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A Digital Endocranial Cast of the Early Paleocene (Puercan) 'Archaic' Mammal Onychodectes Tisonensis (Eutheria: Taeniodonta)

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A Digital Endocranial Cast of the Early Paleocene (Puercan) 'Archaic' Mammal Onychodectes Tisonensis (Eutheria: Taeniodonta) J Mammal Evol DOI 10.1007/s10914-017-9381-1 ORIGINAL PAPER A Digital Endocranial Cast of the Early Paleocene (Puercan) ‘Archaic’ Mammal Onychodectes tisonensis (Eutheria: Taeniodonta) James G. Napoli1,2 & Thomas E. Williamson3 & Sarah L. Shelley 2 & Stephen L. Brusatte2,3 # The Author(s) 2017. This article is published with open access at Springerlink.com Abstract Eutherian mammals—placentals and their closest olfactory bulb dimensions are not strongly correlated with body extinct relatives—underwent a major radiation following the size or fossorial digging for shelter, but relative bulb width is end-Cretaceous extinction, during which they evolved dispa- significantly greater in taxa that habitually dig to forage for rate anatomy and established new terrestrial ecosystems. food. The anatomical description and statistical results allow Much about the timing, pace, and causes of this radiation us to present an ecological model for Onychodectes and similar remain unclear, in large part because we still know very little taeniodonts, in which they are animals of simple behavior about the anatomy, phylogenetic relationships, and biology of that rely on a strong sense of smell to locate buried food the so-called ‘archaic’ eutherians that prospered during the before extracting and processing it with their specialized ~10 million years after the extinction. We describe the first skeletal anatomy. digital endocranial cast of a taeniodont, a bizarre group of eutherians that flourished in the early Paleogene, reconstruct- Keywords Taeniodonta . K-Pg extinction . Paleocene . ed from a computed tomography (CT) scan of a late Puercan Digital endocast . Mammal radiation (65.4 million year old) specimen of Onychodectes tisonensis that recovered most of the forebrain and midbrain and portions of the inner ear. Notable features of the endocast include long, Introduction broad olfactory bulbs, dorsally-positioned rhinal fissures, and a lissencephalic cerebrum. Comparison with other taxa shows In the wake of the end-Cretaceous (K-Pg) mass extinction, that Onychodectes possessed some of the largest olfactory mammals dramatically diversified, filling newly-open niche bulbs (relative to cerebral size) of any known mammal. space vacated by the dinosaurs (e.g., Simpson 1949;Romer Statistical analysis of modern mammals shows that relative 1966; Rose 2006; Wible et al. 2007;O’Leary et al. 2013). This is an exemplary radiation in the fossil record, and it set the stage for the mammal-rich world of today. It is clear that one Electronic supplementary material The online version of this article — (doi:10.1007/s10914-017-9381-1) contains supplementary material, particular type of mammal the eutherians, which include which is available to authorized users. placentals and their closest extinct relatives—blossomed in the post-extinction world, and their dominance continues to * James G. Napoli the present. But many questions remain about the timing, [email protected] pace, and drivers of the eutherian radiation (e.g., Archibald and Deutschman 2001;Goswami2012; dos Reis et al. 2014; 1 Department of Earth, Environmental, and Planetary Sciences, Brown Grossnickle and Newham 2016). In large part, this is because University, 69 Brown Street, Box 5385, Providence, Rhode we still know very little about the anatomy, phylogenetic re- Island 02912, USA lationships, and biology of the eutherians that prospered dur- 2 School of GeoSciences, Grant Institute, University of Edinburgh, ing the ~10 million years after the extinction, in the Paleocene James Hutton Road, Edinburgh EH9 3FE, Scotland, UK (Rose 2006). 3 New Mexico Museum of Natural History and Science, 1801 Most eutherian mammals that diversified during the early Mountain Road, NW, Albuquerque, NM 87104-1375, USA Paleocene were so-called ‘archaic’ species, which were clearly J Mammal Evol larger and more diverse than the Cretaceous eutherians but Methods whose relationships to the modern placental orders are poorly understood (Rose 2006). Among the most familiar Specimen and distinctive of the ‘archaic’ groups are the taeniodonts, a clade of small-to-medium-sized mammals endemic to This study is based on AMNH 785, a nearly-complete skull of North America that originated in the Late Cretaceous, sur- Onychodectes tisonensis that was collected from fossil zone B vived the extinction, and proliferated in the Paleocene of De-Na-Zin Wash of the Bisti/De-Na-Zin Wilderness Area (Schoch 1986; Lucas et al. 1998;FoxandNaylor2003). of northwestern New Mexico, USA (Williamson 1996). This Taeniodonts were among the first mammals to evolve den- horizon represents the classic BPolymastodon horizon^ tal adaptations for feeding on tough vegetation (e.g., crown (Polymastodon = Taeniolabis) of Sinclair and Granger hypsodonty) and postcranial specializations for scratch (1914) and yields the type fauna of the late Puercan biozone digging and/or burrowing. At least nine valid taeniodont of the Puercan North American Land Mammal Age (Lofgren genera are currently recognized (Schoch 1986; Rook and et al. 2004). Preliminary results from 40Ar/39Ar geochronolo- Hunter 2014; Williamson and Brusatte 2013). They became gy from ash and detrital sanidine closely associated with fossil larger and more specialized over time (e.g., Patterson 1949), zone B indicate that it dates from within the first 500,000 years culminating in the last-surviving middle Eocene Stylinodon, of the Paleocene (Heizler et al. 2013). which was about 50–100 kg in body mass and possessed The cranium AMNH 785 was collected by a crew from the rootless teeth, large gliriform canines, and powerful forelimbs American Museum of Natural History led by Joseph Wortman with enlarged claws (Schoch 1986; Lucas et al. 1998; in 1892, and was initially described by Osborn and Earle Turnbull 2004). (1895); it was later redescribed by both Matthew (1937)and Over the past few decades, the discovery of important Schoch (1986). The cranium of AMNH 785 is crushed and new fossils of taeniodonts and other ‘archaic’ mammals mediolaterally distorted, and is broken in several locations. has helped to elucidate some aspects of their paleobiology, Both zygomatic arches, most of the teeth, and the occiput whichinturnisstartingtogiveinsightintotheirevolution- are missing entirely, along with a portion of the posterior skull ary radiation (e.g., Williamson 1996). One area that has roof. The preservation of the skull made it impossible to been little explored, however, is the neuroanatomy and completely reconstruct neural anatomy, but many of the major neurosensory capabilities of these animals. The now wide- anatomical features are visible. spread use of computed tomography (CT) scanning in pa- leontology has allowed for the visualization of internal CT Scanning cranial features and the reconstruction of brains, inner ears, and cranial vasculature, which has revolutionized the field The specimen was scanned by Dr. Hong-yu Yi with a GE (e.g., Witmer et al. 2008). However, few ‘archaic’ Paleocene phoenix v|tome|x micro-CT scanner at the American mammals have been studied this way, so we still know Museum of Natural History Microscopy and Imaging very little about their brains and sense organs, how their Facility. It was scanned with the following parameters: voltage neurosensory abilities compared to modern mammals, and of 170–220 kV, current of 150–260 μA, and voxel size how their neurobiology may have been related to their 51.1 μm. post-Cretaceous radiation. Here, we present the first CT-based study of the brain- CT Reconstruction case of a taeniodont. Using high-resolution X-ray CT, we digitally visualize and describe the endocranial cast of the A dataset of tiff images from the CT scan was imported into basal taeniodont Onychodectes tisonensis.Inmammals, Materialize Mimics 17.0 (Materialize N.V. 2014)atthe the endocranial space is filled with the brain and other University of Edinburgh School of GeoSciences. This was soft-tissue structures, which leave an impression on the done to visualize and examine the internal structures of the internal surface of the surrounding skull bones (Jerison braincase and to construct three-dimensional digital render- 1973;Macrinietal.2007). An endocast, therefore, gives ings of the specimen. This work was carried out by JGN. an indication of the general shape and volume of the The slices containing the braincase were manually seg- brain. Because the brain is responsible for controlling mented into several label maps. While the majority of the andregulatingthebody,andplaysanimportantrolein anterior braincase was preserved, there were many slices most organismal functions, including sensory processing in which a section of it was missing, or where cracks and analysis, motor control, and learning, studying its size traversed the entirety of the skull. In these cases, a straight and shape provides vital clues to the neurosensory abili- line was drawn between preserved edges. We then used ties of this critical mammal that thrived soon after the Mimics to generate surface files from the label maps, and end-Cretaceous extinction. smoothed the surfaces to reduce noise and allow proper J Mammal Evol Table 1 Comparisons of brain measurements among taxa included in given in mm. Body mass estimates in kg. Extinct species are not scored comparison set. Abbreviations as follows: OBL, olfactory bulb length; for forage style or mode of life. All data for extant species come from the OBW, olfactory bulb width; CL, cerebrum length; CW, cerebrum width; Major National Resources
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