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Petrosal Anatomy of the Paleocene Eutherian Mammal Deltatherium Fundaminis (Cope, 1881)

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Petrosal Anatomy of the Paleocene Eutherian Mammal Deltatherium Fundaminis (Cope, 1881) Journal of Mammalian Evolution https://doi.org/10.1007/s10914-021-09568-3 ORIGINAL PAPER Petrosal Anatomy of the Paleocene Eutherian Mammal Deltatherium fundaminis (Cope, 1881) Sarah L. Shelley1,2 · Ornella C. Bertrand1 · Stephen L. Brusatte1,3 · Thomas E. Williamson3 Accepted: 1 August 2021 © The Author(s) 2021 Abstract We describe the tympanic anatomy of the petrosal of Deltatherium fundaminis, an enigmatic Paleocene mammal based on cranial specimens recovered from New Mexico, U.S.A. Although the ear region of Deltatherium has previously been described, there has not been a comprehensive, well-illustrated contribution using current anatomical terminology. The dental and cranial anatomy of Deltatherium is a chimera, with morphological similarities to both ‘condylarth’ and ‘cimo- lestan’ taxa. As such, the phylogenetic relationships of this taxon have remained elusive since its discovery, and it has vari- ably been associated with Arctocyonidae, Pantodonta and Tillodontia. The petrosal of Deltatherium is anteriorly bordered by an open space comprising a contiguous carotid opening and pyriform fenestra. The promontorium features both a small rostral tympanic process and small epitympanic wing but lacks well-marked sulci. A large ventral facing external aperture of the canaliculus cochleae is present and bordered posteriorly by a well-developed caudal tympanic process. The hiatus Fallopii opens on the ventral surface of the petrosal. The tegmen tympani is mediolaterally broad and anteriorly expanded, and its anterior margin is perforated by a foramen for the ramus superior of the stapedial artery. The tympanohyal is small but approximates the caudal tympanic process to nearly enclose the stylomastoid notch. The mastoid is widely exposed on the basicranium and bears an enlarged mastoid process, separate from the paraoccipital process. These new observations provide novel anatomical data corroborating previous hypotheses regarding the plesiomorphic eutherian condition but also reveal subtle diferences among Paleocene eutherians that have the potential to help inform the phylogeny of Deltatherium. Keywords Paleogene · Deltatherium · Auditory region · Condylarthra · Basicranium Introduction compared to its scantily known postcrania (Cope 1881a; Matthew 1937). However, closer consideration of Deltathe- Deltatherium, meaning ‘triangular-beast’ in reference to rium reveals it to be among the most enigmatic Paleocene its tritubercular teeth, is a eutherian mammal known from mammals, a chimera of morphologies similar to both ‘con- Paleocene-aged deposits in the San Juan Basin of New dylarth’ and ‘cimolestan’ taxa. As such, its phylogenetic Mexico, U.S.A (Cope 1881a; Matthew 1937; Williamson relationships have remained elusive since its discovery. 1996). At frst glance, the fossils of Deltatherium allude to Determining the phylogenetic position of Deltatherium is a rather innocuous opossum-sized animal with dental and a key step in resolving broader issues of Paleocene mammal cranial adaptations indicative of an omnivorous to carnivo- phylogeny and requires detailed anatomical descriptions to rous diet and a disproportionately large and robust head inform character data. The specifc objective of this study is to describe the Deltatherium fundaminis * Sarah L. Shelley tympanic petrosal anatomy of . [email protected] Matthew (1937) briefy described the auditory region of Deltatherium in his seminal posthumous monograph on the 1 School of GeoSciences, University of Edinburgh, Edinburgh, Paleocene faunas of the San Juan Basin. Here, we update United Kingdom and expand upon Matthew’s description and include detailed 2 Carnegie Museum of Natural History, Pittsburgh, comparisons to other Paleogene mammals. The petrosal Pennsylvania, United States of America bone is a rich source of comparative anatomical data and 3 New Mexico Museum of Natural History and Science, continues to be a useful source of phylogenetic information. Albuquerque, New Mexico, United States of America Vol.:(0123456789)1 3 Journal of Mammalian Evolution Nevertheless, since it is not the goal of this paper to resolve of Tillodontia as well as some pantodonts (Gazin 1953). the phylogenetic position of Deltatherium, we do not lend Tillodonts are medium to large-sized eutherian mammals credence to any phylogenetic hypotheses at present and known from North America and Asia distinguished by refrain from making any statements regarding that matter. their rodent-like second upper incisors and other fea- We will, however, describe and discuss the comparative tures more reminiscent of carnivores or ungulates (Gazin anatomy of Deltatherium within a phylogenetic context, 1953; Lucas and Schoch 1998; Rose 2006). Pantodonts and we intend to use information collected from this study are mostly medium to large-sized herbivorous mammals in future work evaluating the phylogenetic relationships of known from North and South America and Asia and were Paleocene mammals. among the frst eutherian mammals to attain truly large body sizes following the end-Cretaceous mass extinction Historical Background (Simons 1960; Lucas 1998; Rose 2006). E. L. Simons (1960) also alluded to some shared, potentially plesio- The taxonomy and systematics of Deltatherium have a long morphic, features of the dentition and basicranium of history of study; however, it is useful to detail this informa- Deltatherium and members of Pantodonta. Regarding the tion to understand why the phylogenetic afnities of this auditory region, Simons noted that both Deltatherium taxon have remained so elusive. Deltatherium was first and Titanoides shared a broad, expanded region on the described in 1881 by E. D. Cope based on the upper denti- anterointernal surface of the petrosal beyond the region tion (Cope 1881a). The upper dentition indicated a more of the promontorium, which houses the cochlea (Simons omnivorous-carnivorous diet, and Cope considered Del- 1960). It is likely that Simons was referring to the rostral tatherium a member of Leptictidae within Creodonta rather tympanic process and medial shelf. than a member of ‘Condylarthra’, which was established L. Van Valen (1978) went a step further and based on taxa with more bunodont dentitions and tendencies hypothesised a pantodont affinity for Deltatherium. He towards a more omnivorous-herbivorous diet (Cope 1881a). speculated on a close relationship with the arctocyonid In the same year, Cope described the lower dentition of Del- Oxyprimus (a member of the Oxyclaeninae) and tatherium based on a partial dentary preserving the three illustrated Deltatherium giving rise to Pantodonta (Van lower molars; he named it Lipodectes penetrans and consid- Valen 1978). It is worth noting that Arctocyonidae had ered it to be a mesonychid creodont (Cope 1881b). With the been transferred from Creodonta to ‘Condylarthra’, a discovery of new material, Cope recognised the association position that is still broadly upheld, prior to Van Valen between the upper and lower dentition of Deltatherium and hypothesizing on the origins of pantodonts (Simpson reasserted a leptictid classifcation within Creodonta (Cope 1936; Gazin 1941; Van Valen 1966, 1969). Van Valen did 1884). not provide any explanation for his hypothesis but implied In 1887, M. Schlosser revised Creodonta, moving Del- some dental characteristics shared by Deltatherium and tatherium from Leptictidae to Proviverridae which also early Torrejonian pantodonts such as Pantolambda. included Triisodon and Didelphodus (Schlosser 1887). Cope Cifelli (1983) questioned an arctocyonid/pantodont acknowledged Schlosser’s referral of Deltatherium to Pro- affinity for Deltatherium, instead proposing a broadly viverridae and also referred Mioclaenus, Onychodectes, Tri- defined ‘insectivore’ affinity for the taxon, possibly isodon and Chriacus to the group (Cope 1888). W. B. Scott alluding to a cimolestan ancestry. McKenna (1975) had retained Cope’s concept of Proviverridae except for moving previously included Deltatherium within Pantodonta, Chriacus to Oxyclaenidae (Scott 1892). H. F. Osborn and which he classified within Cimolesta, but did not provide C. Earle followed suit and retained a Proviverridae classif- any rationale for his assertion. Szalay (1977) disputed cation (Osborn and Earle 1895). W. D. Matthew was indi- McKenna’s cimolestan classification of Pantodonta rectly skeptical that Deltatherium pertained to Proviverridae and provided discussion in support of Van Valen’s (Matthew 1897), and in 1899, he shifted Deltatherium from arctocyonid-Deltatherium-pantodont hypothesis based on Proviverridae to Oxyclaenidae within Creodonta and alluded dental and postcranial observations. Shortly thereafter, to a close afnity with Chriacus (Matthew 1899). In 1937, Chow et al. (1977) proposed a close relationship between Matthew reviewed the taxonomic history of Deltatherium in pantodonts and tillodonts within Cimolesta. Van Valen detail and described the specimen studied here, which was (1988) subsequently reasserted an arctocyonid affinity found during the 1913 expeditions to the San Juan Basin for Deltatherium and erected the subfamily Deltatheriinae (Matthew 1937). Matthew transferred Deltatherium to the within Arctocyonidae, which he considered closely subfamily Chriacinae within Arctocyonidae, which was still related to Pantodonta, presumably based on dental considered a member of Creodonta at the time. similarities, although this was not explicitly stated. Following a mid-century hiatus, C. L. Gazin noted a Muizon and Marshall (1992)
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