigm across a diverse breadth of biological disciplines but also reveals a potential and highly prevalent limitation running through past studies. Specifically, we find that quantification of muscle architectural properties is almost universally based on an extremely small number of fibre measurements. Despite the volume of research into muscle properties, across a diverse breadth of research disciplines, the fundamental assumption that a small proportion of fibre measurements can accurately represent the architectural properties of a muscle has never been quantitatively tested. Subsequently, we use a combination of medical imaging, statistical analysis, and physics-based computer simulation to address this issue for the first time. By combining diffusion tensor imaging (DTI) and deterministic fibre tractography we generated a large number of fibre measurements (>3000) rapidly for individual human lower limb muscles. Through statistical subsampling simulations of these measurements, we demonstrate that analysing a small number of fibres (n < 25) typically used in previous studies may lead to extremely large errors in the characterisation of overall muscle architectural properties such as mean fibre length and physiological cross-sectional area. Through dynamic musculoskeletal simulations of human walking and jumping, we demonstrate that recovered errors in fibre architecture characterisation have significant implications for quantitative predictions of in-vivo dynamics and muscle fibre function within a species. Furthermore, by applying data-subsampling simulations to comparisons of muscle function in humans and chimpanzees, we demonstrate that error magnitudes significantly impact both qualitative and quantitative assessment of muscle specialisation, potentially generating highly erroneous conclusions about the absolute and relative adaption of muscles across species and evolutionary transitions. Our findings have profound implications for how a broad diversity of research fields quantify muscle architecture and interpret muscle function.

Revision of hip flexor anatomy and function in modern humans, and implications for the evolution of hominin bipedalism.

Hip flexor musculature was instrumental in the evolution of hominin bipedal gait and in endurance running for hunting in the genus Homo. The iliacus and psoas major muscles were historically considered to have separate tendons with different insertions on the lesser trochanter. However, in the early 20th century, it became "common knowledge" that the two muscles insert together on the lesser trochanter as the "iliopsoas" tendon. We revisited the findings of early anatomists and tested the more recent paradigm of a common "iliopsoas" tendon based on dissections of hips and their associated musculature (n = 17). We rediscovered that the tendon of the psoas muscle inserts only into a crest running from the superior to anterior aspect of the lesser trochanter, separate from the iliacus. The iliacus inserts fleshly into the anterior portion of the lesser trochanter and into an inferior crest extending from it. We developed 3D multibody dynamics biomechanical models for: (a) the conjoint "iliopsoas" tendon hypothesis and (b) the separate insertion hypothesis. We show that the conjoint model underestimates the iliacus' capacity to generate hip flexion relative to the separate insertion model. Further work reevaluating the primate lower limb (including human) through dissection, needs to be performed to develop those datasets for reconstructing anatomy in fossil hominins using the extant phylogenetic bracket approach, which is frequently used for tetrapods clades outside of paleoanthropology.

Intra-subject sample size effects in plantar pressure analyses.

BACKGROUND: Recent work using large datasets (>500 records per subject) has demonstrated seemingly high levels of step-to-step variation in peak plantar pressure within human individuals during walking. One intuitive consequence of this variation is that smaller sample sizes (e.g., 10 steps per subject) may be quantitatively and qualitatively inaccurate and fail to capture the variance in plantar pressure of individuals seen in larger data sets. However, this remains quantitatively unexplored reflecting a lack of detailed investigation of intra-subject sample size effects in plantar pressure analysis. METHODS: Here we explore the sensitivity of various plantar pressure metrics to intra-subject sample size (number of steps per subject) using a random subsampling analysis. We randomly and incrementally subsample large data sets (>500 steps per subject) to compare variability in three metric types at sample sizes of 5-400 records: (1) overall whole-record mean and maximum pressure; (2) single-pixel values from five locations across the foot; and (3) the sum of pixel-level variability (measured by mean square error, MSE) from the whole plantar surface. RESULTS: Our results indicate that the central tendency of whole-record mean and maximum pressure within and across subjects show only minor sensitivity to sample size >200 steps. However, <200 steps, and particularly <50 steps, the range of overall mean and maximum pressure values yielded by our subsampling analysis increased considerably resulting in potential qualitative error in analyses of pressure changes with speed within-subjects and in comparisons of relative pressure magnitudes across subjects at a given speed. Our analysis revealed considerable variability in the absolute and relative response of the single pixel centroids of five regions to random subsampling. As the number of steps analysed decreased, the absolute value ranges were highest in the areas of highest pressure (medial forefoot and hallux), while the largest relative changes were seen in areas of lower pressure (the midfoot). Our pixel-level measure of variability by MSE across the whole-foot was highly sensitive to our manipulation of sample size, such that the range in MSE was exponentially larger in smaller subsamples. Random subsampling showed that the range in pixel-level MSE only came within 5% of the overall sample size in subsamples of >400 steps. The range in pixel-level MSE at low subsamples (<50) was 25-75% higher than that of the full datasets of >500 pressure records per subject. Overall, therefore, we demonstrate a high probability that the very small sample sizes (n < 20 records), which are routinely used in human and animal studies, capture a relatively low proportion of variance evident in larger plantar pressure data set, and thus may not accurately reflect the true population mean.

Back to the bones: do muscle area assessment techniques predict functional evolution across a macroevolutionary radiation?

Measures of attachment or accommodation area on the skeleton are a popular means of rapidly generating estimates of muscle proportions and functional performance for use in large-scale macroevolutionary studies. Herein, we provide the first evaluation of the accuracy of these muscle area assessment (MAA) techniques for estimating muscle proportions, force outputs and bone loading in a comparative macroevolutionary context using the rodent masticatory system as a case study. We find that MAA approaches perform poorly, yielding large absolute errors in muscle properties, bite force and particularly bone stress. Perhaps more fundamentally, these methods regularly fail to correctly capture many qualitative differences between rodent morphotypes, particularly in stress patterns in finite-element models. Our findings cast doubts on the validity of these approaches as means to provide input data for biomechanical models applied to understand functional transitions in the fossil record, and perhaps even in taxon-rich statistical models that examine broad-scale macroevolutionary patterns. We suggest that future work should go back to the bones to test if correlations between attachment area and muscle size within homologous muscles across a large number of species yield strong predictive relationships that could be used to deliver more accurate predictions for macroevolutionary and functional studies.

Foot anatomy, walking energetics, and the evolution of human bipedalism.

Interspecies differences in locomotor efficiency have been extensively researched, but within-species variation in the metabolic cost of walking and its underlying causes have received much less attention. This is somewhat surprising given the importance of walking energetics to natural selection, and the fact that the mechanical efficiency of striding bipedalism in modern humans is thought to be related in some part to the unique morphology of the human foot. Previous studies of human running have linked specific anatomical traits in the foot to variations in locomotor energetics to provide insight into form-function relationships in human evolution. However, such studies are relatively rare, particularly for walking. In this study, relationships between a range of functional musculoskeletal traits in the human lower limb and the energetics of walking over compliant and noncompliant substrates are examined, with particular focus on the lower limb and foot. Twenty-nine young, healthy individuals walked across three surfaces-a noncompliant laboratory floor, and compliant 6 cm and 13 cm thick foams-at self-selected speeds while oxygen consumption was measured, from which the metabolic cost of transport was calculated. Lower limb lengths, calcaneus lengths, foot shape indices, and maximum isometric plantarflexion torques were also measured and subsequently tested for relationships with metabolic cost over these surfaces using linear regression. It was found that metabolic cost varied considerably between individuals within and across substrate types, but this variation was not statistically related to or explained by variations in musculoskeletal parameters considered to be adaptively important to efficient bipedal locomotion. This therefore provides no supportive evidence that variations in these gross anatomical parameters confer significant advantages to the efficiency of walking, and therefore suggest caution in the use of similar metrics to infer differences in walking energetics in closely related fossil species.

Evolutionary versatility of the avian neck.

Bird necks display unparalleled levels of morphological diversity compared to other vertebrates, yet it is unclear what factors have structured this variation. Using three-dimensional geometric morphometrics and multivariate statistics, we show that the avian cervical column is a hierarchical morpho-functional appendage, with varying magnitudes of ecologically driven osteological variation at different scales of organization. Contrary to expectations given the widely varying ecological functions of necks in different species, we find that regional modularity of the avian neck is highly conserved, with an overall structural blueprint that is significantly altered only by the most mechanically demanding ecological functions. Nevertheless, the morphologies of vertebrae within subregions of the neck show more prominent signals of adaptation to ecological pressures. We also find that both neck length allometry and the nature of neck elongation in birds are different from other vertebrates. In contrast with mammals, neck length scales isometrically with head mass and, contrary to previous work, we show that neck elongation in birds is achieved predominantly by increasing vertebral lengths rather than counts. Birds therefore possess a cervical spine that may be unique in its versatility among extant vertebrates, one that, since the origin of flight, has adapted to function as a surrogate forelimb in varied ecological niches.