Adaptive landscapes unveil the complex evolutionary path from sprawling to upright forelimb function and posture in mammals.

The 'sprawling-parasagittal' postural transition is a key part of mammalian evolution, associated with sweeping reorganization of the postcranial skeleton in mammals compared to their forebears, the non-mammalian synapsids. However, disputes over forelimb function in fossil synapsids render the precise nature of the 'sprawling-parasagittal' transition controversial. We shed new light on the origins of mammalian posture, using evolutionary adaptive landscapes to integrate 3D humerus shape and functional performance data across a taxonomically comprehensive sample of fossil synapsids and extant comparators.

Synchrotron X-ray micro-computed tomography enhances our knowledge of the skull anatomy of a Late Triassic ecteniniid cynodont with hypercanines.

Hypercanines, or hypertrophied canines, are observed in a wide range of both extinct and extant synapsids. In non-mammaliaform cynodonts, the Permo-Triassic forerunners of mammals, long canines are not uncommon, appearing in several unrelated taxa within the clade. Among them is Trucidocynodon riograndensis, a carnivorous ecteniniid cynodont from the Late Triassic of Brazil, which exhibits a specialized dentition, including spear-shaped incisors, very long and narrow canines, and sectorial postcanines with distally oriented cusps, all of which have finely serrated margins.

Origins of mammalian vertebral function revealed through digital bending experiments.

Unravelling the functional steps that underlie major transitions in the fossil record is a significant challenge for biologists owing to the difficulties of interpreting functional capabilities of extinct organisms. New computational modelling approaches provide exciting avenues for testing function in the fossil record. Here, we conduct digital bending experiments to reconstruct vertebral function in non-mammalian synapsids, the extinct forerunners of mammals, to provide insights into the functional underpinnings of the synapsid-mammal transition.

Derived faunivores are the forerunners of major synapsid radiations.

Evolutionary radiations generate most of Earth's biodiversity, but are there common ecomorphological traits among the progenitors of radiations? In Synapsida (the mammalian total group), 'small-bodied faunivore' has been hypothesized as the ancestral state of most major radiating clades, but this has not been quantitatively assessed across multiple radiations. To examine macroevolutionary patterns in a phylogenetic context, we generated a time-calibrated metaphylogeny ('metatree') comprising 1,888 synapsid species from the Carboniferous through the Eocene (305-34 Ma) based on 269 published character matrices. We used comparative methods to investigate body size and dietary evolution during successive synapsid radiations.

Diversification of the ruminant skull along an evolutionary line of least resistance.

Clarifying how microevolutionary processes scale to macroevolutionary patterns is a fundamental goal in evolutionary biology, but these analyses, requiring comparative datasets of population-level variation, are limited. By analyzing a previously published dataset of 2859 ruminant crania, we find that variation within and between ruminant species is biased by a highly conserved mammalian-wide allometric pattern, CREA (craniofacial evolutionary allometry), where larger species have proportionally longer faces. Species with higher morphological integration and species more biased toward CREA have diverged farther from their ancestors, and Ruminantia as a clade diversified farther than expected in the direction of CREA.

The topography of diet: Orientation patch count predicts diet in turtles.

Use of quantitative morphological methods in biology has increased with the availability of 3D digital data. Rotated orientation patch count (OPCr) leverages such data to quantify the complexity of an animal's feeding surface, and has previously been used to analyze how tooth complexity signals diet in squamates, crocodilians, and mammals. These studies show a strong correlation between dental complexity and diet.

Fossil bone histology reveals ancient origins for rapid juvenile growth in tetrapods.

Patterns of growth throughout the lifetime of an animal reflect critical life history traits such as reproductive timing, physiology, and ecological interactions. The ancestral growth pattern for tetrapods has traditionally been described as slow-to-moderately paced, akin to modern amphibians, with fast growth and high metabolic rates considered a specialized physiological trait of amniotes. Here, we present bone histology from an ontogenetic series of the Early Carboniferous stem tetrapod Whatcheeria deltae, and document evidence of fibrolamellar bone-primary bone tissue associated with fast growth.

Inner ear biomechanics reveals a Late Triassic origin for mammalian endothermy.

Endothermy underpins the ecological dominance of mammals and birds in diverse environmental settings. However, it is unclear when this crucial feature emerged during mammalian evolutionary history, as most of the fossil evidence is ambiguous. Here we show that this key evolutionary transition can be investigated using the morphology of the endolymph-filled semicircular ducts of the inner ear, which monitor head rotations and are essential for motor coordination, navigation and spatial awareness.

A Shrewd Inspection of Vertebral Regionalization in Large Shrews (Soricidae: Crocidurinae).

The regionalization of the mammalian spinal column is an important evolutionary, developmental, and functional hallmark of the clade. Vertebral column regions are usually defined using transitions in external bone morphology, such as the presence of transverse foraminae or rib facets, or measurements of vertebral shape. Yet the internal structure of vertebrae, specifically the trabecular (spongy) bone, plays an important role in vertebral function, and is subject to the same variety of selective, functional, and developmental influences as external bone morphology.

Living fast in the Triassic: New data on life history in Lystrosaurus (Therapsida: Dicynodontia) from northeastern Pangea.

Lystrosaurus was one of the few tetrapods to survive the Permo-Triassic mass extinction, the most profound biotic crisis in Earth's history. The wide paleolatitudinal range and high abundance of Lystrosaurus during the Early Triassic provide a unique opportunity to investigate changes in growth dynamics and longevity following the mass extinction, yet most studies have focused only on species that lived in the southern hemisphere. Here, we present the long bone histology from twenty Lystrosaurus skeletal elements spanning a range of sizes that were collected in the Jiucaiyuan Formation of northwestern China.

Effect of captivity on the vertebral bone microstructure of xenarthran mammals.

Captive specimens in museum collections facilitate study of rare taxa, but the lifestyles, diets, and lifespans of captive animals differ from their wild counterparts. Trabecular bone architecture adapts to in vivo forces, and may reflect interspecific variation in ecology and behavior as well as intraspecific variation between captive and wild specimens. We compared trunk vertebrae bone microstructure in captive and wild xenarthran mammals to test the effects of ecology and captivity.

The limits of convergence: the roles of phylogeny and dietary ecology in shaping non-avian amniote crania.

Cranial morphology is remarkably varied in living amniotes and the diversity of shapes is thought to correspond with feeding ecology, a relationship repeatedly demonstrated at smaller phylogenetic scales, but one that remains untested across amniote phylogeny. Using a combination of morphometric methods, we investigate the links between phylogenetic relationships, diet and skull shape in an expansive dataset of extant toothed amniotes: mammals, lepidosaurs and crocodylians. We find that both phylogeny and dietary ecology have statistically significant effects on cranial shape.

Histology and μCT reveal the unique evolution and development of multiple tooth rows in the synapsid Endothiodon.

Several amniote lineages independently evolved multiple rows of marginal teeth in response to the challenge of processing high fiber plant matter. Multiple tooth rows develop via alterations to tooth replacement in captorhinid reptiles and ornithischian dinosaurs, but the specific changes that produce this morphology differ, reflecting differences in their modes of tooth attachment. To further understand the mechanisms by which multiple tooth rows can develop, we examined this feature in Endothiodon bathystoma, a member of the only synapsid clade (Anomodontia) to evolve a multi-rowed marginal dentition.

Local Superimpositions Facilitate Morphometric Analysis of Complex Articulating Structures.

Articulating structures, such as the vertebrate skeleton or the segmented arthropod exoskeleton, comprise a majority of the morphological diversity across the eukaryotic tree of life. Quantifying the form of articulating structures is therefore imperative for a fuller understanding of the factors influencing biological form. A wealth of freely available 3D data capturing this morphological diversity is stored in online repositories such as Morphosource, but the geometric morphometric analysis of an articulating structure is impeded by arbitrary differences in the resting positions of its individual articulating elements.

Phylogeny, function and ecology in the deep evolutionary history of the mammalian forelimb.

Mammals are the only living members of the larger clade Synapsida, which has a fossil record spanning 320 Ma. Despite the fact that much of the ecological diversity of mammals has been considered in the light of limb morphology, the ecological comparability of mammals to their fossil forerunners has not been critically assessed. Because of the wide use of limb morphology in testing ecomorphological hypothesis about extinct tetrapods, we sought: (i) to estimate when in synapsid history, modern mammals become analogues for predicting fossil ecologies; (ii) to document examples of ecomorphological convergence; and (iii) to compare the functional solutions of distinct synapsid radiations.

Evidence from South Africa for a protracted end-Permian extinction on land.

Earth's largest biotic crisis occurred during the Permo-Triassic Transition (PTT). On land, this event witnessed a turnover from synapsid- to archosauromorph-dominated assemblages and a restructuring of terrestrial ecosystems. However, understanding extinction patterns has been limited by a lack of high-precision fossil occurrence data to resolve events on submillion-year timescales.

Adaptive landscapes challenge the "lateral-to-sagittal" paradigm for mammalian vertebral evolution.

The evolution of mammals from their extinct forerunners, the non-mammalian synapsids, is one of the most iconic locomotor transitions in the vertebrate fossil record. In the limb skeleton, the synapsid-mammal transition is traditionally characterized by a shift from a sprawling limb posture, resembling that of extant reptiles and amphibians, to more adducted limbs, as seen in modern-day mammals. Based on proposed postural similarities between early synapsids and extant reptiles, this change is thought to be accompanied by a shift from ancestral reptile-like lateral bending to mammal-like sagittal bending of the vertebral column.

Effects of taphonomic deformation on geometric morphometric analysis of fossils: a study using the dicynodont (Therapsida, Anomodontia).

Taphonomic deformation, the distortion of fossils as a result of geological processes, poses problems for the use of geometric morphometrics in addressing paleobiological questions. Signal from biological variation, such as ontogenetic trends and sexual dimorphism, may be lost if variation from deformation is too high. Here, we investigate the effects of taphonomic deformation on geometric morphometric analyses of the abundant, well known Permian therapsid .

Deciphering an extreme morphology: bone microarchitecture of the hero shrew backbone (Soricidae: ).

Biological structures with extreme morphologies are puzzling because they often lack obvious functions and stymie comparisons to homologous or analogous features with more typical shapes. An example of such an extreme morphotype is the uniquely modified vertebral column of the hero shrew , which features numerous accessory intervertebral articulations and massively expanded transverse processes. The function of these vertebral structures is unknown, and it is difficult to meaningfully compare them to vertebrae from animals with known behavioural patterns and spinal adaptations.

Author Correction: Regionalization of the axial skeleton predates functional adaptation in the forerunners of mammals.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Regionalization of the axial skeleton predates functional adaptation in the forerunners of mammals.

The evolution of semi-independent modules is hypothesized to underlie the functional diversification of serially repeating (metameric) structures. The mammal vertebral column is a classic example of a metameric structure that is both modular, with well-defined morphological regions, and functionally differentiated. How the evolution of regions is related to their functional differentiation in the forerunners of mammals remains unclear.

Stepwise shifts underlie evolutionary trends in morphological complexity of the mammalian vertebral column.

A fundamental concept in evolutionary biology is that life tends to become more complex through geologic time, but empirical examples of this phenomenon are controversial. One debate is whether increasing complexity is the result of random variations, or if there are evolutionary processes which actively drive its acquisition, and if these processes act uniformly across clades. The mammalian vertebral column provides an opportunity to test these hypotheses because it is composed of serially-repeating vertebrae for which complexity can be readily measured.

Antiquity of forelimb ecomorphological diversity in the mammalian stem lineage (Synapsida).

Mammals and their closest fossil relatives are unique among tetrapods in expressing a high degree of pectoral girdle and forelimb functional diversity associated with fully pelagic, cursorial, subterranean, volant, and other lifestyles. However, the earliest members of the mammalian stem lineage, the "pelycosaur"-grade synapsids, present a far more limited range of morphologies and inferred functions. The more crownward nonmammaliaform therapsids display novel forelimb morphologies that have been linked to expanded functional diversity, suggesting that the roots of this quintessentially mammalian phenotype can be traced to the pelycosaur-therapsid transition in the Permian period.