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doi: 10.7717/peerj.4648. eCollection 2018.

A new species ofGulo from the Early Pliocene Gray Fossil Site (Eastern United States); rethinking the evolution of wolverines

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A new species ofGulo from the Early Pliocene Gray Fossil Site (Eastern United States); rethinking the evolution of wolverines

Joshua X Samuels et al. PeerJ..

Abstract

The wolverine (Gulo gulo) is the largest living terrestrial member of the Mustelidae; a versatile predator formerly distributed throughout boreal regions of North America and Eurasia. Though commonly recovered from Pleistocene sites across their range, pre-Pleistocene records of the genus are exceedingly rare. Here, we describe a new species ofGulo from the Gray Fossil Site in Tennessee. Based on biostratigraphy, a revised estimate of the age of the Gray Fossil Site is Early Pliocene, near the Hemphillian-Blancan transition, between 4.9 and 4.5 Ma. This represents the earliest known occurrence of a wolverine, more than one million years earlier than any other record. The new species of wolverine described here shares similarities with previously described species ofGulo, and with early fishers (Pekania). As the earliest records of bothGulo andPekania are known from North America, this suggests the genus may have evolved in North America and dispersed to Eurasia later in the Pliocene. Both fauna and flora at the Gray Fossil Site are characteristic of warm/humid climates, which suggests wolverines may have become 'cold-adapted' relatively recently. Finally, detailed comparison indicatesPlesiogulo, which has often been suggested to be ancestral toGulo, is not likely closely related to gulonines, and instead may represent convergence on a similar niche.

Keywords: Carnivora; Guloninae; Mustelidae.

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Conflict of interest statement

The authors declare there are no competing interests.

Figures

Figure 1
Figure 1. Distribution ofGulo through time in (A) North America and (B) Eurasia.
Current range (light gray) and historic range (medium gray) are based on Pasitschniak-Arts & Larivière (1995), Zigouris et al. (2013); Pleistocene range (dark gray) is based on localities where fossils ofGulo have been found and reported in a wide range of literature sources. The only two known Pliocene occurrences, the Gray Fossil Site in Tennessee and Udunga Fauna of Russia, are highlighted with an asterisk. Terrain base map from ESRI ArcMap 10.5.
Figure 2
Figure 2. Stratigraphic ranges of selected mammals from the Gray Fossil Site in Tennessee.
The black bars indicate stratigraphic ranges of genera and species based on first and last appearance dates (data sources are listed in the Geologic Setting section). Overlap in ranges of taxa, between 4.9 and 4.5 Ma, is highlighted with a gray bar.
Figure 3
Figure 3. Holotype ofGulo sudorus (ETMNH 3663) from the Gray Fossil Site, Tennessee.
Specimen consists of a right P2 and maxilla fragment with P4. (A) Lateral view. (B) Occlusal view. (C) Original illustration of the specimen by Keila Bredehoeft. Scale bar equals 1 cm. Photographs by Joshua Samuels.
Figure 4
Figure 4. Occlusal morphology of gulonine mustelids andPlesiogulo.
(A)Gulo gulo (ETVP 291). (B)Pekania pennanti (ETVP 601). (C)Plesiogulo marshalli (composite of FAM 23386 and 49230). (D)Martes americana (NAUQSP 2015). Images are scaled to the same P2–M1 length. Original illustrations by Keila Bredehoeft.
Figure 5
Figure 5. Phylogeny of gulonine mustelids with estimated divergence times and geologic ages of known fossils.
Phylogenetic relationships of extant taxa based on cladistics analyses of Koepfli et al. (2008), Sato et al. (2012) and Li et al. (2014). Note that the placements of extinct taxa are not based on a cladistic analysis, rather they are based morphological comparisons made in this manuscript and prior studies. Divergence times (indicated by horizontal boxes) are based on protein coding region data set in Li et al. (2014), except divergence time of Guloninae from other “crown” mustelids (Helictidinae, Ictonychinae, Lutrinae, and Mustelinae) based on multidivtime analysis of Sato et al. (2012). Age ranges of fossil taxa (thick black lines) are based on occurrences and references listed in Table S2 and data sources listed in the Geologic Setting section.
Figure 6
Figure 6.Gulo gulo (ETVP 269) with an atavistic external median rootlet on the upper P4, which is exposed through the lateral surface of the maxilla.
Scale bar equals 1 cm. Photograph by Joshua Samuels.
See this image and copyright information in PMC

References

    1. Alcalá L, Montoya P, Morales J. New large mustelids from the Late Miocene of Teruel Basin (Spain) Comptes Rendus de l’Académie des Sciences de Paris. 1994;319(série II):1093–1100.
    1. Anderson E. Quaternary evolution of the genus Martes (Carnivora, Mustelidae) Acta Zoologica Fennica. 1970;130:1–132.
    1. Anderson E. Evolution, prehistoric distribution and systematics of Martes. In: Buskirk SW, Harestad AS, Raphael MG, Powell RA, editors. Martens, sables and fishers: biology and conservation. Cornell University Press; Ithaca: 1994. pp. 13–25.
    1. Anderson E. Quaternary wolverines (Gulo gulo) from Idaho. In: Akersten WA, McDonald HG, Meldrum DJ, Flint MET, editors. Papers on the vertebrate paleontology of Idaho honoring John H. White, volume 1. Idaho Museum of Natural History Occasional Paper 36; Pocatello, Idaho: 1998. pp. 173–185.
    1. Baryshnikov GF. Late Pleistocene Ursidae and Mustelidae remains (Mammalia, Carnivora) from Geographical Society Cave in the Russian Far East. Proceedings of the Zoological Institute RAS. 2015;319(1):3–22.

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