Morphology of the Bony Labyrinth of Select Species of the Superorder Xenarthra Christopher T Wilkinson | Joe Sertich PhD MSMHA MS Candidate 2015 | DMNS Background & Rationale Figure 1. The currently accepted phylogenetic relationships among Xenarthra. The genera Choloepus and Bradypus are highlighted in pink. Other genera looked at in this study are highlighted in green. Figure adapted from Gaudin, 2003 • Sloths, Anteaters, and Armadillos are all part of the superorder Xenarthra • There are only two extant tree sloth genera (Bradypus and Choloepus) • Distantly related, (Fig. 1) but share many traits: • Appearance • Shoulder morphology7 • Habitat9 • Inner Ear structure2 • Previous studies have established adaptations specific to the inner ear of extant tree sloths1,2 • However, they don’t resemble the inner ear of other extant Xenarthrans • A recent study looked at Megalonyx jeffersoni (an extinct ground sloth) to examine whether the tree sloth inner ear morphology was shared with extinct ground sloths1 • The inner ear of Megalonyx jeffersoni more resembles the inner ear of an anteater than that of a tree sloth (either genera)1 • Goals of this study: • Increase the number of models of Xenarthran inner ears • Take a closer look at the inner ear morphology of supposed ground sloths • Hypothesis- The inner ear morphology of tree sloths is unique among Xenarthrans. Project Approach • Obtained CT scans of Xenarthran skulls and isolated temporal bones currently in collections at the Denver Museum of Nature and Science • Modeled 46 bony labyrinths • The Half Maximum Height method was used to estimate the border between air and bone3,5,8 • Used these HMH values to segment out the bony labyrinth in Avizo4 • Fig. 2 illustrates landmarks as they appear in coronal slices • All bony labyrinths were modeled by the same observer • Measurements were made in Geomagic Design X 64* • A first PCA was conducted looking at all measured traits • A second PCA was conducted looking at • the relationship between ampulla and canal diameter (3 measurements) • the relationship between canal size and total inner ear size (3 measurements) *All measurements can be found in the electronic supplementary materials along with additional graphics displaying how measurements were made in Geomagic Figure 2. (left) This figure displays select slices from a sequence of scans of the right inner ear of a Tamandua mexicana. The number in the lower right corner of each screen capture is the slice number and the labels correspond to landmarks as they appear in CT cross sections. All of these images are taken from coronal sections and the full bony labyrinth in this specimen was more than 250 slices. IAM-Internal acoustic meatus. SSC superior semicircular canal. SSCa-Superior semicircular canal ampulla. LSC lateral semicircular canal. LSCa lateral semicircular canal ampulla.PSC-posterior semicircular canal. PSCa-posterior semicircular canal ampulla. Results Results • 26 CT scans Scans of the temporal region were taken for each specimen(Fig. 3) • 46 inner ear models Both inner ears were modeled whenever possible (Fig. 3) • A visual comparison shows the specimens’ bony labyrinths falling in one of two categories (Fig. 4) • Tree sloth ear type • Canals that vary greatly from an orthogonal orientation • Canals whose cross sectional diameter is close to the cross sectional diameter of their ampulla • Canals whose overall size is small relative to the inner ear size (RATIO). • Anteater ear type • Canals that are roughly orthogonal • Canals that are much smaller in cross sectional diameter then their corresponding ampulla • Canals whose overall size is large relative to the total inner ear size. • The first PCA (Fig. 7 A) was run using twelve factors (SSCR, PSCR, LSCR, LSC/ PSC angle, LSC/SSC angle, SSC/PSC angle, LSC c/a ratio, PSC c/a ratio, SSC c/a ratio, IEH, Coc turns, and CocR) • Divided the specimens into four groups • tree sloths • ground sloths • anteaters • armadillos • The tree sloths separated from the other Xenarthrans with only a slight overlap • The second PCA (Fig. 7 B) used only six factors, looking specifically at the size of the semicircular canals (LSCR, SSCR, PSCR, LSC c/a ratio, PSC c/a ratio, and SSC c/a ratio). • This separates the tree sloths even more clearly • The three leftmost points within the ground sloth grouping are all Acratocnus B Figure 3. (Right )The bony labyrinth of a Choloepus hoffmani within a transparent cranium. Only half of the skull was scanned to obtain the highest resolution possible. The top picture shows posterior view of the cranium and the bottom picture shows a right lateral view. A D A Figure 7. Part A shows the PCA using twelve factors while part B looks specifically at factors related to SSC size Conclusions • The tree sloth inner ear is represented by a distinct set of adaptations • Acratocnus (classified as a ground sloth) shares these adaptations • Acratocnus likely exhibits a locomotor repertoire similar to extant tree sloths References & Acknowledgements B 1) Billet G, Germain D, Ruf I, de Muizon C, and Hautier L. 2013. The inner ear of Megatherium and the evolution of the vestibular system in sloths. J. Anat. 557-567. 2) Billet G, Hautier L, Asher R. 2012. High morphological variation of vestibular system accompanies slow and infrequent locomotion in three-toed sloths. Proc Biol Sci 279: 3932-3939. E C 3) Coleman M and Colbert M. 2007. Technical note: CT thresholding protocols for taking measurements on three-dimensional models. Am J Phys Anthropol 133: 723-725. 4) Ekdale E. 2013. Comparative anatomy of the bony labyrinth (inner ear) of placental mammals. PLoS ONE 8: 1-100. 5) Fajardo R, Ryan T, and Kappelman J. 2002. Assessing the accuracy of high-resolution X-ray commuted tomography of primate trabecular bone in comparisons with histological sections. Am J Phys Anthropol 118: 1-10. 6) Gaudin T. 2003. Phylogenetic relationships among sloths (Mammalia, Xenarthra, Tardigrada): the craniodental evidence. Zool J Linn Soc 255-305. 7) Nyakatura J, and Fischer M. 2011. Functional morphology of the muscular sling at the pectoral girdle in tree sloths: convergent morphological solutions to new functional demands. J. Anat 219: 360-374. 8) Spoor C, Zonneveld F, and Macho G. 1993. Linear measurements of cortical bone and dental enamel by computed tomography: applications and problems. Am J Phys Anthropol 91: 469-484. 9) Superina M. 2006. Biology of Xenarthra. Edentata 7:59-60. Figure 4. This figure depicts the bony labyrinth models created for five different genera. The specimens grouped on the left have the tree sloth characteristics, those on the right hand side have the anteater type characteristics. The species depicted are: Choloepus hoffmani (A), Bradypus variegatus (B), Acratocnus (C), Megalonyx jeffersoni (D), and Tamandua Mexicana (E). The authors thank Kendra Huber and Ramesh Karki for assisting with data collection and analysis. Support by NIH AG090000 to M.C. McGwire.
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