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Deciduous Dentition and Dental Eruption of Hyainailouroidea (Hyaenodonta, “Creodonta,” Placentalia, Mammalia)

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Deciduous Dentition and Dental Eruption of Hyainailouroidea (Hyaenodonta, “Creodonta,” Placentalia, Mammalia) Palaeontologia Electronica palaeo-electronica.org Deciduous dentition and dental eruption of Hyainailouroidea (Hyaenodonta, “Creodonta,” Placentalia, Mammalia) Matthew R. Borths and Nancy J. Stevens ABSTRACT Deciduous dental morphology and the sequence of permanent dental eruption offer developmental and phylogenetic insights into mammalian evolution. Early life his- tory in Hyaenodonta, a morphologically diverse and biogeographically widespread clade of carnivorous mammals known from the Paleogene and early Neogene, has only been examined closely in one genus: Hyaenodon. Here we describe the decidu- ous dental morphology for six hyainailouroids, a clade of hyaenodonts distantly related to Hyaenodon that dominated early Cenozoic meat-eating niches of Afro-Arabia for nearly 20 Ma. Results suggest significant variation among hyainailouroids in expres- sion of the dP3 and dP4 protocone, paracone, and metacone and in expression of the dP3 and dP4 paraconid, metaconid, and talonid. Deciduous dental characters were incorporated into a Bayesian phylogenetic analysis. The topology was congruent with earlier systematic examinations of hyaenodont evolution, suggesting that deciduous dental features can be phylogenetically informative and that taxa known only from deciduous teeth can be incorporated into studies of mammalian systematics. Based on the specimens described herein, the dental eruption sequence of Hyainailouroidea is reconstructed. P4 and P3 erupt after M3, evidence that the deciduous carnassial com- 3 plex formed between dP and dP4 was retained until late in development, permitting hyaenodonts to heavily wear shearing blades on these teeth before replacement. This differs from Carnivora, a group that tends to replace deciduous dentition earlier in development, effectively limiting carnivorans to a single set of carnassials throughout most of their lifespan. This developmental difference between carnivorans and hyaeno- donts can now be examined in a larger systematic and ecological context. Matthew R. Borths. Department of Biomedical Sciences, Heritage College of Osteopathic Medicine, Ohio University, Athens, Ohio 45701, USA; Center for Ecology and Evolutionary Studies, Ohio University, Athens, Ohio 45701, USA. [email protected] Nancy J. Stevens. Department of Biomedical Sciences, Heritage College of Osteopathic Medicine, Ohio University, Athens, Ohio 45701, USA; Center for Ecology and Evolutionary Studies, Ohio University, Athens, Ohio 45701, USA. [email protected] Borths, Matthew R. and Stevens, Nancy J. 2017. Deciduous dentition and dental eruption of Hyainailouroidea (Hyaenodonta, “Creodonta,” Placentalia, Mammalia). Palaeontologia Electronica 20.3.55A: 1-34. https://doi.org/10.26879/776 palaeo-electronica.org/content/2017/2056-deciduous-hyaenodont-teeth Copyright: November 2017 Society for Vertebrate Paleontology. This is an open access article distributed under the terms of Attribution-NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0), which permits users to copy and redistribute the material in any medium or format, provided it is not used for commercial purposes and the original author and source are credited, with indications if any changes are made. creativecommons.org/licenses/by-nc-sa/4.0/ BORTHS & STEVENS: DECIDUOUS HYAENODONT TEETH Keywords: Ontogeny; Carnivory; Egypt; Africa; Paleogene Submission: 26 April 2017 Acceptance: 31 October 2017 INTRODUCTION Afro-Arabia, making it the longest-lived radiation of meat-eating placental mammals besides Carnivora Fossilized remains of animals are often inter- (Lewis and Morlo, 2010). Parsing out the evolution, preted by paleontologists as phenotypes shaped life history, and extinction of Hyaenodonta is critical by natural selection (e.g., Stern and Susman, for building a larger understanding of the evolution 1983; Shubin et al., 2006; Tseng et al., 2016), with of the niches occupied by modern carnivorous hypotheses surrounding adaptation, niche occupa- mammals. Previous work on the dental develop- tion, evolution, and extinction of a species typically ment of hyaenodonts focused on the genus Hyae- examined through the lens of the adult phenotype nodon, a genus from the Eurasian and North (e.g., Ross, 1996; Finarelli and Flynn, 2009; American clade Hyaenodontinae (Mellet, 1977; Goswami and Polly, 2010). Yet examination of Bastl et al., 2011; Bastl and Nagel, 2013; Bastl et extant organisms demonstrates that natural selec- al., 2014); deciduous dental materials preserved in tion acts throughout the life cycle, and fitness is other hyaenodonts (e.g., Lesmesodon, Cynohyae- significantly impacted by developmental stages nodon, Paroxyaena) have received somewhat less that precede maturity (Lindström, 1999). Hence the attention (but see Morlo and Habersetzer, 1999; adaptive strategies employed by a species or lin- Lavrov, 2007). Recent systematic research demon- eage can only truly be understood through deeper strates that Hyaenodon is only distantly related to knowledge of the ontogeny of a species or lineage Hyainailouroidea, a separate hyaenodont clade (Sánchez-Villagra, 2010a; Codron et al., 2013; that contains hypercarnivorous taxa and all Afro- Urdy et al., 2013; Hone et al., 2016). As a heritable Arabian hyaenodonts (Rana et al., 2015; Borths et aspect of an animal’s biology, developmental data al., 2016). But only a few deciduous teeth have is also a rich source of phylogenetic information, been described from Afro-Arabian hyaenodonts providing insight into evolutionary relationships (Morales et al., 2010; Solé et al., 2016), making it (e.g., de Queiroz, 1985; Smith, 2000; Asher and difficult to compare the life histories of Afro-Arabian Lehmann, 2008). hyainailouroids to Eurasian and North American In mammals, the reduction of dental replace- hyaenodont lineages and to use deciduous dental ment to diphyodonty has built a readily preserved morphology and dental eruption to probe phyloge- ontogenetic calibration point into the mouths of netic relationships among key taxa in this important most taxa (Anders et al., 2011). The replacement group of early Cenozoic predators. of deciduous teeth, eruption of adult dentition, and In this study, we describe and refer hyaeno- subsequent wear contribute information on the rate dont deciduous dental specimens from the late of maturation across lineages (e.g., Slaughter et Eocene-early Oligocene (Fayum Depression, al., 1974, Smith, 2000; Asher and Lehmann, 2008; Egypt) and the early Miocene (Rusinga Island and Ciancio et al., 2012; Veitschegger and Sánchez- Fort Ternan, Kenya) to five known hyainailouroid Villagra, 2016). Dental eruption sequences can be genera: Apterodon, Leakitherium, Anasinopa, Dis- compared with other proxies for developmental sopsalis, Masrasector, and Metasinopa. We com- maturity, such as long bone epiphyseal fusion pare the new deciduous dental material to (Coutinho et al., 1993; Uhen, 2000) and cranial Eurasian and North American hyaenodont materi- suture fusion (Sánchez-Villagra, 2010b; Rager et als to reconstruct dental development patterns al., 2013), to identify key developmental phases in among hyainailouroids. Building upon the efforts of extinct taxa that can be used to reconstruct life his- Bastl et al. (2014), we incorporate this new mor- tories and phylogenetic relationships (Fink, 1982; phological information into a phylogenetic analysis Smith, 1989; O’Leary et al., 2013). that includes deciduous dental characters. Finally, Hyaenodonta is a diverse radiation of carnivo- we use morphological and developmental data pre- rous placental mammals found in terrestrial eco- served in these specimens to discuss how hyaeno- systems in Eurasia, North America, and Afro- dont dental development informs our under- Arabia throughout the Paleogene (Rose, 2006). standing of life history strategies in the clade. The clade persisted into the Neogene in Asia and 2 PALAEO-ELECTRONICA.ORG MATERIALS AND DESCRIPTIVE METHODS Other included species. Apterodon altidens Schlosser, 1910; Apterodon intermedius Lange- Permissions Badré and Böhme, 2005; Apterodon langebadreae Specimens bearing DPC reference numbers Grohé et al., 2012; Apterodon macrognathus described herein are available at the Division of Andrews, 1904; Apterodon saghensis Simons and Fossil Primates at the Duke Primate Center in Gingerich, 1974. Raleigh, NC and were collected and exported with Apterodon macrognathus Andrews, 1904 permission granted by the Egyptian Mineral Figures 1-2, Tables 1-2 Resources Authority (formerly the Egyptian Geo- logical Survey and Minding Authority) and the Referred specimens. DPC 4126, right maxilla Egyptian Geological Museum. Specimens bearing fragment with dP4 (Fayum Depression, Jebel KNM reference numbers described herein are Qatrani Formation, Quarry M); DPC 8217, left den- available at the Nairobi National Museum, part of tary fragment with dP3, dP4, M1 (Fayum Depres- the National Museums of Kenya, and were exam- sion, Jebel Qatrani Formation, Quarry M). Note: ined under research permits issued by the Kenyan These specimens were not found in association. National Commission for Science, Technology and Description Innovation and as research affiliates of the National Museums of Kenya. In the maxilla of Apterodon macrognathus (DPC 4126; Figure 1) the infraorbital foramen is Institutional Abbreviations superior and slightly anterior to the mesial root of BMNH, Natural History Museum, London; DPC, dP4. The parastyle is mesiodistally short and Duke Lemur Center, Division of Fossil Primates, divided by a buccolingually
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