Forelimb morphology is an indicator for terrestrial locomotor ecology. The limb morphology of the enigmatic tapir (Perissodactyla: Tapirus) has often been compared to that of basal perissodactyls, despite the lack of quantitative studies comparing forelimb variation in modern tapirs. Here, we present a quantitative assessment of tapir upper forelimb osteology using three-dimensional geometric morphometrics to test whether the four modern tapir species are monomorphic in their forelimb skeleton. The shape of the upper forelimb bones across four species (T. indicus; T. bairdii; T. terrestris; T. pinchaque) was investigated. Bones were laser scanned to capture surface morphology and 3D landmark analysis was used to quantify shape. Discriminant function analyses were performed to reveal features which could be used for interspecific discrimination. Overall our results show that the appendicular skeleton contains notable interspecific differences. We demonstrate that upper forelimb bones can be used to discriminate between species (>91% accuracy), with the scapula proving the most diagnostic bone (100% accuracy). Features that most successfully discriminate between the four species include the placement of the cranial angle of the scapula, depth of the humeral condyle, and the caudal deflection of the olecranon. Previous studies comparing the limbs of T. indicus and T. terrestris are corroborated by our quantitative findings. Moreover, the mountain tapir T. pinchaque consistently exhibited the greatest divergence in morphology from the other three species. Despite previous studies describing tapirs as functionally mediportal in their locomotor style, we find osteological evidence suggesting a spectrum of locomotor adaptations in the tapirs. We conclude that modern tapir forelimbs are neither monomorphic nor are tapirs as conserved in their locomotor habits as previously described. J. Morphol. 277:1469-1485, 2016. © 2016 Wiley Periodicals, Inc.

Morphological responses of nonmammalian herbivores to external ecological drivers have not been quantified over extended timescales. Herbivorous nonavian dinosaurs are an ideal group to test for such responses, because they dominated terrestrial ecosystems for more than 155 Myr and included the largest herbivores that ever existed. The radiation of dinosaurs was punctuated by several ecologically important events, including extinctions at the Triassic/Jurassic (Tr/J) and Jurassic/Cretaceous (J/K) boundaries, the decline of cycadophytes, and the origin of angiosperms, all of which may have had profound consequences for herbivore communities. Here we present the first analysis of morphological and biomechanical disparity for sauropodomorph and ornithischian dinosaurs in order to investigate patterns of jaw shape and function through time. We find that morphological and biomechanical mandibular disparity are decoupled: mandibular shape disparity follows taxonomic diversity, with a steady increase through the Mesozoic. By contrast, biomechanical disparity builds to a peak in the Late Jurassic that corresponds to increased functional variation among sauropods. The reduction in biomechanical disparity following this peak coincides with the J/K extinction, the associated loss of sauropod and stegosaur diversity, and the decline of cycadophytes. We find no specific correspondence between biomechanical disparity and the proliferation of angiosperms. Continual ecological and functional replacement of pre-existing taxa accounts for disparity patterns through much of the Cretaceous, with the exception of several unique groups, such as psittacosaurids that are never replaced in their biomechanical or morphological profiles.

The absence of preserved soft tissues in the fossil record is frequently a hindrance for palaeontologists wishing to investigate morphological shifts in key skeletal systems, such as the limbs. Understanding the soft tissue composition of modern species can aid in understanding changes in musculoskeletal features through evolution, including those pertaining to locomotion. Establishing anatomical differences in soft tissues utilising an extant phylogenetic bracket can, in turn, assist in interpreting morphological changes in hard tissues and modelling musculoskeletal movements during evolutionary transitions (e.g. digit reduction in perissodactyls). Perissodactyls (horses, rhinoceroses, tapirs and their relatives) are known to have originated with a four‐toed (tetradactyl) forelimb condition. Equids proceeded to reduce all but their central digit, resulting in monodactyly, whereas tapirs retained the ancestral tetradactyl state. The modern Malayan tapir (Tapirus indicus) has been shown to exhibit fully functional tetradactyly in its forelimb, more so than any other tapir, and represents an ideal case‐study for muscular arrangement and architectural comparison with the highly derived monodactyl Equus. Here, we present the first quantification of muscular architecture of a tetradactyl perissodactyl (T. indicus), and compare it to measurements from modern monodactyl caballine horse (Equus ferus caballus). Each muscle of the tapir forelimb was dissected out from a cadaver and measured for architectural properties: muscle‐tendon unit (MTU) length, MTU mass, muscle mass, pennation angle, and resting fibre length. Comparative parameters [physiological cross‐sectional area (PCSA), muscle volume, and % muscle mass] were then calculated from the raw measurements. In the shoulder region, the infraspinatus of T. indicus exhibits dual origination sites on either side of the deflected scapular spine. Within ungulates, this condition has only been previously reported in suids. Differences in relative contribution to limb muscle mass between T. indicus and Equus highlight forelimb muscles that affect mobility in the lateral and medial digits (e.g. extensor digitorum lateralis). These muscles were likely reduced in equids during their evolutionary transition from tetradactyl forest‐dwellers to monodactyl, open‐habitat specialists. Patterns of PCSA across the forelimb were similar between T. indicus and Equus, with the notable exceptions of the biceps brachii and flexor carpi ulnaris, which were much larger in Equus. The differences observed in PCSA between the tapir and horse forelimb muscles highlight muscles that are essential for maintaining stability in the monodactyl limb while moving at high speeds. This quantitative dataset of muscle architecture in a functionally tetradactyl perissodactyl is a pivotal first step towards reconstructing the locomotor capabilities of extinct, four‐toed ancestors of modern perissodactyls, and providing further insights into the equid locomotor transition.

Throughout their evolution, tetrapods have repeatedly colonised a series of ecological niches in marine ecosystems, producing textbook examples of convergent evolution. However, this evolutionary phenomenon has typically been assessed qualitatively and in broad-brush frameworks that imply simplistic macroevolutionary landscapes. We establish a protocol to visualize the density of trait space occupancy and thoroughly test for the existence of macroevolutionary landscapes. We apply this protocol to a new phenotypic dataset describing the morphology of short-necked plesiosaurians, a major component of the Mesozoic marine food webs (ca. 201 to 66 Mya). Plesiosaurians evolved this body plan multiple times during their 135-million-year history, making them an ideal test case for the existence of macroevolutionary landscapes. We find ample evidence for a bimodal craniodental macroevolutionary landscape separating latirostrines from longirostrine taxa, providing the first phylogenetically-explicit quantitative assessment of trophic diversity in extinct marine reptiles. This bimodal pattern was established as early as the Middle Jurassic and was maintained in evolutionary patterns of short-necked plesiosaurians until a Late Cretaceous (Turonian) collapse to a unimodal landscape comprising longirostrine forms with novel morphologies. This study highlights the potential of severe environmental perturbations to profoundly alter the macroevolutionary dynamics of animals occupying the top of food chains.