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ALS DENVER Museum of Nature & SCIENCE ANNALS DENVER MUSEUM OF NATURE & SCIENCE ANNALS DENVER MUSEUM OF NATURE & SCIENCE ANNALS DENVER NUMBER 3, SEPTEMBER 19, 2013 MU Denver Museum of Nature & Science Annals WWW.DMNS.ORG/SCIENCE/MUSEUM-PUBLICATIONS SE U (Print) ISSN 1948-9293 M 2001 Colorado Boulevard OF Denver, CO 80205 Denver Museum of Nature & Science Annals N AT (Online) ISSN 1948-9307 U RE & S CIENCE The Denver Museum of Nature & Science inspires curiosity and excites minds of all ages through ANN A scientific discovery and the presentation and LS • preservation of the world’s unique treasures. Our N U vision is to create a community of critical thinkers M B ER who understand the lessons of the past and act as 3 • S responsible stewards of the future. Cover photo: Excavation crew from the Denver EPTEM Museum of Nature & Science at the Buck Spring B Quarry Site in the Wind River Formation, Wind River ER 19, 2013 19, Basin, Wyoming. Type stratigraphic level and site for Acrodentis rosenorum and Nyctitherium krishtalki. The Denver Museum of Nature & Science Annals is an Frank Krell, PhD, Editor-in-Chief open-access, peer-reviewed scientific journal publishing original papers in the fields of anthropology, geology, EDITORIAL BOARD: paleontology, botany, zoology, space and planetary James Hagadorn, PhD (subject editor, Paleontology and Revision of the Wind River Faunas, sciences, and health sciences. Papers are either authored Geology) by DMNS staff, associates, or volunteers, deal with DMNS Bridget Coughlin, PhD (subject editor, Health Sciences) Early Eocene of Central Wyoming. specimens or holdings, or have a regional focus on the John Demboski, PhD (subject editor, Vertebrate Zoology) Part 15. New Nyctitheriidae Rocky Mountains/Great Plains ecoregions. David Grinspoon, PhD (subject editor, Space Sciences) Frank Krell, PhD (subject editor, Invertebrate Zoology) (?Lipotyphla) with Analysis of the The journal is available online at www.dmns.org/Science/ Steve Nash, PhD (subject editor, Anthropology and Relationships of North American Taxa Museum-Publications free of charge. Paper copies are Archaeology) exchanged via the DMNS Library exchange program Michael G. Christiansen and ([email protected]) or are available for purchase EDITORIAL AND PRODUCTION: Richard K. Stucky from our print-on-demand publisher Lulu (www.lulu.com). Frank Krell, PhD: production DMNS owns the copyright of the works published in the Julie Whitman-Zai, PhD: copy editor Annals, which are published under the Creative Commons Attribution Non-Commercial license. For commercial use of published material contact the Alfred M. Bailey Library & WWW.DMNS.ORG/SCIENCE/MUSEUM-PUBLICATIONS Archives at [email protected]. DENVER MUSEUM OF NATURE & SCIENCE ANNALS NUMBER 3, SepteMBER 19, 2013 Michael G. Christiansen1 and Revision of the Wind River Faunas, Early Richard K. Stucky2 Eocene of Central Wyoming Part 15. New Nyctitheriidae (?Lipotyphla) with Analysis of the Relationships of North American Taxa ABSTRACT—Among the fossilized remains of early Eocene mammals collected from the Buck Spring Quarries of Wyoming are the dentitions of several previously undescribed nyctitheriids. Comparison of this material (from the Lost Cabin Member of the Wind River Formation, late Wasatchian Land Mammal Age (LMA), Lostcabinian Land Mammal Subage (LMSA, Wa-7)) to closely related taxa requires the description of a new genus and species of nyctitheres, Acrodentis rosenorum, as well as a new species of Nyctitherium, N. krishtalkai. A. rosenorum is similar to closely related Nyctitherium and Leptacodon, but is set apart by its distinctively shaped anterodorsally curving paraconid, together with a protoconid and metaconid that project away from one another forming an open trigonid. In the upper teeth, the paracone and metacone also project in slightly different directions, suggesting the association of upper and lower molar morphology. The protocone is nearly centered laterally between the paracone and metacone, unlike Leptacodon, and the hypoconal shelf is less broadly expanded than in Nyctitherium. N. krishtalkai, though similar to N. velox and N. serotinum, differs 1Department of Materials Science and from these species in that the cristid obliqua terminates where it strikes Engineering, Massachusetts Institute of the postvallid, the hypoconulid does not as closely twin the entoconid, Technology, Cambridge, MA 02139, and the entoconid occurs slightly higher than the hypoconid. In the U.S.A. [email protected] upper teeth, the conules and conular wings are more developed than in previously described species, the hypocones, though broadly expanded 2Department of Earth Sciences, into shelves on M1–2, are less developed, and the paracone and metacone Denver Museum of Nature & Science, of M3 are less reduced. The evolutionary context of these two new groups Denver, Colorado 80205, U.S.A. was investigated with a cladistic analysis based on dental characters, [email protected] including species from the described genera Nyctitherium, Leptacodon, Plagioctenodon, Plagioctenoides, Pontifactor, Wyonycteris, and Lima- conyssus. Palaeictops spp. served as the outgroup. The results of this analysis suggest a close relationship between Acrodentis, Leptacodon, and Nyctitherium, in which A. rosenorum appears closely related to an Christiansen & Stucky New Nyctheriidae from the Early Eocene of Wyoming ancestor intermediate between Leptacodon and Nycti- material assumed to belong to nyctitheres in addition to therium. N. krishtalkai is the most primitive of its genus dental material. Sigé (1976) considered the nyctitheres and is the most closely related to A. rosenorum. Whereas to be erinaceomorph lipotyphlans and Butler (1988) a cladistic analysis involving only nyctitheres should not regarded them as soricomorphs, as has Lopatin (2006) be used as basis to divide the family, two major clades in a review of the Asian taxa. More recently, in a review within the Nyctitheriidae of North America may exist: of all North American small insectivorous mammals, one including the genera Leptacodon, Plagioctenodon, Gunnell et al. (2008) also discussed the nyctytheres Nyctitherium, and Acrodentis and the other including within the Soricomorpha. Missiaen & Smith (2005) Wyonycteris, Limaconyssus, and “Plagioctenoides.” performed a cladistic analysis indicating that the Asian nyctitheres (Asionyctiinae) likely represent a monophy- letic group within the Nyctitheriidae. Lopatin (2006) The Wind River Basin’s Buck Spring quarries site included nyctitheres in his comprehensive review of (DMNH Locality 115, Natrona County, Wyoming, U.S.A.) the Asian insectivorous mammals of the Paleogene, in is the most significant locality of the Wind River Forma- which he proposed subdivision of the family into five tion for well preserved fossilized remains from the late subfamilies: the Nyctitheriinae, Amphidozotheriinae, early Eocene (Wa-7, late Wasatchian Land Mammal Asionyctiinae, Eosoricodontinae, and the Praolestinae. Age) (Stucky et al. 1990; Gunnell et al. 2009). Detailed In this arrangement, all North American taxa were study of this material has revealed several new mam- placed in the Nyctitheriinae. malian taxa (e.g., Dawson 2006). Among the remains Among North American taxa, researchers described produced by this locality are several exceptionally well a number of novel genera in the decades following Rob- preserved dentitions of a new genus and species of nyc- inson’s assessment, including Pontifactor (West 1974), titheres and a new species of Nyctitherium. The purpose Plagioctenodon (Bown 1979), Plagioctenoides (Bown of this paper is to describe the morphology of these new 1979), and Limaconyssus (Gingerich 1987). These new taxa and to present an analysis of the relationships of additions, as well as the description of new species of the North American nyctitheres. Nyctitherium (Krishtalka & Setogouchi 1977), Lepta- The nyctitheres have a particularly colorful history codon (Krishtalka 1976; Gingerich 1987; Secord 2008), of taxonomic allocation and subsequent reallocation, and others, produced a host of different ideas about the which has been reviewed by Robinson (1968) and relationships among the Nyctitheriidae. For instance, Krishtalka (1976). For instance, Nyctitherium was first Krishtalka suggested in 1976 that there was a natural described by Marsh in 1872, a genus which he thought division forming a group consisting of Leptacodon to have represented primitive bats. This same genus was packi, L. tener, L. catulus, and Nyctitherium and a later regarded as a talpid by Matthew (1909: 536), a group containing L. munusculum and Pontifactor. soricid by Simpson (1945: 50), an erinaceid by Van Valen Alternatively, Bown and Schankler soon after advocated (1967), and an ancestral talpid with erinaceid affinities a division into a natural group that consisted of Lepta- by Robinson (1968). After Robinson published his 1968 codon tener, Plagioctenodon, and Plagioctenoides, and paper, in which he revised the nyctitheriids, only N. velox another consisting of “L.” packi, “L.” munusculum, and N. serotinum remained as valid species in the genus Pontifactor, and Nyctitherium, although they cast some Nyctitherium. In a 1972 paper, Butler classified the nyc- doubt on whether this would constitute a truly monophy- titheres as primitive soricoids, an arrangement that has letic grouping due to a “complex of crossing characters” been maintained by subsequent researchers (McKenna (Bown & Schankler 1982: 58). The absorption into the & Bell 1997; Rose 2006). Hooker (2001) suggested the
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