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1. Locomotor shifts, stylopod proportions, and the evolution of allometry in Synapsida

   https://doi.org/10.1002/ar.70006  

   Bishop, P_J; Pierce, S_E (June 2025, The Anatomical Record)

   Abstract Locomotor evolution in synapsids involved numerous functional shifts associated with the transition from sprawled to erect limb postures on the line to therian mammals. Given that bone structure frequently reflects functional requirements, this study investigated evolutionary changes in synapsid humerus and femur proportions as a lens to evaluate functional shifts through time. A total of 936 bones were measured, representing 330 species across the full 320+ million years of synapsid history. This dataset was used to test whether transformations in stylopod proportions are consistent with inferred changes in bone loading mechanics, alignment of joint and muscle forces, muscular control of the shoulder and hip, and differential support of body weight by the fore‐ and hindlimbs. As variation in bone dimensions may also correlate with bone or body size, this study first developed a novel approach for calculating species‐specific, size‐corrected measures of bone proportions. By disentangling the effect of body size from functional signals recorded in bone geometry, this then enabled a node‐to‐node appraisal of how bone allometry itself evolved through time. Ancestral state reconstruction of size‐corrected stylopod proportions reveals trends that broadly support many hypothesized shifts in locomotor biomechanics along the therian stem lineage. However, patterns of transformation are frequently complex, suggesting functional mosaicism, and stylopod proportions that typify therians as a whole are often not achieved until crown Theria itself. Several instances of temporary trend reversal are also inferred, particularly within non‐mammalian cynodonts, indicating greater functional or ecological diversification in this group. 

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2. Broad similarities in shoulder muscle architecture and organization across two amniotes: implications for reconstructing non-mammalian synapsids

   https://doi.org/10.7717/peerj.8556  

   Fahn-Lai, Philip; Biewener, Andrew A.; Pierce, Stephanie E. (January 2020, PeerJ)

   The evolution of upright limb posture in mammals may have enabled modifications of the forelimb for diverse locomotor ecologies. A rich fossil record of non-mammalian synapsids holds the key to unraveling the transition from “sprawling” to “erect” limb function in the precursors to mammals, but a detailed understanding of muscle functional anatomy is a necessary prerequisite to reconstructing postural evolution in fossils. Here we characterize the gross morphology and internal architecture of muscles crossing the shoulder joint in two morphologically-conservative extant amniotes that form a phylogenetic and morpho-functional bracket for non-mammalian synapsids: the Argentine black and white tegu Salvator merianae and the Virginia opossum Didelphis virginiana . By combining traditional physical dissection of cadavers with nondestructive three-dimensional digital dissection, we find striking similarities in muscle organization and architectural parameters. Despite the wide phylogenetic gap between our study species, distal muscle attachments are notably similar, while differences in proximal muscle attachments are driven by modifications to the skeletal anatomy of the pectoral girdle that are well-documented in transitional synapsid fossils. Further, correlates for force production, physiological cross-sectional area (PCSA), muscle gearing (pennation), and working range (fascicle length) are statistically indistinguishable for an unexpected number of muscles. Functional tradeoffs between force production and working range reveal muscle specializations that may facilitate increased girdle mobility, weight support, and active stabilization of the shoulder in the opossum—a possible signal of postural transformation. Together, these results create a foundation for reconstructing the musculoskeletal anatomy of the non-mammalian synapsid pectoral girdle with greater confidence, as we demonstrate by inferring shoulder muscle PCSAs in the fossil non-mammalian cynodont Massetognathus pascuali . 

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3. The fossil record of appendicular muscle evolution in Synapsida on the line to mammals: Part II —Hindlimb

   https://doi.org/10.1002/ar.25310  

   Bishop, Peter J; Pierce, Stephanie E (September 2023, The Anatomical Record)

   Abstract This paper is the second in a two‐part series that charts the evolution of appendicular musculature along the mammalian stem lineage, drawing upon the exceptional fossil record of extinct synapsids. Here, attention is focused on muscles of the hindlimb. Although the hindlimb skeleton did not undergo as marked a transformation on the line to mammals as did the forelimb skeleton, the anatomy of extant tetrapods indicates that major changes to musculature have nonetheless occurred. To better understand these changes, this study surveyed the osteological evidence for muscular attachments in extinct mammalian and nonmammalian synapsids, two extinct amniote outgroups, and a large selection of extant mammals, saurians, and salamanders. Observations were integrated into an explicit phylogenetic framework, comprising 80 character–state complexes covering all muscles crossing the hip, knee, and ankle joints. These were coded for 33 operational taxonomic units spanning >330 Ma of tetrapod evolution, and ancestral state reconstruction was used to evaluate the sequence of muscular evolution along the stem lineage from Amniota to Theria. The evolutionary history of mammalian hindlimb musculature was complex, nonlinear, and protracted, with several instances of convergence and pulses of anatomical transformation that continued well into the crown group. Numerous traits typically regarded as characteristically “mammalian” have much greater antiquity than previously recognized, and for some traits, most synapsids are probably more reflective of the ancestral amniote condition than are extant saurians. More broadly, this study highlights the utility of the fossil record in interpreting the evolutionary appearance of distinctive anatomies. 

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4. Can Geometric Morphometric Analyses of Limb Shape Reveal Ecomorphological Patterns Across the Evolutionary History of Synapsida?

   Lungmus, J. K.; Angielczyk, K. D. (April 2019, Journal of morphology)

   Extant Mammalia are the only living representatives of the larger clade known as Synapsida, which has a continuous fossil record from around 320 million years ago to today. Despite the fact that much of the ecological diversity of mammals has been considered in light of limb morphology, the deep time origin of synapsid limb diversity and its influence on ecological diversity has received less attention. Here, we present shape analyses focusing on the forelimbs of the two earliest synapsid radiations (“pelycosaurs”, and pre-mammaliaforme Therapsida) in comparison to a broad sample of extant Mammalia. Using an expansive geometric morphometric data set, comprised of 384 fossil specimens and 148 extant mammalian specimens, we sought evidence for ecomorphological signals that could provide insight on the ecology of the earliest synapsids. Collecting shape data of humeral and ulnar elements from an extant sample representing multiple known eco morphologies provided the framework for a comparative exploration of extinct ecomorphologies, associated specifically with locomotion. Our results show that distal humeral shape is not informative of broad locomotor ecomorphologies in early fossil Synapsida. In contrast, proximal humeral shape shows a more complex pattern that suggests shape similarity between basal synapsids and members of extant Perissodactyla, and certain highly derived fully fossorial mammals, as just two examples. Overall, however, our findings suggest general shape analyses may have limited utility when analyzing for ecological-signal across deep time. Considering skeletal morphology in a holistic framework that considers unique combinations of shapes, as well as the use of biomechanically focused indices (such are functional proportions), may help to elucidate the more nuanced ways that locomotor ecology influenced limb shape in some of the earliest amniote radiations. 

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5. Shoulder joint range of motion in fossil synapsids and the origins of mammalian locomotor diversity

   Brocklehurst, R. J. (July 2020, Journal of Vertebrate Paleontology, Program and Abstracts, 2020)

   null (Ed.)

   Extant mammals are both taxonomically and ecologically diverse, having evolved a remarkable array of locomotor ecologies (e.g., swimming, digging, and flying). Evolution of the therian-type forelimb, with a highly reduced pectoral girdle and ball-and-socket shoulder joint, has been heralded as a key innovation that enabled mammals to co-opt their forelimbs for diverse functions. The acquisition of the mammal forelimb can be traced through their forerunners, the non-mammalian synapsids (NMS), but exactly how this musculoskeletal transformation proceeded and its impact on functional diversification have not be quantitatively tested. To explore the evolution of forelimb functional diversity in synapsids, we measured shoulder joint osteological range of motion (ROM) in a range of extant amniotes (lizards, monotremes, therian mammals), and compared their patterns of joint mobility to exemplars from each of the major grades of NMS: ‘pelycosaurs’, basal therapsids, and non-mammalian cynodonts. Three-dimensional models of the shoulder girdles and humeri were digitally aligned in an anatomical ‘neutral pose’ using a semi-automated approach based on articular surface morphology. ROM was then determined for the shoulder joint using a fully automated method, where the humerus was moved in flexion-extension, adduction-abduction, and pronation-supination until bone-to-bone contact occurred. Relative degree and directionality of mobility were then compared across taxa. We find an increase in total shoulder joint ROM through synapsid evolution, suggesting that more derived NMS could perform a wider range of limb movements. However, we also see more complex trends in directionality of shoulder mobility that may be indicators of forelimb posture. Extant lepidosaurs and monotremes had the greatest ROM in abduction-adduction, whereas therians had more ROM in flexion-extension, likely related to ‘sprawling’ vs. ‘erect’ gaits. Therapsids and cynodonts both had greatest ROM in abduction-adduction, matching previous reconstructions of these taxa as sprawling to semi-erect. However, ‘pelycosaurs’ had the greatest ROM in flexion-extension, despite having abducted forelimbs, suggesting they did not move their forelimbs in same manner as modern sprawling animals. Our results demonstrate the complex nature of forelimb evolution in synapsids and provide novel insights into the functional transformation and diversification of the mammalian forelimb. Funding Sources Funding information: NSF DEB- 1757749 (S.E.P) and NSF DEB-1754502 (K.D.A). 

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