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Oligocene–Miocene) of Western North America A new tribe of castoroidine beavers from the late Arikareean to Hemphillian (Oligocene–Miocene) of western North America WILLIAM W. KORTH Korth, W.W. 2017. A new tribe of castoroidine beavers from the late Arikareean to Hemphillian (Oligocene–Miocene) of western North America. Acta Palaeontologica Polonica 62 (2): 249–258. Two genera of Miocene beavers (Castoridae), Hystricops and Priusaulax are referred to a new tribe Hystricopini of the subfamily of giant beavers, Castoroidinae based on cranial and dental morphologies. Previously undescribed and more complete specimens of Hystricops demonstrate that it is referable to the Castoroidinae rather than the Castorinae. Along with Priusaulax, it is believed to represent a group of basal castoroidines, distinct from later Castoroidinae by the retention of the primitive morphology of the contact of the lacrimal bone with the jugal on the cranium, as well as more primitive dental morphologies (lower crown-height and simple occlusal pattern of cheek teeth). A partial cranium and several isolated teeth from the Hemingfordian of Colorado, originally referred to Anchitheriomys?, are referred here to a new species, Priusaulax wilsoni sp. nov. Monosaulax senrudi from the Barstovian of Montana is also referred to Priusaulax. Hystricops is limited to the type species H. venustus from the Clarendonian and Barstovian North American Land Mammal Age (NALMA), and H. browni from the Hemphillian NALMA. Key words: Mammalia, Rodentia, Castoridae, Castoroidinae, cranial anatomy, dental terminology, North American Land Mammal Ages, Colorado. William W. Korth [[email protected]], Rochester Institute of Vertebrate Paleontology, 265 Carling Road, Roch- ester, New York 14610, USA. Received 10 January 2017, accepted 9 February 2017, available online 11 April 2017. Copyright © 2017 W.W. Korth. This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. ing a new species from the Hemphillian, H. browni Shotwell, Introduction 1963. The first larger sample (more than ten specimens) of Hystricops was reported from the late Clarendonian Ash Hystricops was named by Leidy (1858) as a subgenus of the Hollow Formation of Nebraska (Korth 1997, 1998b). This Recent Old World porcupine Hystrix Linneaus, 1758, based collection included the first specimens other than isolated on two isolated cheek teeth from the “Niobrara River beds” teeth. It was suggested that these specimens were likely from in northern Nebraska. This horizon was later interpreted the same horizon as the holotype material of H. venustus, as the Ogallala Group and divided into the Valentine and and referred to that species (Korth 1998b). Ash Hollow formations, representing the middle to early– Two other species have been suggested as referable to late Miocene: Barstovian to Clarendonian, repectively (see Hystricops by some authors: Monosaulax senrudi Wood, Voorhies 1990, for historical review). Matthew (1902) was 1945, from the Barstovian of Montana (see Stout and Stone the first to suggest that the teeth of Hystricops were more 1971) and Anchitheriomys? sp. (Wilson 1960) from the likely those of a beaver (= Castoridae) because of the implau- Hemingfordian of Colorado (also see Korth 1998b). sibility of having a genus of an Old World family occurring Due to the sparse record of Hystricops (predominantly in Tertiary rocks of North America. Stirton (1935) definitely isolated cheek teeth), its systematic position within the fam- referred the genus to the Castoridae based on the morphol- ily Castoridae has been in question (Stirton 1935; Korth ogy of the cheek teeth, even though no additional specimens 1994). McKenna and Bell (1998) placed it in the clade along had been recovered at the time of his publication. Additional with the Recent beavers in the subfamily Castorinae. Korth isolated teeth referred to Hystricops were later reported (2001b) included it in the Agnotocastorinae along with the from the Clarendonian and Hemphillian (late Miocene) of most primitive genera of the family. More recently, Flynn Oregon (Shotwell and Russell 1963; Shotwell 1963), includ- and Jacobs (2008) included the genus in “Basal Castoroids”, Acta Palaeontol. Pol. 62 (2): 249–258, 2017 https://doi.org/10.4202/app.00339.2017 250 ACTA PALAEONTOLOGICA POLONICA 62 (2), 2017 again suggesting inclusion among the more primitive gen- Tribe Hystricopini nov. era. However, in the cladogram presented by the latter Type genus: Hystricops Leidy, 1958. Hystricops (Flynn and Jacobs 2008: fig. 23.2), was figured Included genera: Hystricops Leidy, 1958; Priusaulax Korth and Bai- as the sister-taxon of the Recent Castor Linnaeus, 1758, sim- ley, 2006. ilar to the position suggested by McKenna and Bell (1998). Diagnosis Rybczynski (2007: fig. 3) placed Hystricops within the sub- .—Derived features shared with castorines, cas- family Castorinae. toroidines, and palaeocastorines: P3 lacking; P4/p4 larger Priusaulax was named by Korth and Bailey (2006) than molars; upper tooth rows slightly diverge posteriorly; from the latest Arikareean and early Hemingfordian (latest upper diastema arched; posterior palatine foramina inter- Oligocene–early Miocene) of Nebraska. It was regarded as sected by maxillary-palatine suture; palatal surface dis- basal castoroidine based on dental and cranial features as tinctly grooved; length of incisive foramen less than 30% well as age of occurrence. Rybczynski (2007: fig. 3), in a cla- length of upper diastema; loss of stapedial foramen in au- distic analysis of the Castoridae, had a similar result, plac- ditory bulla. Derived characters shared with castoroidines: ing Priusaulax as the sister-taxon to the clade that included larger size (compared to contemporaneous species); angle both the Castoroidinae and Castorinae. Similarities between of dentary laterally expanded with flattened ventral sur- Priusaulax and Hystricops appear to ally the two genera, face, mandibular condyle medial to angle and ascending and suggest a systematic position near the Castoroidinae. ramus (= zig-zag pattern). Retained primitive features of castorids: cheek teeth mesodont in crown-height; occlusal Institutional abbreviations.—AMNH, American Museum of morphology of cheek teeth with single, persistent re-entrant Natural History, New York, USA; F:AM, Frick Collections, (lingual hypoflexus on upper; buccal hypoflexid on lowers) American Museum of Natural History, New York, USA; and three persistent, transversely oriented fossettes (-ids); KU, University of Kansas Museum of Natural History, jugal bone contacts lacrimal at its anterior end; enamel sur- Lawrence, Kansas, USA; UNSM, University of Nebraska face of incisors smooth, and unornamented; nasal bones State Museum, Lincoln, Nebraska, USA; UOMNH, Uni- extend more posteriorly than premaxillaries on dorsal sur- versity of Oregon Museum of Natural History, Eugene, face of cranium; rostrum elongated; upper incisors slightly Oregon, USA; UW, University of Wyoming, Geological procumbent. Museum, Laramie, Wyoming, USA. Discussion.—The Hystricopini share nearly all of the de- rived cranial characters of the Castoroidinae (Korth 2001b: 289, fig. 4). The only major difference is in the contact of the Material and methods lacrimal bone with the jugal. In more derived castoroidines the jugal ends anteriorly well before joining the lacrimal. The majority of the fossil material discussed herein has This feature in hystricopines is clearly primitive among cas- been previously cited with the exception of three specimens torids. In the original description of Priusaulax, it was sug- of Hystricops venustus from Nebraska, housed in the Frick gested that the presence of most of the castoroidine features Collections of the American Museum of Natural History and presence of the lacrimal-jugal contact suggested a basal and the University of Nebraska State Museum, the bulk of position of the genus within the castoroidines. In addition, which were collected in the 1940s and 1950s. The holotype the cheek teeth of both Hystricops and Priusaulax are sim- Priusaulax and topotypic specimens of the new species of ilar to other early castoroidines (e.g., Monosaulax Stirton, from Colorado are housed at the University of Kansas 1935) being mesodont in crown-height and retaining a sin- Museum of Natural History, originally described by Wilson gle re-entrant and three transversely elongated enamel fos- P. browni (1960). Those of are at the University of Nebraska settes after minimal to moderate wear. State Museum (Korth and Bailey 2006). The holotype and Hystricopini differ from the nothodipoidine castoroi- referred specimens of H. browni are at the University of dines in being larger and lacking the fossorial adaptions Oregon Museum of Natural History, collected in the 1950s of the skull of the latter (smaller size; higher-crowned and early 1960s (Shotwell 1963). cheek teeth; low, wide cranium; more procumbent in- Dental terminology follows that of Stirton (1935), and cisors; greater postorbital constriction; diastema more cranial terminology follows that of Wahlert (1977). Division strongly arched; Korth 2007: 1536), as well as retaining a of North American Land Mammal Ages follows that of more primitive occlusal pattern of the cheek teeth (never Janis et al. (2008; e.g., Ar1, Cl2, etc.). All measurements in attaining the
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