Comparative kinetics of humans and non-human primates during vertical climbing.

Climbing represents a critical behavior in the context of primate evolution. However, anatomically modern human populations are considered ill-suited for climbing. This adaptation can be attributed to the evolution of striding bipedalism, redirecting anatomical traits away from efficient climbing. Although prior studies have speculated on the kinetic consequences of this anatomical reorganization, there is a lack of data on the force profiles of human climbers. This study utilized high-speed videography and force plate analysis to assess single limb forces during climbing from 44 human participants of varying climbing experience and compared these data with climbing data from eight species of non-human primates (anthropoids and strepsirrhines). Contrary to expectations, experience level had no significant effect on the magnitude of single limb forces in humans. Experienced climbers did, however, demonstrate a predictable relationship between center of mass position and peak normal forces, suggesting a better ability to modulate forces during climbing. Humans exhibited significantly higher peak propulsive forces in the hindlimb compared with the forelimb and greater hindlimb dominance overall compared with non-human primates. All species sampled demonstrated exclusively tensile forelimbs and predominantly compressive hindlimbs. Strepsirrhines exhibited a pull-push transition in normal forces, while anthropoid primates, including humans, did not. Climbing force profiles are remarkably stereotyped across humans, reflecting the universal mechanical demands of this form of locomotion. Extreme functional differentiation between forelimbs and hindlimbs in humans may help to explain the evolution of bipedalism in ancestrally climbing hominoids.

Center of mass position does not drive energetic costs during climbing.

Climbing animals theoretically should optimize the energetic costs of vertical climbing while also maintaining stability. Many modifications to climbing behaviors have been proposed as methods of satisfying these criteria, focusing on controlling the center of mass (COM) during ascent. However, the link between COM movements and metabolic energy costs has yet to be evaluated empirically. In this study, we manipulated climbing conditions across three experimental setups to elicit changes in COM position, and measured the impact of these changes upon metabolic costs across a sample of 14 humans. Metabolic energy was assessed via open flow respirometry, while COM movements were tracked both automatically and manually. Our findings demonstrate that, despite inducing variation in COM position, the energetic costs of climbing remained consistent across all three setups. Differences in energetic costs were similarly not affected by body mass; however, velocity had a significant impact upon both cost of transport and cost of locomotion, but such a relationship disappeared when accounting for metabolic costs per stride. These findings suggest that climbing has inescapable metabolic demands driven by gaining height, and that attempts to mitigate such a cost, with perhaps the exception of increasing speed, have only minimal impacts. We also demonstrate that metabolic and mechanical energy costs are largely uncorrelated. Collectively, we argue that these data refute the idea that efficient locomotion is the primary aim during climbing. Instead, adaptations towards effective climbing should focus on stability and reducing the risk of falling, as opposed to enhancing the metabolic efficiency of locomotion.

Mechanical and morphometric approaches to body mass estimation in rhesus macaques: A test of skeletal variables.

OBJECTIVES: Estimation of body mass from skeletal metrics can reveal important insights into the paleobiology of archeological or fossil remains. The standard approach constructs predictive equations from postcrania, but studies have questioned the reliability of traditional measures. Here, we examine several skeletal features to assess their accuracy in predicting body mass. MATERIALS AND METHODS: Antemortem mass measurements were compared with common skeletal dimensions from the same animals postmortem, using 115 rhesus macaques (male: n = 43; female: n = 72). Individuals were divided into training (n = 58) and test samples (n = 57) to build and assess Ordinary Least Squares or multivariate regressions by residual sum of squares (RSS) and AIC weights. A leave-one-out approach was implemented to formulate the best fit multivariate models, which were compared against a univariate and a previously published catarrhine body-mass estimation model. RESULTS: Femur circumference represented the best univariate model. The best model overall was composed of four variables (femur, tibia and fibula circumference and humerus length). By RSS and AICw, models built from rhesus macaque data (RSS = 26.91, AIC = -20.66) better predicted body mass than did the catarrhine model (RSS = 65.47, AIC = 20.24). CONCLUSION: Body mass in rhesus macaques is best predicted by a 4-variable equation composed of humerus length and hind limb midshaft circumferences. Comparison of models built from the macaque versus the catarrhine data highlight the importance of taxonomic specificity in predicting body mass. This paper provides a valuable dataset of combined somatic and skeletal data in a primate, which can be used to build body mass equations for fragmentary fossil evidence.

Beakiation: how a novel parrot gait expands the locomotor repertoire of living birds.

Occupation of arboreal habitats poses myriad locomotor challenges, driving both anatomical and behavioural innovations across various tetrapod lineages. Here, we report and biomechanically assess a novel, beak-driven locomotor mode-'beakiation'-by which parrots advance along the underside of narrow arboreal substrates. Using high-speed videography and kinetic analyses, we describe the limb loading patterns and pendular mechanics of beakiation, and compare the biomechanical characteristics of this gait with other suspensory behaviours (namely, forelimb-driven brachiation and inverted quadrupedal walking). We report that the parrot beak experiences comparable force magnitudes (approx. 150% body weight in the normal plane; approx. 50% body weight in the fore-aft plane) to the forelimbs of brachiating primates. Parrot beakiation is also characterized by longer-than-expected pendular periods, similar to observations of gibbon brachiation. However, in terms of mechanical energy recovery, beakiation is typified by lower levels of energetic recovery than brachiating specialists: a product of its slower, more careful nature. The observation of this novel behaviour-which adds to a growing base of literature regarding beak-assisted locomotor strategies in birds-highlights the extraordinary behavioural plasticity of birds, the functional versatility of the avian beak, and the difficulties in reconstructing an animal's locomotor repertoire from morphological characteristics alone.

What does climbing mean exactly? Assessing spatiotemporal gait characteristics of inclined locomotion in parrots.

At what inclination does climbing begin? In this paper, we investigate the transition from walking to climbing in two species of parrot (Agapornis roseicollis and Nymphicus hollandicus) that are known to incorporate both their tail and their craniocervical system into the gait cycle during vertical climbing. Locomotor behaviors ranging in inclination were observed at angles between 0° and 90° for A. roseicollis, and 45°-85° degrees for N. hollandicus. Use of the tail in both species was observed at 45° inclination, and was joined at higher inclinations (> 65°) by use of the craniocervical system. Additionally, as inclination approached (but remained below) 90°, locomotor speeds were reduced while gaits were characterized by higher duty factors and lower stride frequency. These gait changes are consistent with those thought to increase stability. At 90°, A. roseicollis significantly increased its stride length, resulting in higher overall locomotor speed. Collectively these data demonstrate that the transition between horizontal walking and vertical climbing is gradual, incrementally altering several components of gait as inclinations increase. Such data underscore the need for further investigation into how exactly "climbing" is defined and the specific locomotor characteristics that differentiate this behavior from level walking.

Ecological correlates of three-dimensional muscle architecture within the dietarily diverse Strepsirrhini.

Analysis of muscle architecture, traditionally conducted via gross dissection, has been used to evaluate adaptive relationships between anatomical form and behavioral function. However, gross dissection cannot preserve three-dimensional relationships between myological structures for analysis. To analyze such data, we employ diffusible, iodine-based contrast-enhanced computed tomography (DiceCT) to explore the relationships between feeding ecology and masticatory muscle microanatomy in eight dietarily diverse strepsirrhines: allowing, for the first time, preservation of three-dimensional fascicle orientation and tortuosity across a functional comparative sample. We find that fascicle properties derived from these digital analyses generally agree with those measured from gross-dissected conspecifics. Physiological cross-sectional area was greatest in species with mechanically challenging diets. Frugivorous taxa and the wood-gouging species all exhibit long jaw adductor fascicles, while more folivorous species show the shortest relative jaw adductor fascicle lengths. Fascicle orientation in the parasagittal plane also seems to have a clear dietary association: most folivorous taxa have masseter and temporalis muscle vectors that intersect acutely while these vectors intersect obliquely in more frugivorous species. Finally, we observed notably greater magnitudes of fascicle tortuosity, as well as greater interspecific variation in tortuosity, within the jaw adductor musculature than in the jaw abductors. While the use of a single specimen per species precludes analysis of intraspecific variation, our data highlight the diversity of microanatomical variation that exists within the strepsirrhine feeding system and suggest that muscle architectural configurations are evolutionarily labile in response to dietary ecology-an observation to be explored across larger samples in the future.

The dual function of prokinesis in the feeding and locomotor systems of parrots.

Prokinesis, a mode of avian cranial kinesis involving motion between the neurocranium and upper beak, has long been investigated in biomechanical analyses of avian feeding and drinking. However, the modern avian beak is also used for non-feeding functions. Here, we investigate the dual function of prokinesis in the feeding and locomotor systems of the rosy-faced lovebird (Agapornis roseicollis). Lovebirds and other parrots utilize their beak both during feeding and as a third limb during vertical climbing. Thus, we experimentally measured both force-generating potential and movement of the rosy-faced lovebird mandible and maxilla (via prokinetic flexion of the craniofacial hinge) during tripedal climbing and mandibular/maxillary adduction. We found that whereas the maxilla is primarily responsible for generating force during locomotion, the mandible is primarily responsible for generating force during forceful jaw adduction, hinting at a remarkable capacity to alter prokinetic function with differing neuromuscular control. The ability of the prokinetic apparatus to perform functions with competing optimality criteria via modulation of motor control illustrates the functional plasticity of the avian cranial kinesis and sheds new light on the adaptive significance of cranial mobility.

A functional framework for interpreting phalangeal form.

Across tetrapods, the proportional lengths of the manual and pedal phalanges are highly constrained, following a generalized blueprint of shortening in a proximodistal gradient. Despite this, several lineages of both mammals (e.g. sloths, bats and colugos) and birds (e.g. raptors, parrots and woodpeckers) have broken this pattern, shortening the proximal phalanx while elongating more distal elements. As yet, no unifying explanation for this convergence has been empirically evaluated. This study combines a comparative phylogenetic assessment of phalangeal morphology across mammals and birds with a novel bioinspired robotics approach to explicitly test functional hypotheses relating to these morphotypes. We demonstrate that shortening the proximal phalanx allows taxa to maximize forces produced at the proximal interphalangeal joint, while elongation of subsequent elements maintains total ray length-ensuring arboreal species can still enclose large-diameter supports. Within suspensory and vertically clinging mammals, we additionally observe a secondary adaptation towards maximizing grip strength: namely increasing the height of the trochleae to increase the moment arm of digital flexor muscles that cross the joint. Together, our analyses highlight that numerous tetrapod lineages independently converged upon this morphotype to maximize proximal gripping strength, an adaptation to support specialized hunting and locomotor behaviours.

Skeletal age during hurricane impacts fluctuating asymmetry in Cayo Santiago rhesus macaques.

As natural disasters become more frequent due to climate change, understanding the biological impact of these ecological catastrophes on wild populations becomes increasingly pertinent. Fluctuating asymmetry (FA), or random deviations from bilateral symmetry, is reflective of developmental instability and has long been positively associated with increases in environmental stress. This study investigates craniofacial FA in a population of free-ranging rhesus macaques (Macaca mulatta) that has experienced multiple Category 3 hurricanes since the colony's inception on Cayo Santiago, including 275 individuals from ages 9 months to 31 years (F = 154; M = 121). Using geometric morphometrics to quantify FA and a linear mixed-effect model for analysis, we found that sex, age, and decade of birth did not influence the amount of FA in the individuals included in the study, but the developmental stage at which individuals experienced these catastrophic events greatly impacted the amount of FA exhibited (p = .001). Individuals that experienced these hurricanes during fetal life exhibited greater FA than any other post-natal developmental period. These results indicate that natural disasters can be associated with developmental disruption that results in long-term effects if occurring during the prenatal period, possibly due to increases in maternal stress-related hormones.

A medida que los desastres naturales se vuelven más frecuentes debido al cambio climático, entender el impacto biológico de estas catástrofes ecológicas en poblaciones silvestres va en aumento pertinente. La asimetría fluctuante (AF), o desviaciones aleatorias de simetría bilateral, es reflejo de inestabilidad durante el desarrollo y se ha asociado positivamente con incrementos en estrés ambiental durante mucho tiempo. Este estudio investiga AF craneofacial en una población de macacos rhesus (Macaca mulatta) en libertad que ha experimentado múltiples huracanes categoría 3 desde el inicio de la colonia en Cayo Santiago, e incluye 275 individuos de 9 meses a 31 años de edad (F = 154; M = 121). Usando morfometría geométrica para cuantificar AF y un modelo lineal de efectos mixtos para análisis, encontramos que el sexo, la edad y la década de nacimiento no influyeron en la cantidad de AF en los individuos incluidos en el estudio, pero la etapa de desarrollo en la que los individuos experimentaron estos eventos catastróficos impactó altamente la cantidad de AF exhibida (p = .001). Los individuos que experimentaron estos huracanes durante el período fetal exhibieron mayor AF que cualquier otro período de desarrollo posnatal. Estos resultados indican que los desastres naturales pueden asociarse con trastornos del desarrollo que tienen efectos a largo plazo si ocurren durante el período prenatal, posiblemente debido al aumento de hormonas maternas relacionadas con el estrés.

Patterns and Factors Influencing Parrot (Order: Psittaciformes) Success in Establishing Thriving Naturalized Populations within the Contiguous United States.

Parrots (Order: Psittaciformes) represent one of the most striking and ecomorphologically diverse avian clades, spanning more than two orders of magnitude in body size with populations occupying six continents. The worldwide diaspora of parrots is largely due to the pet trade, driven by human desire for bright, colorful, and intelligent animals as companions. Some introduced species have aptly inserted themselves into the local ecosystem and established successful breeding colonies all around the globe. Notably, the United States is home to several thriving populations of introduced species including red-masked parakeets (Psittacara erythrogenys), monk parakeets (Myiopsitta monachus), nanday conures (Aratinga nenday), and red-crowned amazons (Amazona viridigenalis). Their incredible success globally begs the question as to how these birds adapt so readily to novel environments. In this commentary, we trace parrots through evolutionary history, contextualize existent naturalized parrot populations within the contiguous United States, and provide a phylogenetic regression analysis of body mass and brain size based on success in establishing breeding populations. The propensity for a parrot species to become established appears to be phylogenetically driven. Notably, parrots in the family Cacatuidae and Neotropical Pyrrhua appear to be poor at establishing themselves in the United States once released. Although brain size among Psittaciformes did not show a significant impact on successful breeding in the continental United States, we propose that the success of parrots can be attributed to their charismatic nature, significant intelligence relative to other avian lineages, and behavioral flexibility.

Tail feather strength in tail-assisted climbing birds is achieved through geometric, not material change.

Birds encompass vast ecomorphological diversity and practise numerous distinct locomotor modes. One oft-cited feature seen in climbing birds is an increase in tail 'stiffness', yet it remains unclear to what extent these feathers are altered, and the specific mechanism by which differences in functional performance are attained. We collected a broad taxonomic sample of tail feathers (6525 total, from 774 species representing 21 avian orders and ranging in size from approximately 3 g to greater than 11 kg) and present data on their material properties, cross-sectional geometry and morphometrics. Ordinary and phylogenetic least-squares regressions of each variable versus body mass were conducted to assess scaling relationships and demonstrate that tail-supported climbers exhibit longer tail feathers with a wider rachis base and tip, and a greater second moment of area and maximum bending moment. However, no differences were observed in the material properties of the keratin itself. This suggests that tail-supported arboreal climbing birds of multiple orders have independently adopted similar morphologies. Moreover, these geometric relationships follow the same allometric scaling relationships as seen in the long bones of mammalian limbs, suggesting that the morphology of these developmentally and evolutionarily distinct structures are governed by similar functional constraints of weight support.

Dynamics of horizontal walking and vertical climbing in the Australian green tree frog (Ranoidea caerulea).

Despite the high mechanical demands associated with climbing, the ability to ascend vertically has evolved independently in most major animal lineages. However, little is known about the kinetics, mechanical energy profiles or spatiotemporal gait characteristics of this locomotor mode. In this study, we explored the dynamics of horizontal locomotion and vertical climbing on both flat substrates and narrow poles in five Australian green tree frogs (Ranoidea caerulea). Vertical climbing is associated with slow, deliberate movements (i.e. reduced speed and stride frequency and increased duty factors) and propulsive fore-aft impulses in both the forelimb and hindlimb. By comparison, horizontal walking was characterized by a braking forelimb and a propulsive hindlimb. In the normal plane, tree frogs mirrored other taxa in exhibiting a net pulling forelimb and a net pushing hindlimb during vertical climbing. In terms of mechanical energy, tree frogs matched theoretical predictions of climbing dynamics (i.e. the total mechanical energetic cost of vertical climbing was predominantly driven by potential energy, with negligible kinetic contributions). Utilizing power as a means of estimating efficiency, we also demonstrate that Australian green tree frogs show total mechanical power costs only slightly above the minimum mechanical power necessary to climb, highlighting their highly effective locomotor mechanics. This study provides new data on climbing dynamics in a slow-moving arboreal tetrapod and raises new testable hypotheses about how natural selection can act upon a locomotor behavior that is notably constrained by external physical forces.

Behavioral correlates of fascicular organization: The confluence of muscle architectural anatomy and function.

Muscle is a complex tissue that has been studied on numerous hierarchical levels: from gross descriptions of muscle organization to cellular analyses of fiber profiles. In the middle of this space between organismal and cellular biology lies muscle architecture, the level at which functional correlations between a muscle's internal fiber organization and contractile abilities are explored. In this review, we summarize this relationship, detail recent advances in our understanding of this form-function paradigm, and highlight the role played by The Anatomical Record in advancing our understanding of functional morphology within muscle over the past two decades. In so doing, we honor the legacy of Editor-in-Chief Kurt Albertine, whose stewardship of the journal from 2006 through 2020 oversaw the flourishing of myological research, including numerous special issues dedicated to exploring the behavioral correlates of myology across diverse taxa. This legacy has seen the The Anatomical Record establish itself as a preeminent source of myological research, and a true leader within the field of comparative anatomy and functional morphology.

Three toes and three modes: Dynamics of terrestrial, suspensory, and vertical locomotion in brown-throated three-toed sloths (Bradypodidae, Xenarthra).

Living sloths exhibit numerous anatomical specializations towards inverted quadrupedalism, however, previous studies have noted a more varied locomotor repertoire than previously anticipated. In this study, we present spatiotemporal gait characteristics and triaxial kinetic data from the brown-throated three-toed sloth (Bradypus variegatus) across three locomotor modes: terrestrial quadrupedal "crawling", suspensory walking, and vertical climbing. Compared to quadrupedal crawling and suspensory walking, B. variegatus adopted longer contact times and stride durations, larger duty factors, and greater speed during vertical climbing. Net fore-aft impulses were significantly greater during vertical climbing in both limb pairs than in quadrupedal crawling and suspensory walking. Functionally, during quadrupedal crawling and vertical climbing, both limb pairs served propulsive roles, while differentiation between a propulsive forelimb and braking hindlimb was observed during suspension. Net tangential forces differentiated vertical climbing kinetics from the other modes of locomotion, with the introduction of bidirectional pulling and pushing forces in the forelimb and hindlimb, respectively. The net mediolateral impulses were similar in vertical climbing and quadrupedal crawling as both limb pairs directed forces in one direction, whereas during suspensory walking, the laterally dominant forelimb was opposed by the medially dominant hindlimb. In total, this study provides novel data on the diverse locomotor dynamics in a slow-moving arboreal tetrapod and posits new testable hypotheses about the neuroplasticity and ease of transitioning between locomotor behaviors. The strikingly similar kinetic profiles of quadrupedal crawling and suspensory walking compared to vertical climbing suggest shared neuromuscular and mechanical demands between these mirrored locomotor modes.

Ecomorphological correlates of inner and middle ear anatomy within phyllostomid bats.

Echolocation is the primary sense used by most bats to navigate their environment. However, the influence of echolocating behaviors upon the morphology of the auditory apparatus remains largely uninvestigated. While it is known that middle ear ossicle size scales positively with body mass across mammals, and that peak call frequency scales negatively with body mass among bats, there are still large gaps in our understanding of the degree to which allometry or ecology influences the morphology of the chiropteran auditory apparatus. To investigate this, we used µCT datasets to quantify three morphological components of the inner and middle ear: ossicle size, ossicle shape, and cochlear spirality. These data were collected across 27 phyllostomid species, spanning a broad range of body sizes, habitats, and dietary categories, and the relationships between these variables and ear morphology were assessed using a comparative phylogenetic approach. Ossicle size consistently scaled with strong negative allometry relative to body mass. Cochlear spirality was significantly (p = .025) associated with wing aspect ratio (a proxy for habitat use) but was not associated with body mass. From a morphological perspective, the malleus and incus exhibited some variation in kind with diet and call frequency, while stapes morphology is more closely tied to body size. Future work will assess these relationships within other chiropteran lineages, and investigate potential morphological differences in the middle and inner ear of echolocating-vs-non-echolocating taxa.

The influence of substrate size upon pulling and gripping forces in parrots (Psittaciformes: Agapornis roseicollis).

The ability to securely grasp substrates of variable diameter is critical to arboreal animals. Arboreal specialists have emerged across several vertebrate lineages - including mammals, lizards and amphibians - and several attempts have been made to quantify their grasping performance, by measuring either gripping (i.e. forces generated about an object or substrate enclosed within the digits) or pulling (i.e. the ability to resist being removed from a substrate) forces. In this study, we present data on both pulling and gripping performance across a range of substrate diameters (0.5-17.5 mm) within a model parrot species (Agapornis roseicollis). Parrots represent an ancient arboreal lineage, allowing us to compare their abilities with those of arboreal specialists within other tetrapod groups. Data were collected using 3D-printed perches of variable diameter, and forces were registered using either an AMTI low-load force plate (grip force) or a Harvard Apparatus portable strength tester (pull force). Gripping forces peaked at a 5 mm diameter perch, while pulling forces were greatest at a 2.5 mm diameter. All forces strongly diminished above 10 mm size, suggesting grip force is optimized when utilizing small perches, a finding which corresponds to observational studies of preferential perching habits among free-ranging parrots. Relative grasping performance (adjusted for body size) in parrots is roughly equivalent to that of other arboreal specialists from other tetrapod lineages, but low when compared with that of raptorial birds that utilize their feet during aerial prey capture. Further taxonomic sampling is encouraged to contextualize how grasping performance varies in an adaptive evolutionary context.

Positional Behavior of Introduced Monk Parakeets (Myiopsitta monachus) in an Urban Landscape.

Positional behaviors have been broadly quantified across the Order Primates, and in several other mammalian lineages, to contextualize adaptations to, and evolution within, an arboreal environment. Outside of Mammalia, however, such data are yet to be reported. In this study, we present the first quantitative report of positional behavior within Aves, presenting 11,246 observations of scan sampling data from a colony of Monk Parakeets (Myiopsitta monachus) from Brooklyn, New York City. Each scan recorded locomotor and postural behavior and information about weather condition, temperature, and substrate properties (e.g., type, size, orientation). A distinction was also recorded between natural and artificial substrates. Parrots exhibited a strong preference for small and terminal branches, a selection which may reflect targeted foraging of new fruit growth and leaf-buds. We further observed that the gait transition from walking to sidling appears primarily driven by substrate size, with the former preferred on the ground and on large, broad substrates and the latter used to navigate smaller branches. Finally, we observed an increase in locomotor diversity on artificial versus naturally occurring substrates. This demonstrates the importance of a flexible behavioral repertoire in facilitating a successful transition towards an urban landscape in introduced species.

A morphological analysis of carnivoran ossicles from Rancho La Brea.

The morphology of the mammalian middle ear-including the size, shape, and stiffness of individual ossicles-controls their vibrational response to sound and, is closely related to an animal's auditory capabilities. While the relationship between middle ear morphology and hearing frequency has been explored in living carnivorans, the size and shape of ossicles in fossil carnivorans have been sparsely documented. In this study, we present the first morphological data on four iconic carnivoran taxa from the Rancho La Brea Tar Pits: Smilodon fatalis, Panthera atrox, Canis dirus, and Arctodus simus. These data are contextualized with samples of extant felids, canids, and ursids to determine the extent to which the ossicles of these iconic fossil taxa resemble their living relatives. Six, five, and seven linear measurements were taken from the malleus, incus, and stapes, respectively. Comparisons of geometric means reveal that the ossicles of fossil canids and felids are similar in size to living analogs, but those of A. simus are significantly larger than those of any living ursid. Further, principal components analyses demonstrate close morphological affinities between fossil and extant taxa within canids and felids, and again, a greater disparity between fossil and extant ursids. Canids and ursids occupy distinct regions of the morphospace, yet both overlap the morphological range spanned by felids. While some elements-for example, the stapes-require further specimens to facilitate more nuanced interpretations of variation, our findings underscore the need for concerted efforts towards identifying and preserving these bones within fossil assemblages.

Decoupled evolution of the cranium and mandible in carnivoran mammals.

The relationship between skull morphology and diet is a prime example of adaptive evolution. In mammals, the skull consists of the cranium and the mandible. Although the mandible is expected to evolve more directly in response to dietary changes, dietary regimes may have less influence on the cranium because additional sensory and brain-protection functions may impose constraints on its morphological evolution. Here, we tested this hypothesis by comparing the evolutionary patterns of cranium and mandible shape and size across 100+ species of carnivoran mammals with distinct feeding ecologies. Our results show decoupled modes of evolution in cranial and mandibular shape; cranial shape follows clade-based evolutionary shifts, whereas mandibular shape evolution is linked to broad dietary regimes. These results are consistent with previous hypotheses regarding hierarchical morphological evolution in carnivorans and greater evolutionary lability of the mandible with respect to diet. Furthermore, in hypercarnivores, the evolution of both cranial and mandibular size is associated with relative prey size. This demonstrates that dietary diversity can be loosely structured by craniomandibular size within some guilds. Our results suggest that mammal skull morphological evolution is shaped by mechanisms beyond dietary adaptation alone.