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1. The fossil record of appendicular muscle evolution in Synapsida on the line to mammals: Part I—Forelimb

   https://doi.org/10.1002/ar.25312  

   Bishop, Peter_J; Pierce, Stephanie_E (September 2023, The Anatomical Record)

   Abstract This paper is the first 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 forelimb. Understanding forelimb muscular anatomy in extinct synapsids, and how this changed on the line to mammals, can provide important perspective for interpreting skeletal and functional evolution in this lineage, and how the diversity of forelimb functions in extant mammals arose. 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 73 character–state complexes covering all muscles crossing the shoulder, elbow, and wrist 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. In addition to producing a comprehensive documentation of osteological evidence for muscle attachments in extinct synapsids, this work has clarified homology hypotheses across disparate taxa and helped resolve competing hypotheses of muscular anatomy in extinct species. The evolutionary history of mammalian forelimb musculature was a complex and nonlinear narrative, punctuated by multiple instances of convergence and concentrated phases of anatomical transformation. More broadly, this study highlights the great insight that a fossil‐based perspective can provide for understanding the assembly of novel body plans. 

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2. RECONSTRUCTIONS OF HINDLIMB MUSCULATURE IN EXTINCT PRE-THERIAN SYNAPSIDS

   https://doi.org/10.3099/MCZ82  

   Bishop, P J; Pierce, S E (October 2024, Bulletin of the Museum of Comparative Zoology)

   Previously, the authors published a framework for phylogenetically informed reconstruction of appendicular musculature in extinct synapsids. In the present study, this framework is employed to rigorously infer the arrangement of muscles in the hindlimb of eight exemplar taxa that capture the evolutionary transformation from basal synapsids to crown therians. Muscle maps detailing origins and insertions are presented for each taxon. These will aid the interpretation of fossil material in the future, especially fragmentary remains of related species, and provide a foundation for functional analysis of the appendicular skeleton and its evolution. 

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3. 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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4. Late acquisition of erect hindlimb posture and function in the forerunners of therian mammals

   https://doi.org/10.1126/sciadv.adr2722  

   Bishop, Peter J; Pierce, Stephanie E (October 2024, Science Advances)

   The evolutionary transition from early synapsids to therian mammals involved profound reorganization in locomotor anatomy and function, centered around a shift from “sprawled” to “erect” limb postures. When and how this functional shift was accomplished has remained difficult to decipher from the fossil record alone. Through biomechanical modeling of hindlimb force-generating performance in eight exemplar fossil synapsids, we demonstrate that the erect locomotor regime typifying modern therians did not evolve until just before crown Theria. Modeling also identifies a transient phase of increased performance in therapsids and early cynodonts, before crown mammals. Further, quantifying the global actions of major hip muscle groups indicates a protracted juxtaposition of functional redeployment and conservatism, highlighting the intricate interplay between anatomical reorganization and function across postural transitions. We infer a complex history of synapsid locomotor evolution and suggest that major evolutionary transitions between contrasting locomotor behaviors may follow highly nonlinear trajectories. 

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5. 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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