Abstract
The relationship between ecology and morphology of the limbs in living placental mammals is well established and has been used to infer aspects of palaeobiology for many extinct species. However, few studies have applied these principles to extinct marsupials. Palorchestids are a poorly understood extinct family of megafaunal marsupials of particular interest due to their highly robust and unusual limb morphology. Using a comparative sample of humeri and femora from living mammals potentially analogous to palorchestids, we applied three-dimensional geometric morphometric techniques in a phylogenetic comparative framework. We established that humerus and femur shape in living species showed significant associations with size, phylogeny, and substrate use, and found a weak association between humeral shape and diet. We then examined patterns in morphological disparity, modularity, covariance and morphospace occupation in palorchestids relative to comparative living mammals as well as their closest extinct marsupial relatives. We found palorchestid femora to be unremarkable in shape, while their overall and proximal humeral morphology were strongly divergent from all other mammals sampled. Over their evolutionary history, palorchestid distal humeri increasingly resembled those of mammals adapted for tearing and hook-and-pull digging, while other analyses showed various arboreal-like and fossorial-like affinities in humeral shape. Our findings indicate strong asymmetric selection acting on the fore- and hindlimbs in palorchestids, and their unique combination of shape traits suggests they used their forelimbs in a specialised manner that has no direct equivalence either with their extinct relatives or among other mammals alive today.
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Data availability
All code and data associated with this study are available in the Online Resources and from Figshare (10.26180/19946171), with specimen 3D meshes available on Morphosource.org where possible (project ID: 000445426).
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Acknowledgements
For access to museum collections, we thank: Sandy Ingleby & Matt McCurry (Australian Museum); Marisa Suvoy, Sara Ketelsen & Eleanor Hoeger (American Museum of Natural History); Adam Yates (Museum and Art Gallery of the Northern Territory); Tim Ziegler, Katie Date, Karen Roberts & Ricky-Lee Erickson (Museums Victoria); Tammy Gordon (Queen Victoria Museum and Art Gallery); Mary-Anne Binnie (South Australian Museum); Rolan Eberhard & Nikki King-Smith (Tasmanian Museum and Art Gallery). We are grateful to Stephanie Pierce and Peter Bishop (Harvard University) for access to Didelphis meshes, Jacob Van Zoelen and Gavin Prideaux for sharing meshes of Kolopsis and Plaisiodon from the MAGNT collection, and to Alex McDonald for CT scanning several extant marsupials used in this study. CT scanning was performed with the valuable assistance of Dr Michael de Veer at Monash Biomedical Imaging. The authors acknowledge the facilities and scientific and technical assistance of the National Imaging Facility (NIF), a National Collaborative Research Infrastructure Strategy (NCRIS) capability at Monash Biomedical Imaging (MBI), a Technology Research Platform at Monash University. Additional meshes were sourced from online databases (Morphosource.org, Sketchfab.com, Digital Morphology Museum KUPRI), detailed in Online Resource 1. HLR thanks Kathleen Garland, Tahlia Pollock and James Rule for their generous advice on data analysis and phylogenetic comparative methods, Laura Wilson for methods assistance and comments on as earlier draft. We are also grateful to Nestor Toledo and an anonymous reviewer for their input that greatly improved this paper.
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This research was supported by an Australian Government Research Training Program Scholarship and Monash University-Museums Victoria PhD Research Scholarship (HLR), internal funds from the Department of Anatomy and Developmental Biology, Monash University (JWA), and the Australian Research Council DP180101797 (ARE).
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HLR led the study, gathered and processed data, performed analyses, constructed figures and drafted the manuscript. HLR, DSR and ARE wrote code and interpreted results. JWA and ARE provided supervision. All authors contributed to and approved the final manuscript text.
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Richards, H.L., Rovinsky, D.S., Adams, J.W. et al. Inferring the palaeobiology of palorchestid marsupials through analysis of mammalian humeral and femoral shape. J Mammal Evol 30, 47–66 (2023). https://doi.org/10.1007/s10914-022-09640-6
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DOI: https://doi.org/10.1007/s10914-022-09640-6