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1. Trabecular bone confers different functional advantages across small mammals

   Smith, Stephanie M; Angielczyk, Kenneth D; Stayton, Tristan (March 2025, Society for Integrative and Comparative Biology)

   The relative contributions of trabecular (spongy) and cortical (compact) bone to bone strength and stiffness are poorly understood across mammalian body size. In mammals, some small species have notably reduced their vertebral trabecular bone structure, resulting in mostly hollow medullary cavities. To assess the importance of trabecular structure to the mechanical properties of small mammalian vertebrae, we conducted finite element analysis on the lumbar vertebrae of 25 species of shrews (Soricidae) weighing 2-100g. We analyzed two sets of models: vertebrae with the trabecular structure intact (full), and vertebrae with all trabeculae excised from the centrum (hollow). All models were scaled to the same ratio of load to surface area. The cranial end of the centrum was immobilized, and a 5N craniocaudally-oriented load was applied to the caudal end of the centrum. We measured mean von Mises stress (MVMS) to capture strength, and total strain energy to capture stiffness. MVMS and total strain energy both decrease as body size increases, and hollow models experience higher stresses and strains than full models. With increasing body size, the difference in total strain energy between full and hollow models decreases, but the difference in MVMS slightly increases. This suggests a difference in the functional advantage conferred by trabeculae among small mammals, as well as a possible selective pressure for different functional emphasis in very small and larger mammalian bones. 

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

   https://doi.org/10.1002/ar.24817  

   Zack, Ellianna H.; Smith, Stephanie M.; Angielczyk, Kenneth D. (November 2021, The Anatomical Record)

   null (Ed.)

   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. We collected μCT scans of the last six presacral vertebrae in 13 fossorial, terrestrial, and suspensorial xenarthran species (body mass: 120 g to 35 kg). For each vertebra, we measured centrum length; bone volume fraction (BV.TV); trabecular number and mean thickness (Tb.Th); global compactness (GC); cross-sectional area; mean intercept length; star length distribution; and connectivity and connectivity density. Wild specimens have more robust trabeculae, but this varies with species, ecology, and pathology. Wild specimens of fossorial taxa (Dasypus) have more robust trabeculae than captives, but there is no clear difference in bone microstructure between wild and captive specimens of suspensorial taxa (Bradypus, Choloepus), suggesting that locomotor ecology influences the degree to which captivity affects bone microstructure. Captive Tamandua and Myrmecophaga have higher BV.TV, Tb.Th, and GC than their wild counterparts due to captivity-caused bone pathologies. Our results add to the understanding of variation in mammalian bone microstructure, suggest caution when including captive specimens in bone microstructure research, and indicate the need to better replicate the habitats, diets, and behavior of animals in captivity. 

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3. Effects of captivity on the bone microstructure of xenarthrous vertebrae

   Zack, Ellianna H; Smith, Stephanie M; Angielczyk, Kenneth D (March 2021, Integrative and comparative biology)

   Captive specimens in natural history collections allow researchers to inspect the morphologies of rare taxa, but the lifestyles,diets, and lifespans of captive animals differ from those of their wild counterparts. To quantify these differences, we compared bone microstructure of trunk vertebrae in captive and wild xenarthran mammals (sloths, armadillos, anteaters). Because trabecular bone architecture (TBA) adapts to in vivo forces, bone microstructure reflects ecology and behavior, but this means that it may differ between captive and wild specimens of the same species. We collected μCT scans of the last six presacral vertebrae in 13 species of fossorial, terrestrial, and suspensorial xenarthrans ranging in body mass from 120g (Chlamyphorus) to 35kg (Myrmecophaga). For each vertebra, we measured bone volume fraction (BVF); trabecular number, mean thickness (TbTh), and orientation; global compactness; and cross sectional area. Wild specimens generally have more robust trabeculae, but this differs based on species, vertebral position, ecology, and pathology. The wild specimens of fossorial taxa (Dasypus) have more robust trabeculae than their captive counterparts, but there is no clear difference in TBA of wild and captive specimens in suspensorial and terrestrial taxa (Bradypus, Choloepus, Cyclopes). These data suggest that locomotor ecology affects the level to which captivity affects bone microstructure. The captive specimens of both Tamandua and Myrmecophaga have higher BVF and TbTh than their wild counterparts, indicating more brittle trabeculae due to bone pathologies caused by captivity. Our results add to the overall understanding of variation in mammalian bone microstructure and suggest caution when including captive specimens in research on TBA. 

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4. Adventures inside shrew vertebrae: trabecular bone morphology and regionalization in Soricidae

   Smith, Stephanie M; Angielczyk, Kenneth D (March 2021, Integrative and comparative biology)

   The regionalized vertebral column is a hallmark of mammalian morphology and reflects functional differentiation of the vertebral regions. Mammalian vertebrae are serially homologous and morphologically patterened by Hox expression, but also vary in number and gross morphology across species. The trabecular bone inside vertebral centra is more plastic than gross vertebral bone, and structurally adapts to better withstand forces it experiences during life. However, the functional regionalization of vertebral trabecular bone is poorly examined. Are there trabecular "regions” reflecting the differing functions and in-vivo stress patterns of gross morphological vertebral regions? Or is trabecular morphology homogeneous throughout the spine, suggesting that differences in functional demands are borne exclusively by external characteristics? To address these questions, we collected μCT scans and linear measurements of cervical, thoracic, and lumbar vertebrae in four species of large shrews, including two species of the hero shrew Scutisorex, which has a highly modified vertebral column. We compared linear measurements and trabecular bone characteristics of the cranial and caudal ends of each centrum across species. To detect unique vertebral regions, we executed principal coordinates analysis and segmented regression on three versions of our data set: trabecular bone data only, external measurements only, and the two combined. We found that some regionalization is recovered using only trabecular bone data, but trabecular bone regions do not correspond exactly to gross vertebral regions. This reflects divergence between the functional signals of internal and external vertebral bone morphology, which should be further examined in a kinematic context. 

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5. The roles of phylogeny, body size and substrate use in trabecular bone variation among Philippine ‘earthworm mice’ (Rodentia: Chrotomyini)

   https://doi.org/10.1093/biolinnean/blad033  

   Smith, Stephanie M; Rowsey, Dakota M; Nations, Jonathan A; Angielczyk, Kenneth D; Heaney, Lawrence R (July 2023, Biological Journal of the Linnean Society)

   Abstract Trabecular bone is modelled throughout an animal’s life in response to its mechanical environment, but like other skeletal anatomy, it is also subject to evolutionary influences. Yet the relative strengths of factors that affect trabecular bone architecture are little studied. We investigated these influences across the Philippine endemic murine rodent clade Chrotomyini. These mammals have robustly established phylogenetic relationships, exhibit a range of well-documented substrate-use types, and have a body size range spanning several hundred grammes, making them ideal for a tractable study of extrinsic and intrinsic influences on trabecular bone morphology. We found slight differences in vertebral trabecular bone among different substrate-use categories, with more divergent characteristics in more ecologically specialized taxa. This suggests that the mechanical environment must be relatively extreme to affect trabecular bone morphology in small mammals. We also recovered allometric patterns that imply that selective pressures on bone may differ between small and large mammals. Finally, we found high intrataxonomic variation in trabecular bone morphology, but it is not clearly related to any variable we measured, and may represent a normal degree of variation in these animals rather than a functional trait. Future studies should address how this plasticity affects biomechanical properties and performance of the skeleton. 

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