First record of immigrant Phoberogale (Mammalia, Ursidae, ) from Southern California Author(s): Xiaoming Wang, Robert M. Hunt Jr, Richard H. Tedford and E. Bruce Lander Source: Geodiversitas, 31(4):753-773. Published By: Muséum national d'Histoire naturelle, Paris DOI: http://dx.doi.org/10.5252/g2009n4a753 URL: http://www.bioone.org/doi/full/10.5252/g2009n4a753

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Xiaoming WANG Natural History Museum of Los Angeles County, Department of Vertebrate Paleontology, 900 Exposition Boulevard, Los Angeles, California 90007 (USA) and Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044 (China) [email protected]

Robert M. HUNT Jr University of Nebraska, Division of Vertebrate Paleontology, W436 Nebraska Hall, Lincoln, Nebraska 68588-0514 (USA) [email protected]

Richard H. TEDFORD American Museum of Natural History, Department of Vertebrate Paleontology, Central Park West at 79th Street, New York, New York 10024 (USA) [email protected]

E. Bruce LANDER Paleo Environmental Associates Inc., 2248 Winrock Avenue, Altadena, California 91001-3205 (USA) and Natural History Museum of Los Angeles County, Department of Vertebrate Paleontology, 900 Exposition Boulevard, Los Angeles, California 90007 (USA) [email protected]

Wang X., Hunt R. M. Jr, Tedford R. H. & Lander E. B. 2009. — First record of immigrant Phobero- gale (Mammalia, Ursidae, Carnivora) from Southern California. Geodiversitas 31 (4) : 753-773.

ABSTRACT Phoberogale Ginsburg & Morales, 1995, is a primitive ursid named to separate hypercarnivorous species from a rather heterogeneous group formerly included in the genus Jourdan, 1862. Th e presence of Cephalogale (s.l.) in the Great Plains of the United States has been documented since the late 1980s, often in the context of immigrant taxa and biochronology. However, no formal description or illustration of the respective specimens was published. Recently, an

GEODIVERSITAS • 2009 • 31 (4) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 753 Wang X. et al.

associated skull and left dentary were recovered from an early terrestrial deposit in southern California. We take this opportunity to describe the Califor- nia specimen, which represents a new species, Phoberogale shareri n. sp. Th is new species belongs to the Phoberogale- lineage and is distinguished from species of Cephalogale (s.s.) by its larger size, less-reduced pre-carnassial premolars, and less-elongated M2. Within the genus Phoberogale, P. shareri n. sp. is more KEY WORDS derived than Europeans species (P. bonali, P. depereti, P. ginesticus) as indicated Mammalia, by its larger size and more diff erentially enlarged p4 compared to the p3. It dif- Carnivora, Ursidae, fers from Pakistani P. bugtiensis in having a less prominent notch between the systematics, anterior base of the P4 paracone and protocone, a more distinct anterior ridge on zoogeography, the P4 paracone, and more mediolaterally shortened M1-2. Phoberogale shareri Miocene, California, n. sp. represents the fi rst known Pacifi c Coast occurrence of this lineage and has new species. important zoogeographic and biochronologic implications.

RÉSUMÉ Première trace de l’immigrant Phoberogale (Mammalia, Ursidae, Carnivora) en Californie du sud. Phoberogale Ginsburg & Morales, 1995 est un genre d’ursidés primitifs créé pour distinguer les espèces hypercarnivores d’un groupe assez hétérogène intégré auparavant dans le genre Cephalogale Jourdan, 1862. La présence de Cepha- logale (s.l.) dans les Grandes Plaines des États-Unis a été documentée dès la fi n des années 80, souvent à propos de taxons immigrants et de la biochronologie du Néogène, mais aucune description formelle ou illustration des spécimens n’a été publiée. Depuis, un crâne associé à un dentaire gauche a été découvert dans des dépôts continentaux du Miocène inférieur de Californie méridionale. Nous profi tons de cette opportunité pour présenter les spécimens de Californie, qui représentent une nouvelle espèce, Phoberogale shareri n. sp. Cette nouvelle espèce appartient à la lignée Phoberogale-Phoberocyon et se distingue des espèces de Cephalogale (s.s.) par une plus grande taille, des prémolaires pré-carnassières moins réduites et une M2 moins allongée. Au sein du genre Phoberogale, P. shareri n. sp. est plus dérivé que les espèces européennes (P. bonali, P. depereti, P. gin- MOTS CLÉS esticus) comme le montrent sa plus grande taille et une diff érence de taille plus Mammalia, nette entre p4 et p3. Elle diff ère de P. bugtiensis du Pakistan par une encoche Carnivora, Ursidae, moins proéminente entre les bases antérieures du paracône et du protocône de systématique, P4, une crête antérieure du paracône de P4 plus distincte et des M1-2 moins zoogéographie, élargies médiolatéralement. Phoberogale shareri n. sp. représente la première Miocène, Californie, apparition connue de cette lignée sur la Côte pacifi que, ce qui a des implica- espèce nouvelle. tions zoogéographiques et biochronologiques importantes.

INTRODUCTION dae. It is characterized by its cursorial adaptations, such as digitigrade posture, and hypercarnivorous Th e genus Cephalogale Jourdan, 1862 s.l., (includ- dentition, in contrast to extant ursids that are all ing Phoberogale Ginsburg & Morales, 1995), is a plantigrade and possess the most hypocarnivorous medium- to large-sized arctoid carnivore generally teeth known in the Order Carnivora (Hunt 1998a; regarded as an early member of the Family Ursi- Ginsburg 1999). Such a tendency toward curso-

754 GEODIVERSITAS • 2009 • 31 (4) Phoberogale (Mammalia, Ursidae, Carnivora) from California

Santa Ynez Mtns Pine Mtn. WESTERN A B 91 Topa TRANSVERSE Topa Mtns RANGES Windy Ridge Santa Barbara PROVINCE (OCPC locality 2039) South Mtn. Ventura Simi Valley 241 -Oak Ridge 55 Santa Ana Santa Rosa Santa Monica Mtns Mountains Island C Los Angeles NORTHERN PENINSULAR 241 RANGES PROVINCE Loma Ridge San Joaquin SantaB Ana 261 Pacific Ocean Mtns Tustin Plain 0 50 km Hills 5 405 241 133 C San Joa Santa Monica Mtns quin Hills 27 N-1 Piuma 5 Road

1 1 74 1 Pacific Ocean 10 Pacific Ocean 0 5 km 0 5 km Point Dume

FIG. 1. — Location of ETC Windy Ridge (OCPC 2039) and Piuma Road localities, and exposures of Sespe Formation (shaded) in southern California. Modifi ed from Calvano et al. (2008). rial locomotion in Cephalogale might be linked Based on dental dimensions, yet providing no de- to its widespread geographic distribution. Indeed, tailed description, Hunt (1998a) noted that there beyond its original discovery in Europe, where up are possibly three species from the late Arikareean to 10 species have been described in the Oligo- and early Hemingfordian of Nebraska, Wyoming, cene through early Miocene (Bonis 1973), an and New Mexico in the United States, based on early Miocene species is also known from Pakistan specimens in the Frick Collection at the American (Forster-Cooper 1923). More recently, hints of its Museum of Natural History (F:AM) and in the existence in strata of Georgia, Mongolia, University of Nebraska State Museum (UNSM), and China also have begun to surface (Gabuniya and that some North American forms are larger 1964, 1966; Wang & Qiu 2003; Wang et al. 2005), than the European species. suggesting a pan-Eurasian distribution. As better So far, no record of Cephalogale is known west records become available from Asia, additional of the Continental Divide, despite its apparently discoveries will likely follow. widespread presence east of the Rocky Mountains. Despite its wide distribution, Cephalogale had It is thus signifi cant that an agency-mandated not been reported in North America until quite paleontologic mitigation program in the Santa recently. It is perhaps not surprising, even logical Ana Mountains of Southern California (Fig. 1) in hindsight, that Tedford et al. (1987) announced resulted in the recovery of a well-preserved skull the existence of Cephalogale in the New World and left dentary of Phoberogale Ginsburg & Mo- and used its fi rst appearance as an immigration rales, 1995, a genus recently erected to encompass event to defi ne the late Arikareean North American hypercarnivorous species formerly included in the Land Age (NALMA) (Ar3 of Tedford et al. rather heterogeneous Cephalogale. Th e associated 2004: fi g. 6.3). Its New World presence was further skull and ramus from Southern California represent documented by Hunt (1998a) in his summary of the only such specimen of its kind in North America North American ursids (see also Hunt 2004). and, therefore, are appropriate as the holotype of a

GEODIVERSITAS • 2009 • 31 (4) 755 Wang X. et al.

Topanga Formation silty claystone, mudstone, clayey siltstone (not measured) siltstone 0 m 2040 sandy mudstone, sandy siltstone

10 silty sandstone

fine-grained sandstone

20 fine- to medium-grainded sandstone 2019 medium-grainded sandstone Yatkolamys 30 medium- to coarse-grained sandstone

coarse-grained sandstone 40 pebble conglomerate

cobble conglomerate 50 fault 2039 OCPC fossil locality

60 (early Hemingfordian) fossil land mammal locality Upper ETC Windy Ridge l.f.

70

80 arenarum or M. elegans 2062 2039 Phoberogale shareri n. sp. 90 2038

100

110

2017 120

130 184 m (604 ft) to base of measured section Lower ETC Windy Ridge l.f. (late Arikareean)

FIG. 2. — Stratigraphic column of ETC Windy Ridge section. After Lander (2003: pl. 10). Colors are simplifi ed approximations of the original colors observed in outcrops of the formation.

756 GEODIVERSITAS • 2009 • 31 (4) Phoberogale (Mammalia, Ursidae, Carnivora) from California

new species, P. shareri n. sp. We dedicate this paper SYSTEMATICS to a festschrift volume in honor of Professeur Louis de Bonis, a long-time colleague and a preeminent Order CARNIVORA Bowdich, 1821 expert on fossil carnivores, including Cephalogale. Infraorder ARCTOIDEA Flower, 1869 Parvorder URSIDA Tedford, 1976 Superfamily URSOIDEA MATERIAL AND METHODS Fischer de Waldheim, 1817 Family Frick, 1926 In this paper, we provide a detailed description Subfamily PHOBEROCYONINAE of the unique Southern California specimen of Ginsburg & Morales, 1995 Phoberogale. Although we make occasional reference Genus Phoberogale Ginsburg & Morales, 1995 to other North American specimens from east of the Continental Divide, those specimens, much more Phoberogale shareri n. sp. numerous in the F:AM and the UNSM collections, (Figs 3-5; Tables 1; 2) will be described elsewhere. For comparative pur- poses, we examined selected examples of Cephalogale Cephalogale sp. – Whistler & Lander 2003: 256. and Phoberogale from the classic collections of the HOLOTYPE. — OCPC 21794 and 21795, rostral part Quercy district housed in the Naturhistorisches of skull missing tip of inion and much of basicranial Museum Basel, the British Museum of Natural His- region, but with complete upper dentition except right tory, and the Muséum national d’Histoire naturelle P1, and nearly complete left dentary with c-m2 and in Paris, as well as the more recent, stratigraphically m3 alveolus; fi eld number BMM111396.1; collected by Brett M. Malas, Raytheon Infrastructure Services, controlled collections in the Université de Poitiers. Inc., December 29, 1995. Although the skull (21794) For Asian specimens, we examined the holotype of and dentary (21795) were catalogued with separate Phoberogale bugtiensis Cooper, 1923 and had access numbers, they belong to the same individual, based on to an unpublished manuscript on new specimens their recovery at the same locality and the similar size of Cephalogale from the Hsanda Gol Formation relationships and stages of wear of their teeth. of Mongolia. ETYMOLOGY. — In honor of Mr Kevin W. Sharer for his leadership as Chairman of the Board of Trustees at the ABBREVIATIONS Natural History Museum of Los Angeles County. BMNH B ritish Museum of Natural History, London; ETC Eastern Transportation Corridor, California; TYPE LOCALITY. — OCPC 21794-5 were collected as a F:AM Frick Collection, American Museum of result of mitigation activities conducted in support of Natural History, New York; grading for the Eastern Transportation Corridor (ETC) IVPP Institute of Vertebrate Paleontology and highway construction project (Lander 2003). Th e type Paleoanthropology, Beijing; locality, OCPC locality 2039, is in the northern (ETC l.f. local fauna; Windy Ridge) segment of SR 241, about 8.2 km south MAE Collections of the joint Mongolian Academy of its intersection with State Route 91 (Riverside Free- of Sciences – American Museum of Natural way) and north of its intersection with SR 261 in sec. History Paleontological Expeditions, currently 23 T.4S. R.8W. (33°49’34’’N/117°42’52’’W) in the housed in American Museum of Natural northern Santa Ana Mountains of Orange County, History, New York; coastal southern California (Fig. 1). MN European Neogene mammal zone/unit; MP European Paleogene mammal zone/unit; DIAGNOSIS. — Phoberogale shareri n. sp. is easily distin- NALMA North American Land mammal Age; guished from “Cephalogale” minor Filhol, 1879 [Filhol OCPC Orange County Paleontological Collection, 1879] by its much larger size, enlarged P4 and m1, County of Orange, Resources and Develop- posteriorly located P4 protocone, and diff erentially ment Management Department, Santa Ana, enlarged p4 relative to p3. It diff ers from species of California; Cephalogale (C. geoff royi Jourdan, 1862, C. ursinus Bonis, SR State Route; 1973, C. brevirostris (Croizet, 1836) [Croizet 1836]) UNSM University of Nebraska State Museum, Lin- in its larger size and primitively less reduced pre-car- coln, Nebraska. nassial premolars and less elongated M2. Within the

GEODIVERSITAS • 2009 • 31 (4) 757 Wang X. et al.

A

B

C

FIG. 3. — Phoberogale shareri n. sp., OCPC 21794, holotype (associated with OCPC 21795) from ETC Windy Ridge locality, Orange County, California; lateral (A), dorsal (B), and ventral (C) views of partial skull. Scale bar: 30 mm.

758 GEODIVERSITAS • 2009 • 31 (4) Phoberogale (Mammalia, Ursidae, Carnivora) from California

TABLE 1. — Dental measurements (in mm) of Phoberogale species. Those for P. bonali Helbing, 1928 are from Helbing (1928); those for P. depereti Viret, 1929 from Beaumont (1965); those for P. ginesticus Kuss, 1962 from Kuss (1962).

P. shareri n. sp. P. bugtiensis P. bonali P. depereti P. ginesticus C1 length 10.2 C1 width 8.8 P1 length 7.1 P1 width 3.7 P2 length 8.8 P2 width 4.4 P3 length 10.3 P3 width 5.6 4.9 P4 length 20.3 21.1 W4 width 12.9 12.7 M1 length 15.8 15.5 15.0 M1 width 17.4 18.2 16.5 M2 length 10.1 9.4 M2 width 13.1 14.5 c1 length 10.9 c1 width 7.6 p1 length 6.5 p1 width 3.3 p2 length 8.3 9.0 p2 width 5.1 5.0 p3 length 10.0 10.0 p3 width 5.5 5.5 p4 length 13.4 12.7 13.0 11.5 p4 width 7.5 7.0 6.5 m1 length 24.6 19.2 19.8 22.0 m1 width 10.6 8.0 9.2 9.2 m1 trigonid length 17.4 m2 length 13.5 10.8 12.1 12.5 m2 width 9.7 7.2 7.9 8.0 genus Phoberogale, P. shareri n. sp. is distinguished from and left dentary (OCPC 21795) are catalogued as European species (P. bonali Helbing, 1928, P. depereti separate specimens. Similarities in wear stage on the Viret, 1929, P. ginesticus Kuss, 1962) by its larger size, and more diff erentially enlarged p4 over p3. It diff ers upper and lower teeth and the preservation of bones from Pakistani P. bugtiensis in its less prominent notch suggest that they belong to the same individual. between anterior base of P4 paracone and protocone, OCPC 21794-5 belongs to a young adult with more distinct anterior ridge on P4 paracone, and more permanent dentition fully erupted but showing mediolaterally shortened M1-2. Species of Phoberogale minimal wear. Th e skull (Fig. 3) is missing the are distinguished from more-derived Phoberocyon by their smaller size, absence of a premasseteric fossa, and posterior half of the braincase and the basicranial lack of a distinct m2 paraconid. region, but the remaining parts are more or less intact. Overall, the skull is slightly dorsoventrally DESCRIPTION crushed resulting in a shallower depth, with the Th e type and only specimen of Phoberogale shareri most deformation sustained at the orbital region n. sp. is preserved in deep purplish mudstones, and and the glenoid fossa region. suff ers from minor to modest distortions and cracks. Th e nasals sag slightly along the mid line such Its teeth are nearly perfectly preserved. Although that the left and right nasals meet each other at the basicranial region in OCPC 21794-5 is miss- almost a right angle. Th e anterior tips of the na- ing, it is the only associated skull and dentary in sals are missing and the posterior margins extend North American Phoberogale materials, and is thus back behind the anterior rim of the orbit, situated well suited as a holotype. Th e skull (OCPC 21794) at the same level as the frontal-maxillary suture.

GEODIVERSITAS • 2009 • 31 (4) 759 Wang X. et al.

A

B

FIG. 4. — Phoberogale shareri n. sp., OCPC 21795, holotype (associated with OCPC 21794) from ETC Windy Ridge locality, Orange County, California; lingual (A) and labial (B) views of left dentary. Scale bar: 30 mm.

Th e frontal is very broad (widest distance between In lateral view, the infraorbital foramen is at dorsal rims of the orbits is 59 mm and distance the level of the anterior border of the P4 and the between postorbital processes of the frontals is infraorbital canal is relatively long. In anterior view, 55 mm). Th e frontal is slightly elevated behind the foramen is a narrow oval with the long axis the postorbital process, indicating a modest de- oriented vertically. Th e zygomatic arch is mostly velopment of frontal sinuses. Th e temporal crests missing on both sides, with only the anterior seg- behind the postorbital process are not sharply ment preserved on the right side. Th e remaining delineated and they converge to a single sagittal zygomatic arch is quite deep, 25 mm at its ante- crest about 10 mm anterior to the frontal-parietal rior base and 14 mm at the broken tip. Bones in suture. About 40 mm of the anterior sagittal the orbital region suff er from extensive damage crest is preserved and it is quite low (~4 mm in and the lacrimal area is poorly preserved on both height). sides. Even on the relatively better preserved left

760 GEODIVERSITAS • 2009 • 31 (4) Phoberogale (Mammalia, Ursidae, Carnivora) from California

side, individual bones in the orbital mosaic are form a basal cingulum. Th e upper canine is coni- diffi cult to make out. cal, oval in cross section, curving backward slightly, In ventral view, the palate is not broadened as and has a total length of ~24 mm. An indistinct in other North American Phoberogale (see Com- anterolingual and posterior crest is present along parison below), with 66.4 mm the maximum dis- the length of the tooth. tance between the left and right P4s. Th e incisive P1-3 are simple, single cusped teeth, and gradu- foramina are markedly widened and they form a ally increase in size posteriorly (Figs 3; 5). All circular opening 12.4 mm in greatest diameter. have an anterolingual and a posterior crest. A very Part of the palatines, particularly that on the right indistinct cingulum surrounds the entire P3, and side near the P4-M1, is missing, and the exact that on the P2 is even more indistinct. P4 is large position of the posterior margin is not clear. Judg- and robust but not to the extent seen in the large ing from the remainder, however, the posterior phoberocyonine species from the Nebraska Heming- palatine border seems to stop at a level anterior to fordian. Th e paracone has an anterior ridge but lacks the posterior edge of the M2s. Th e basicranium is a parastyle. Th e protocone is not advanced but is largely missing, except the left basisphenoid area. retracted along the lingual face of the P4, hence An alisphenoid canal is present. Th e postglenoid is posterior to the anterior edge of the paracone, fossae have been forced forward by at least 6 mm, and a slight notch is present between these cusps. resulting in a shortened orbital region, especially Th e protocone is low and rather fl at anteriorly but on the right side. includes a small sharp, ridge-like cusp toward the Th e left dentary is largely intact except the tip of posterior end. A very weak cingulum is present on the ascending ramus and angular process (Fig. 4). Th e the anterior and labial margin, and that along the horizontal ramus gently tapers toward the anterior lingual margin is more distinct. A distinct carnas- end, with the depths and widths below p2, p4, and sial notch is present. m2 being 22.2 × 11.8 mm, 22.7 × 11.2 mm, 25.9 × M1 is subquadrate in occlusal outline. Th e labial 9.7 mm, respectively. Although the bony surface on cingulum and parastyle are poorly developed. Th e the labial side below the m2 is somewhat damaged in paracone is taller crowned than the metacone, al- an area about 15 mm in width, the remaining bone though wear on these cusps does not allow a more surrounding this damaged area shows no sign of a precise description. Th e protocone is a low, slightly premasseteric fossa. Th e bone between the ventral enlarged cusp situated at the lingual termination rim of the masseteric fossa and ventral edge of the of the preprotocrista. Th e metaconule forms a low horizontal ramus is narrow, measuring 8.2 mm in ridge posterior to the protocone, and separated width. Despite the loss of the tip of the ascending from it by a notch. It is anteriorly expanded at the ramus, the overall height of the ascending ramus expense of the postprotocrista. Th e posteriorly di- is estimated to be ~52 mm. Th e tip of the angular rected metaconule ridge adopts a slight labial turn, process is missing. ending at the posterior margin of the tooth rather With the exception of the lower incisors, the entire than at the base of the metacone. A posterolingual upper and lower dentition is well preserved; only cingulum (hypocone) is moderately developed and modest wear ensures that the cusp morphology can be wraps anteriorly around the protocone. M2 has described. Th e dental formula is 3142/3143. Upper an oval outline. Its paracone is much larger than incisors form a relatively straight dental arcade and the metacone, and as in M1, the labial cingulum they are procumbent. Wear on the tips of the I1-2 is weak. Th e protocone-metaconule ridge is low obscures their cusp morphology, but they appear and indistinct. Th e lingual cingulum (hypocone) to be unicuspid. Th e size increase from I1 to I3 is is expanded at its posterolingual corner. relatively modest and the I3 is not exceptionally Th e lower incisors are missing. Th e lower ca- enlarged as seen in some large, hypercarnivorous nine is more posteriorly curved than its upper caniforms. Th e I3 has a medial and lateral ridge, counterpart. Th e p1-3 are single cusped with an and the medial ridge wraps around lingually to indistinct anterior and posterior ridge (Figs 4; 5).

GEODIVERSITAS • 2009 • 31 (4) 761 Wang X. et al.

AB

FIG. 5. — Phoberogale shareri n. sp., OCPC 21794–5, holotype from ETC Windy Ridge locality, Orange County, California; occlusal stereo views of lower (A) and upper teeth (B). Scale bar: 30 mm.

762 GEODIVERSITAS • 2009 • 31 (4) Phoberogale (Mammalia, Ursidae, Carnivora) from California

Size increase from p1-p3 is gradual and slight. Th e (2955 ft) (Schoelhamer et al. 1981). Locally in the p4 is substantially larger and more robust than p3, ETC Windy Ridge section of SR 41, the exposed, and in addition to anterior and posterior ridges, a upper portion of the sequence is 317 m (1039 ft) small posterior accessory cusp is present. Th e p4 thick and is cut by at least three minor unmapped also has a more pronounced cingulum, particularly faults. Th e magnitude and sense of off set along along the lingual border. Th e lower carnassial (m1) these faults are undocumented. Consequently, the is long, with a well-defi ned shearing blade formed amount of section that has been removed or repeated by the labial facet of paraconid and protoconid. by faulting is undetermined. Th e base of the section Th e protoconid is much taller than the paraconid. was not exposed by grading. Whistler & Lander Th e metaconid is low and slightly off set from the (2003) considered the Sespe/Vaqueros Formation protoconid (positioned at the posterolingual aspect in the ETC Windy Ridge section to range in age of the protoconid). A dominant hypoconid occupies from late Uintan to early Hemingfordian. Th e en- more than two thirds the width of the talonid. A tire section is composed of the “Sespe” lithofacies, lower entoconid ridge, with less than one third the which is disconformably overlain by the marine, width of the talonid, encloses a shallow basin. Th e late Hemingfordian Topanga Formation (Calvano entoconid crest is not divided into smaller cusps. et al. 2003). Th e m2 protoconid is larger than the metaconid Th e Phoberogale shareri n. sp. type locality (OCPC in occlusal view. Wear on these cusps prevents an locality 2039) was in a 2.7 m (9 ft) thick, fi ne- estimate of their relative height, but it is likely grained red sandstone 307 m (1006 ft) above the that the protoconid is taller than the metaconid, base of the ETC Windy Ridge section, and a greater, based on comparisons with other North American but undetermined, distance above the base of the Phoberogale specimens. A paraconid is absent, and in formation (Fig. 2). Th is locality is approximately its place is a triangular platform enclosed between 85 m (280 ft) below the disconformably overlying, protoconid and metaconid. As in the m1, the m2 marine Topanga Formation of late Hemingfordian hypoconid is much larger and higher than the ento- age, but the intervening section is cut by the up- conid, which is a low crest. An m3 is indicated by per of the three documented faults. Th e holotype a single-rooted alveolus. (OCPC 21794-5) is preserved in a deep purplish claystone and is the only specimen collected from GEOLOGY, ASSOCIATED FAUNA, AND AGE the site. Stratigraphically correlative sections (ETC Surfi cial geologic mapping of the northern Santa Jamboree Road section, Bolero Lookout, Laguna Ana Mountains is provided by Morton & Miller Hills) yielded magnetostratigraphic records variously (1981) and Schoellhamer et al. (1981). Grading correlated with C5Cr-C5Er (16.73-19.05 Ma) for the northern segment of SR 241 north of Irvine (Prothero & Donohoo 2001; Calvano et al. 2008). Lake cut through one of the thickest sections of However, Whistler & Lander (2003) revised the the undiff erentiated continental Sespe and marine land mammal age assignment for two assemblages Vaqueros formations (Fig. 2). In the northern Santa in the ETC Jamboree Road section and, conse- Ana Mountains, these formations interfi nger exten- quently, changed the chron assignments to C5Cn-6r sively (Schoelhamer et al. 1981) and are not easily (17.28-20.52 Ma). distinguished or mapped separately. Th e upper or Th e upper part of the ETC Windy Ridge section, dominantly marine “Vaqueros” lithofacies consists which contains late Arikareean and early Hem- of interbedded reddish, buff , and gray to white sand- ingfordian land mammal assemblages, is above an stones and greenish-gray to white sandstones and intraformational unconformity that lies at the base siltstones, whereas interbedded reddish, buff , and of a conglomeratic interval and separates this part of gray to white sandstones and claystones constitute the section from the lower part (Calvano et al. 2003). the lower or continental “Sespe” lithofacies (Lander Th e lower part of the section yielded a late Uintan 2003; Whistler & Lander 2003). Th e formations have land mammal assemblage at OCPC locality 2020 a combined maximum thickness of roughly 900 m (Calvano et al. 2003; Whistler & Lander 2003).

GEODIVERSITAS • 2009 • 31 (4) 763 Wang X. et al.

Whistler & Lander (2003) divided the land mam- Yatkolamys and M. elegans are reported in the early mal assemblages from the upper part of the ETC Hemingfordian of north- Windy Ridge section into the upper and lower western Nebraska (Martin & Corner 1980; Lander ETC Windy Ridge local faunas (l.f.s). Th e early 1998: locality CP106). Partly on the basis of the Hemingfordian, upper ETC Windy Ridge l.f. is latter two taxa, Whistler & Lander (2003: fi g. 11.5) represented by taxa from OCPC localities 2019 correlated the upper ETC Windy Ridge l.f. and and 2040, whereas the late Arikareean, lower ETC correlative assemblages (including Santiago Canyon Windy Ridge l.f. spans the “290 m of section below Landfi ll l.f.) in the “Sespe” lithofacies with taxo- locality OCPC 2019” and includes OCPC localities nomically similar assemblages in the Runningwater 2016-2018 (Whistler & Lander 2003). Taxa from Formation. In contrast, latest Arikareean Merychyus the upper ETC Windy Ridge l.f. include Cynelos arenarum, but neither M. elegans nor Yatkolamys, sp., whistleri Wang &Tedford, 2008, occurs in the Anderson Ranch Formation, which Phlaocyon taylori Hayes, 2000, Cuyamalagus dawsoni immediately underlies the Runningwater Forma- Hutchison in Hutchison & Lindsay, 1974, Noto- tion. Found at least 52 m (171 ft) below Yatkolamys, tamias spp., Miospermophilus spp., Yatkolamys sp., the crushed Merychyus skull (OCPC 22097) from and Merychyus elegans Leidy, 1859, while the lower OCPC locality 2062 (Fig. 2) cannot be assigned ETC Windy Ridge l.f. contains Heterosoricidae, confi dently to either species. Consequently, the Cuyamalagus dawsoni Hutchison in Hutchison & Phoberogale shareri n. sp. holotype from no more Lindsay 1974, Gripholagomys? sp. Nototamias spp., than 7 m (23 ft) lower in the section (Fig. 2) does Miospermophilus sp., Schizodontomys sp., Leidymys not occur in association with an age-diagnostic spe- nematodon (Cope, 1879), Merychyus? sp., ?Tanymykter cies of this oreodont. Another age-diagnostic taxon sp., ?Machaeromeryx tragulus Matthew, 1926, and in the upper ETC Windy Ridge l.f. is Cynelos sp., ?Pseudoblastomeryx advena Matthew, 1907 (Whis- which is based on a partial skull and postcranial tler & Lander 2003: table 2; Wang & Tedford 2008). skeleton from OCPC locality 2038, no more than In particular, the murid rodent Yatkolamys sp. from 11 m (35 ft) below the Phoberogale shareri n. sp. OCPC locality 2019 and the oreodontid artiodactyl type locality, OCPC locality 2039 (Fig. 2). Th is Merychyus arenarum or M. elegans from OCPC local- medium-sized amphicyonid carnivore, presum- ity 2062 (Fig. 2) were considered typical of the early ably an immigrant from Eurasia, is fi rst recorded Hemingfordian NALMA and played a key role in in North America by a single isolated lower carnas- separating the upper ETC Windy Ridge l.f. from the sial from the late Arikareean Agate Spring l.f. of lower. Whistler & Lander (2003: 256, fi g. 11.4) as- northwestern Nebraska, and did not become well signed the Phoberogale shareri n. sp. holotype (OCPC represented in the North American fossil record 21794-5) from OCPC locality 2039 to Cephalogale until the early Hemingfordian NALMA of the Great sp., which at that time they regarded as a member of Plains (Hunt 1972; Tedford & Frailey 1976; Hunt the upper ETC Windy Ridge l.f. (Fig. 2). 1998b). Th is skull, based on its proportions and Th e Phoberogale shareri n. sp. type locality (OCPC dentition, suggests a latest Arikareean age. Unfortu- locality 2039) is approximately 56-66 m (185- nately, other age-diagnostic taxa, such as the latest 215 ft) below OCPC locality 2019, which yielded Arikareean oreodont matthewi and dominantly early Hemingfordian Yatkolamys sp. and early Hemingfordian M. magnus and M. proprius, the lowest assemblage assigned to the upper ETC are not recorded from the undiff erentiated Sespe Windy Ridge l.f. by Whistler & Lander (2003), but and Vaqueros formations. about 27-39 m (87-127 ft) above OCPC locality Radiometric data from the Sespe Formation 2017 and the highest assemblage referred to the support a latest Arikareean age for Phoberogale lower ETC Windy Ridge l.f. (Fig. 2). On the other shareri. n. sp. Along Piuma Road in the central hand, an undetermined species of Merychyus occurs Santa Monica Mountains of Los Angeles County at OCPC locality 2062, no more than 7.0 m (23 ft) (Fig. 1), Leidymys nematodon occurs in the lower above OCPC locality 2039 (Lander 2003). part of the Piuma Member of the Sespe Formation,

764 GEODIVERSITAS • 2009 • 31 (4) Phoberogale (Mammalia, Ursidae, Carnivora) from California

TABLE 2. — Craniodental dimensions (in mm) of Phoberogale shareri n. sp. from California, and Phoberogale sp. (F:AM 54464) from the Cephalogale Jourdan, 1862 Ash locality, Dawes Co., Nebraska.

P. shareri F:M 54464 Length of rostrum 91.6 95.2 Palatal breadth across P4s 66.4 76.5 Breadth across zygomatic arches <110 158 Width across upper canines 43.2 57.0 Minimum breadth between dorsal orbital rims 55.5 62.7 Width across postorbital constriction 36.6 42.8 Maximum diameter of upper canine 10.2 14.6 Distance between posterior border of M2 and foramen rotundum 35.5 59.1

whereas Yatkolamys sp. occurs near the top of the III” from fi ve diff erent localities in the overlying upper part of the member (Fig. 1; Fisk et al. 2001; Runningwater Formation or equivalent beds (early Lander et al. 2001a; Lander et al. 2001b; Lander Hemingfordian), Morrill, Sheridan, Dawes, and Box 2003; Calvano et al. 2008: fi g. 10). Th e two taxa are Butte counties, Nebraska. Dental, postcranial and widely separated stratigraphically, as in the Santa Ana rare cranial constitute the North American Mountains. Th e Upper Saddle Peak Tuff lies near material, with the largest sample coming from the the base of the upper part of the Piuma Member, early Hemingfordian Bridgeport Quarries. roughly 255 m (835 ft) below the Yatkolamys record, A comprehensive description of these materials and has been determined to be 21.1 ± 0.2 Ma old is beyond the scope of this study. Instead, we pro- (Lander et al. 2003; Calvano et al. 2008: fi g. 10). vide a brief comparison, leaving a full treatment of Th e level that produced the Phoberogale skull prob- the remaining North American material for future ably correlates with one between the tuff and the studies. A decade ago, Hunt (1998a) noted that the level that produced Yatkolamys sp. largest North American form of Cephalogale (s.l.), his early Hemingfordian species III, exceeded in COMPARISON WITH NORTH AMERICAN FORMS lower carnassial size (m1 length ~23-31 mm) all of Mainly through intensive collecting by the F:AM the known Eurasiatic forms (~12-23 mm). Hunt’s and UNSM, such relatively rare carnivorans as Cephalogale new species I-II were smaller, poorly Phoberogale were accumulated over time. Although represented late Arikareean ursids, known by frag- the specimens were collected many decades ago, their mentary teeth only, whereas the larger, early Hem- existence was announced quite recently and only ingfordian species III was represented by numerous in the context of North American biochronology, specimens from the Bridgeport Quarries. i.e. Cephalogale (now Phoberogale), along with other Phoberogale shareri n. sp. is closest in size to immigrants, was used to defi ne the late Arikareean Hunt’s (1998b) species II from the late Arikareean NALMA (Tedford et al. 1987; 2004). In his summary of Wyoming (a partial maxilla with M2), as well of North American ursids, Hunt (1998a) briefl y as small individuals from the Bridgeport Quarries. discussed the nature of North American Cepha- Compared to the Bridgeport Quarries form, OCPC logale. He listed three unnamed species from eight 21794-95 lacks a parastyle on P4 and consequently localities: “Cephalogale new species I” from Standing lacks a distinct notch between the anterior border Rock Quarry, Zia Sand Formation (late Arikareean), of the paracone and the protocone. Th e California Sandoval County, New Mexico; “Cephalogale new species has a metaconule ridge on M1 closer to the species II” from University Quarry, Upper Harrison metacone, i.e. directed more labially, in contrast to Beds (late Arikareean), Sioux County, Nebraska and the more anteroposterior orientation of the ridge in 16 Mile District, Upper Harrison Beds, Goshen the Bridgeport Quarries species. Th is feature also County, Wyoming; and “Cephalogale new species results in a more developed M1 hypocone (from

GEODIVERSITAS • 2009 • 31 (4) 765 Wang X. et al.

the posterolingual cingulum) at the expense of the greater than 4 mm throughout its preserved length. metaconule in the California species as opposed to Hence, a less pronounced inion seems probable in equal development of metaconule and hypocone in OCPC 21794 and suggests a subtle diff erence in the Bridgeport Quarries carnivore. In addition, the overall proportions of the temporal muscles and Bridgeport hypodigm displays a distinct paraconule posterior cranium in the two carnivores. lacking in P. shareri n. sp. Of the F:AM material, the best preserved specimen is a nearly complete skull and partial skeleton (F:AM DISCUSSION 54464) from the “Cephalogale Ash” locality 1 mile west of the post offi ce at Marsland, Dawes County, Despite over a century and a half of study, records Nebraska. Th is skull shows a rostrum 4% longer for Cephalogale/Phoberogale and related forms are than P. shareri n. sp. and a palate 15% broader. Th is still quite poor, and the majority of published contrast in width is even more pronounced across specimens are fragmentary maxillaries and dentaries, the zygomatic arches where F:AM 54464 exceeds with only one previously fi gured skull of “C.” mi- P. shareri n. sp. by 44%. Similarly, the forehead is nor (Beaumont 1965: fi g. 14) and a partial skull of broader in F:AM 54464 and the upper canines C. gracile (Viret 1929: pl. IX, fi g. 4a). Bonis’ (1973) are more widely spaced (Table 2). Although the systematic synthesis is still the most comprehensive glenoid fossae of OCPC 21794 have been anteri- compared to other recent treatments (Beaumont orly displaced, the left alisphenoid canal and area 1965; Ginsburg & Morales 1995), even though he of the orbital fi ssure are relatively intact. Here the did not attempt to treat every species. We did not distance between the posterior margin of M2 and personally examine most of the European specimens. the foramen rotundum in F:AM 54464 is 66% Consequently, our systematic determination relies greater than in P. shareri n. sp. and this distinction on published descriptions, fi gures, and measure- cannot be explained away by mere preservational ments, while acknowledging the limitations in not eff ects. It is uncertain whether these proportional studying these specimens fi rsthand. diff erences are due to sexual dimorphism, as F:AM 54464 is likely a male with a large upper canine TAXONOMY OF CEPHALOGALE (maximum diameter of 14.6 mm vs. 10.2 mm for (PHOBEROGALE) SPECIES OCPC 21794), a large I3 relative to I1-2, and a Since the fi rst recognition of Cephalogale (Jourdan high sagittal crest. 1862), more than a dozen species have been referred Other than the above proportional diff erences to the genus at one time or another. Beaumont between the California and Nebraska skulls, ad- (1965) was fi rst to attempt a systematic revision ditional characters indicate that they belong to of the genus, and he recognized eight European diff erent species. Th e lateral profi le of the sagittal and one Asian species: C. gracile (Pomel, 1846) crest in F:AM 54464 is essentially horizontal with a [Pomel 1846] from Saint-Gérand-le-Puy (Allier, gentle downward curvature near the frontal-parietal MN2a), C. geoff royi from Billy (Allier, MP30), suture – the crest is prolonged posteriorly where it C. cadurcensis (Filhol, 1877) from an unspeci- forms an overhanging inion. Th is profi le displays a fi ed locality in the Quercy district, C. minor from gradual increase in crest depth to the point where Saint-Gérand-le-Puy, C. brevirostris from Gergovie the surface of the braincase bends downward. Here (Puy-de-Dôme), C. bonali Helbing, 1928 (see also the maximum depth of the crest near the inion is Helbing 1922) from La Milloque (Lot-et-Garonne, ~29 mm. Th e crest profi le for OCPC 21794, on MP29), C. depereti Viret, 1929 from Montaigu-le- the other hand, slopes downward in the posterior Blin in St-Gérand-le-Puy (MN2a), C. gergoviensis direction. Th e posterior terminus of the sagittal crest Viret, 1929 from plateau de Gergovie (Puy-de- is broken at the point where the braincase surface Dôme), and C. bugtiensis Cooper, 1923 from the begins to slope ventrally. Up to this point the sagittal early Miocene Dera Bugti fauna of Baluchistan. crest shows no sign of increasing depth, which is no Since Beaumont’s revision, three additional spe-

766 GEODIVERSITAS • 2009 • 31 (4) Phoberogale (Mammalia, Ursidae, Carnivora) from California

A

BC

FIG. 6. — Phoberogale bugtiensis Forster-Cooper, 1923, BMNH M12338, holotype from Dera Bugti, Baluchistan, Pakistan; lateral (A) and occlusal stereo (B, C) views of left maxillary. Scale bar: 20 mm. cies have been added: C. ginesticus Kuss, 1962 almost certainly includes a stratigraphic component, from Ginestous (MN1) in the Toulouse region in as well as sexual dimorphism, but lacking detailed southwestern France, C. meschethense Gabuniya, documentation for most of the classic European 1964 from the late Oligocene upper Pestrotsvet collections, one cannot be certain. We can turn to Svita of Georgia, and C. ursinus from the early Mi- the Bridgeport sample, however, where m1 length ocene of Paulhiac (Lot-et-Garonne, MN1). More ranges from 23.4 to 30.7 mm (N=16), with a mean recently, in his biometric analysis of Cephalogale, of 28 mm, an SD of 2.16, and a coeffi cient of vari- Bonis (1973) brought in an additional species, C. ation of 7.7. Th ese data support the assumption fi lholi (Munier-Chalmas, 1893) Munier-Chalmas that 12-18 mm may well be a conceivable range 1893), but excluded C. cadurcensis. for a single species. Of the above, the most basal species appears to be C. minor based on numerous specimens from the CEPHALOGALE AND PHOBEROGALE RECORDS IN ASIA classic Quercy fi ssure fi lls. A clear understanding of Five nominal records of Cephalogale or Phobero- the systematics of this species, however, is hampered gale have been reported from Asia, most of which by a lack of stratigraphic data for the majority of are fragmentary. Th e earliest recognized species is material that was collected more than a century ago. Cephalogale bugtiensis Forster-Cooper, 1923 from A sample of several dozen specimens examined by “the early Miocene deposits of Dera Bugti” of Beaumont encompasses a large size range. For ex- Baluchistan. Th e holotype of C. bugtiensis, BMNH ample, Beaumont measured 50 dentaries where m1 M12338, is a fragment of left maxilla with a partial length ranges from 12-18 mm. Such a large range P3 and complete P4-M2 (Fig. 6). Th e poor state of

GEODIVERSITAS • 2009 • 31 (4) 767 Wang X. et al.

preservation and advanced wear on the teeth do not Cephalogale record. Th e status of the Nadu record allow confi dent interpretation of its morphology, thus remains to be confi rmed. particularly the well worn M2. Beaumont (1965) Wang and Qiu (2003) reported the occurrence even questioned its membership in Cephalogale, and of Cephalogale sp., based on a single left M2 (IVPP commented that most of its characters are more V12429), from the late early Oligocene Saint- primitive than the European C. minor. Th e size of Jacques region in the Wulanbulage Formation, C. bugtiensis is nearly identical to OCPC 21794. Inner Mongolia, China. Th e authors remarked that Cephalogale bugtiensis mainly diff ers from P. shareri the Chinese material is closest to C. minor in size. n. sp. in a more pronounced notch between the base However, at a length and width of 7.7 × 5.0 mm, of the P4 paracone and protocone, less prominent IVPP V12429 would be among the smallest indi- anterior ridge on P4 paracone, and more medi- viduals of Cephalogale because M2s in European olaterally broad M1-2. Upper molars in the Bugti C. minor range from 6.1-7.4 mm in length and specimen, therefore, have an anteroposteriorly nar- 8.0-10.0 mm in width (Beaumont 1965). Th e In- row appearance, especially the M2. Th e internal ner Mongolian specimen does have a posteriorly cingulum (hypocone) of M1 is also more thickened oriented postprotocrista-metaconule ridge that ends at the posterolingual corner. Cephalogale bugtiensis at the posterior border of the tooth, an important also appears to have a lower-crowned P4 protocone character for early ursids (Beaumont 1982). Th e M2 compared to the California specimen. Both in size alone is insuffi cient to establish its generic identity, and overall morphology P. shareri n. sp. is closest and Wang et al. (2005: 42) concluded that IVPP to C. bugtiensis, and it is likely that the Bugti form V12429 may be possibly conspecifi c with Pachy- should be referred to Phoberogale. cynodon tedfordi Wang and Qiu, 2003. Th e second record is Cephalogale meschethense Most recently, Wang et al. (2005: fi g. 15a-c) re- Gabuniya, 1964 from the late Oligocene upper ferred to “?Cephalogale sp.” a left mandibular frag- Pestrotsvet Svita of Georgia (see also Gabuniya ment with m2 (MAE SG.97.5396) from the early 1966). Known from one m1, two canines, and a Oligocene Hsanda Gol Formation, eastern Valley fragmentary humerus, C. meschethense cannot be eas- of Lakes, central Mongolian People’s Republic. ily compared with other Asian records and thus has However, as conceded by Wang et al., too little was received little attention after its original description. preserved to be certain of its precise taxonomy. However, from Gabuniya’s fi gure (1966: fi g. 4a), C. meschethense does appear to have a posteriorly PHYLOGENETIC COMMENTS off set metaconid (relative to the protoconid) on Since its initial recognition, Cephalogale Jourdan, m1, a character of basal ursids. 1862 has been variously included in canids, amphi- Tang & Qiu (1979) listed Cephalogale in the cyonids, procyonids, amphicynodontines, and late Eocene Nadu Formation in Guangxi Province, simocyonines (e.g., Schlosser 1888; Teilhard de China. Unfortunately, no description was given Chardin 1915; Simpson 1945; Hough 1948). In for the material, which was subsequently missing the last 50 years, however, it has become increas- (Qiu & Li 2003; Wang & Qiu 2003), and the ingly clear that the genus occupies a basal position authors only commented that “judging from the in the Family Ursidae (Beaumont 1965; Bonis 1973; carnassial, the specimen from Nadu Formation is Beaumont 1982; Ginsburg & Morales 1995, 1998; more primitive than those from Europe and ac- McKenna & Bell 1997; Hunt 1998a). cordingly its age may be earlier than Oligocene” Of the 11 or so species recognized by Bonis (Tang & Qiu 1979: 409). In revisiting the issue, (1973), early workers have long recognized certain the junior author of the original report could not morphologic patterns that might indicate discrete confi rm their initial identifi cation (Qiu 2003: 21). lineages within the genus. Viret (1929: 140) divided Zhai et al. (2003), however, recently described a new Cephalogale into two species groups: those with com- amphicyonid species, Guangxicyon sinoamericanus, plex and trenchant premolars (including C. minor, from the Nadu Formation with no mention of the C. brevirostris, C. gracile, C. bonali, and C. depereti)

768 GEODIVERSITAS • 2009 • 31 (4) Phoberogale (Mammalia, Ursidae, Carnivora) from California

and those with large, simple premolars (including nassials, in contrast to the phoberocyonines with C. gergoviensis and C. geoff royi). Beaumont (1965) more hypercarnivorous dentitions. Th is phylogenetic arrived, less explicitly, at a similar conclusion regard- theme was further elaborated by Ginsburg (1999) ing some of Viret’s taxonomic arrangements. Bonis with slight modifi cations (such as adding C. gracile (1973), and also Bonis & Guinot (1987), on the to the hemicyonine lineage). Ginsburg (1999: 120) other hand, attempted to delineate three lineages diagnosed the Hemicyoninae as “Ursids with carnas- of Cephalogale based on proportional relationships sials lower than in Phoberocyoninae, and elongated of p4, m1, and m2. He recognized a bonali-depereti upper molars,” and “the premolars (P4 excepted) lineage that has a relatively long p4 and low m2/p4 reduced and the upper molars elongated, wider, with ratio. At the other extreme is a brevirostris-gracile- a tendency to become inscribed in a square by the ursinus lineage that features a relatively short p4 increase of the postero-lingual cingulum.” and high m2/p4 ratio. Somewhat in between is Most recently, in a cladogram of North Ameri- a geoff royi-gergoviensis-ginesticus lineage that has can fossil ursids, Hunt (1998a: fi g. 10.3) included intermediate values for the above measurements four genera in the New World Hemicyoninae that and ratios. However, Bonis (1973) refrained from placed Cephalogale (s.l.) as a basal taxon, followed giving generic status to these lineages, even though by Phoberocyon, , and an unnamed ge- he did recognize that certain Cephalogale lineages nus. His hemicyonine clade is defi ned by elongated are antecedent to more advanced hemicyonines. paraxonic feet and digitigrade postures, accompa- In their studies of a new early Miocene Spanish nied by key craniodental traits incorporating the genus and species, Ginsburg & Morales (1995) Type A ursid auditory bulla and ursid upper molar formally recognized three lineages within what pattern. Th e Phoberocyon-Plithocyon clade was de- had been traditionally regarded as a heterogene- fi ned by a premasseteric fossa on the dentary. Hunt ous Cephalogale (s. l.) (see also Ginsburg 1999). A in his cladogram described Cephalogale as having basal “C. minor” (genus undefi ned) group gave rise sectorial carnassials, simple conical premolars, to two clades: subfamilies Phoberocyoninae and M1 subtriangular to subquadrate, M2 quadrate Hemicyoninae. At the base of their newly recognized to subquadrate, a short snout, and auditory bulla phoberocyonines (Phoberocyonini of Kordikova et somewhat infl ated, and presented summary char- al. 2000) is a new genus Phoberogale Ginsburg & acteristics of the genus in greater detail later in Morales, 1995, which includes three species pre- this publication (Hunt 1998a: 180-181). Hunt’s viously placed in Cephalogale: P. bonali (Helbing cladistic hypothesis is similar to that proposed by 1928), P. depereti (Viret 1929), and P. ginesticus (Kuss Ginsburg & Morales (1995), although he did not 1962). Phoberogale in turn gave rise to the more treat individual species of Cephalogale. derived Phoberocyon and Plithocyon. Phoberocyon is Ginsburg & Morales’ (1995) latest attempt at characterized by well-developed posterior accessory sorting individual species of Cephalogale into a cusps on p3-4, elongated m1, distinct paraconid cladistic framework helped to focus previous con- on m2, and m2 protoconid higher than metaconid cepts (such as those of Bonis 1973) that have not (Ginsburg 1955; Tedford & Frailey 1976; Qiu et been taxonomically formalized. In actual practice, al. 1986), characters that are further elaborations their Phoberogale and restricted Cephalogale are from those in Phoberogale. distinguished by size as well as tendencies toward Th e redefi ned Hemicyoninae of Ginsburg & hyper- or hypocarnivorous dentitions (Ginsburg & Morales (1995: fi g. 10), a sister clade to Phobero- Morales 1995: fi g. 9). Th e California Phoberogale cyoninae, now includes the type species Cephalogale shareri n. sp. appears to fall on the phoberocyonine geoff royi plus two related species, C. ursinus and side of the clade with such hypercarnivorous char- C. brevirostris, at the base of the hemicyonine clade, acters as presence of a posterior accessory cusp on followed by more derived forms such as Zaragocyon, p4, p4 enlarged relative to p3, relatively elongated , and . Th e hemicyonines have carnassials at the expense of tuberculate molars, and strong tuberculate molars at the expense of the car- m2 proto conid taller than metaconid.

GEODIVERSITAS • 2009 • 31 (4) 769 Wang X. et al.

ZOOGEOGRAPHIC REMARKS de Bonis, a colleague of the fi rst three authors. His analysis of European Cephalogale in 1973 is still the Th e majority of North American immigrants used most authoritative treatment and forms a founda- to defi ne or characterize Neogene NALMAs (Ted- tion on which this paper is based. It is thus fi tting ford et al. 1987; 2004) are commonly assumed to to describe the new North American Phoberogale come from Asia. However, their actual Asian an- specimen in honor of Louis de Bonis. cestors mostly remain poorly known (Qiu 2003). Mark A. Roeder of Paleo Environmental Associ- Th e emerging records of Eurasian Cephalogale and ates, Inc., supervised the recovery of the holotype, Phoberogale or more accurate calibration of their which subsequently was prepared and cast by Howell ages permit additional comments on the immigra- W. Th omas at the Natural History Museum of Los tion of the latter to North America. Angeles County. Recent fi eld work on the Dera Bugti section by Th e senior author would like to express his appre- French vertebrate paleontologists indicates that the ciation for the support of this research by Mr Kevin lower Miocene terrestrial deposits (~250 m) above Sharer, Chairman of the Board of Trustees of the a sequence of Eocene marine sediments are faunally Natural History Museum of Los Angeles County. correlated to the European MN 3-4 units in the range For collection access, the authors thank the follow- of 17-20 Ma (Welcomme et al. 1997). Unfortunately ing individuals for their generous assistance: Louis no new material of Phoberogale bugtiensis was found de Bonis of the Université de Poitiers; the late John and its precise stratigraphic position is thus still in D. Cooper of California State University, Fuller- question. Based on rodent faunal considerations, Flynn ton, and the OCPC for the loan of the holotype; et al. (1986) also speculated that the Bugti fauna is Andy Currant of the British Museum of Natural between 18-21 Ma, an estimate roughly equivalent History; Burkart Engesser of the Naturhistorisches to the North American occurrence of Phoberogale Museum Basel; Léonard Ginsburg and Pascal Tassy spp. Qiu (2003: 21) also deduced that the dispersal of Muséum national d’Histoire naturelle; and Wang of Cephalogale (Phoberogale) from Europe to North Banyue of Institute of Vertebrate Paleontology and America should have occurred at ~21 Ma. Hunt’s Paleoanthropology. Stéphane Peigné provided a (2004) estimate of 23 Ma indicates a somewhat ear- translation of the French résumé. Th e fi rst three lier immigration event, based on a radioisotopically authors of this paper benefi ted from discussions calibrated occurrence in the Harrison Formation of with Demberelyin Dashzeveg of the Paleontological Nebraska, as yet undescribed. Despite the poor holo- Institute of Mongolian Academy of Sciences, who type of Phoberogale bugtiensis, size and morphological provided them with an unpublished manuscript similarities shared with those of P. shareri n. sp. seem on new examples of Cephalogale from the Hsanda to suggest that P. bugtiensis is the closest known sister Gol Formation. Th e authors thank Lars Werdelin species of the North American clade. If so, it also im- and Manuel J. Salesa for helpful reviews. plies that Phoberogale shareri n. sp. predates P. bugtiensis Travel to museums was supported by funds from and so a much wider geographic distribution for the the Knowledge Innovation Program of the Chinese genus must have existed in the early Miocene. Th is Academy of Sciences (KZCX2-YW-120), Major hitherto unknown record probably existed in much Basic Research Projects (2006CB806400) of MST of Eastern Asia, in latitudes higher than its known of China, the CAS/SAFEA International Partner- South Asia location, in order to avail itself of the op- ship Program for Creative Research Teams, and portunity to migrate to North America. the United States National Science Foundation (EAR 0446699).

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Submitted on 20 January 2009; accepted on 6 June 2009.

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