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Preview Discovery of the First Early Cenozoic Euprimate (Mammalia) from Inner Mongolia

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3571, 11 pp., 3 figures May 16, 2007 Discovery of the First Early Cenozoic Euprimate (Mammalia) from Inner Mongolia XIJUN NI,1,2 K. CHRISTOPHER BEARD,3 JIN MENG,2 YUANQING WANG,1 AND DANIEL L. GEBO4 ABSTRACT Although it is widely thought that euprimates originated in Asia, the fossil record of early euprimates remains sparse there. We describe herein a new omomyid euprimate, Baataromomys ulaanus,n.gen.etsp.,basedonanisolatedrightlowerm2fromBumbanianstrataatWulanboerhe in the Erlian Basin of Inner Mongolia, China. In terms of the size and proportions of m2, Baataromomys ulaanus is intermediate between Eurasian and North American species that are usually assigned to Teilhardina. Morphologically, m2 of Baataromomys differs from that of Teilhardina and North American small-bodied omomyids (including Anemorhysis, Tetonoides, Trogolemur, and Sphacorhysis) in having a smaller paraconid that is less fully connate with the metaconid, a lower entoconid, a weaker crest connecting the metaconid with the entoconid, and aweakerbuccalcingulid.ThenewtaxonismuchsmallerandlowercrownedthanSteinius,agenus commonly regarded as a basal omomyid. Despite the substantial difference in size, the m2s of Baataromomys and Steinius share some important features, including a very broad talonid basin and a relatively low hypoconid and cristid obliqua. Given its early occurrence and primitive anatomy,Baataromomysmayeventuallyhelptoclarifythephylogeneticrelationshipsamongbasal omomyids, but more complete specimens will be required to test this possibility. Baataromomys brandti from the basal Wasatchian zone Wa-0 in the northern Bighorn Basin, Wyoming, was previouslyallocatedtoTeilhardina.However,several dentalfeatures sharedbyB. brandtiandB. ulaanus suggest that they are closely related. The co-occurrence of Baataromomys in Asia and NorthAmericaindicatesthatsmall-bodiedeuprimateswereabletodispersalacrosstheBeringian regionnearthe Paleocene–Eocene boundary. 1Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 100044, China ([email protected]). 2DivisionofPaleontology,AmericanMuseumofNaturalHistory. 3SectionofVertebratePaleontology,CarnegieMuseumofNaturalHistory,Pittsburgh,PA15213. 4DepartmentofAnthropology,NorthernIllinoisUniversity,DeKalb,IL60115. CopyrightEAmericanMuseumofNaturalHistory2007 ISSN0003-0082 2 AMERICAN MUSEUMNOVITATES NO. 3571 INTRODUCTION Altanius hasmore recentlybeeninterpretedas either a basal euprimate or a plesiadapiform Euprimates, or primates of modern aspect, (Rose and Krause, 1984; Gingerich et al., include living lemurs, lorises, tarsiers, and 1991; Beard and Wang, 1995; Ni et al., 2005). anthropoids, as well as extinct taxa such as Aside from the specimen described here, the adapiforms and omomyids that are closely onlyotherBumbanianomomyidsdescribedto relatedtothem(e.g.,Szalayand Delson,1979; date are Kohatius and Teilhardina asiatica. Szalayetal.,1987;RoseandBown,1991;Rose Kohatius is a poorly documented omomyid et al., 1994). To date, the oldest undoubted from the Kuldana Formation of northern euprimate fossils have been recovered from Pakistan that apparently includes multiple earliest Eocene strata in North America, species, only one of which, K. coppensi, has Europe, and Asia (Gingerich, 1986, 1993; Ni been named (Russell and Gingerich, 1980, et al., 2004). One potential exception, the 1987;Thewissenetal.,1997).Thephylogenet- poorly documented Altiatlasius koulchii from ic relationships of Kohatius remain unclear. thelatePaleoceneofMorocco,isoftendebated Teilhardina asiatica is based on a partial skull astheworld’sonlyknownPaleoceneeuprimate and relatively complete lower dentition from (Sige´ etal.,1990;Godinot,1994;Hookeretal., the Lingcha Formation in Hunan Province, 1999; Beard, 2006). The relatively modern China (Ni et al., 2004). According to the anatomy, high taxonomic diversity, and broad phylogenetic analysis of Ni et al. (2004), geographicrangeofearliestEoceneeuprimates T.asiatica is closely related to T. belgica, suggest that the group as a whole originated a species that occurs in earliest Eocene strata sometime before the beginning of the Eocene. of Belgium and France. In agreement with This conclusion is corroborated by molecular Szalay (1982), Ni et al. (2004) found that approachestophylogenyreconstruction,which North American species that are often typically recognize euprimates as one of three referred to Teilhardina may actually be more living members of the Euarchonta, along with closely related to younger and more derived tree shrews (Scandentia) and flying lemurs North American omomyids, such as Tetonius, (Dermoptera) (e.g., Eizirik et al., 2004). Tetonoides, and Anemorhysis. With the excep- Given the wide geographic distribution of tion of ‘‘Teilhardina’’ brandti, which was the earliest fossil euprimates, it is not surpris- originally described as the earliest Teilhardina ing that specialists have so far failed to reach from North America (Gingerich, 1993) and is any consensus regarding their ancestral place transferred to the new genus from Inner oforigin.Thehierarchicallynestedpositionof Mongolia here, systematic revision of these euprimates within the superodinal clades EuarchontaandEuarchontoglires,whoseoth- North Americanspecies is beyond the scope of erprincipalmembersareallwidelythoughtto this study. We provisionally refer them to have originated in Asia, points toward an Teilhardina (e.g., as ‘‘Teilhardina’’ americana) Asian origin for euprimates as well (Beard, in this paper. 1998, 2006; Beard and Dawson, 1999; Ni et The new euprimate was discovered by al., 2005). Despite the strong biogeographic screenwashing matrix from the Gomphos signal provided by this phylogenetic context, bed at Wulanboerhe in the Erlian Basin of some recent workers continue to advocate Inner Mongolia in 2003. Meng et al. (2004) a Gondwanan origin for euprimates (e.g., described dental and postcranial remains of Miller et al., 2005). the gliriform mammal Gomphos elkema from Although phylogenetic evidence suggests this site, which they called ‘‘Huheboerhe’’. that euprimates originated in Asia, the fossil The locality is close to the site ‘‘7 miles record of early euprimates remains sparse southwest (235u) of Camp Margetts’’ of the there. The phylogenetic affinities of Altanius Central Asiatic Expeditions. Locality and orlovi, a tiny species known from Bumbanian stratigraphic data for Wulanboerhe have strata in the Nemegt Basin of Mongolia, are been published (Meng et al., 2004; Bowen et currently disputed. Originally described as an al., 2005). New field observations indicate omomyid (Dashzeveg and McKenna, 1977), that Paleogene beds of the Wulanboerhe- 2007 NI ETAL.: ASIAN CENOZOICEUPRIMATE 3 Huheboerhe-Nuhetingboerhe area consist of American ‘‘Teilhardina’’ species in having three lithological units: the Nomogen, a smaller paraconid that is more closely Arshanto, and Irdin Manha formations. At appressed to the metaconid, and a relatively the Wulanboerhe transect, only the Nomogen lower hypoconid and cristid obliqua. m2 and Arshanto formations are present, con- further differs from that of North American trary to Meng et al. (2004) and Bowen et al. ‘‘Teilhardina’’ species in having a lower en- (2005). The Gomphos bed occurs in the upper toconid, a much weaker crest connecting the part of the Nomogen Formation, the lower entoconid with the metaconid, and a weaker part of which yields a typical late Paleocene buccal cingulid. (Gashatan)fauna,characterizedbythepresence INCLUDEDSPECIES: TeilhardinabrandtiGin- ofLambdopsalis,Prionessus,andPalaeostylops. gerich, 1993:322. Immediately above the Gomphos bed are red GEOLOGICAL RANGE: Bumbanian of Inner beds that contain a species of perissodactyl Mongolia and earliest Wasatchian (Wa-0) of similar to Pataecops parvus and an arctostylo- Wyoming. pid similar to, but significantly larger than, ETYMOLOGY: Baatar,Mongolian,hero,wide- Palaeostylopsmacrodon.WefollowMengetal. ly used as Mongolian first name; and omomys, (2004) in referring the Gomphos bed to the Greek,shoulder-mouse. Bumbanian Land Mammal Age. Institutionalabbreviationsusedinthisreport Baataromomys ulaanus, new species are as follows: IVPP, Institute of Vertebrate figure 1 Paleontology and Paleoanthropology, Chinese Academy ofSciences(Beijing,China);MNHN, Museum National d’Histoire Naturelle (Paris, HOLOTYPE: IVPPV14614,anisolatedright France); UM, University of Michigan Museum m2; only known specimen. ofPaleontology(AnnArbor,Michigan);USGS, TYPE LOCALITY AND AGE: Wulanboerhe, United States Geological Survey (Denver, Erlian Basin, Inner Mongolia. Bumbanian of Colorado); USNM, United States National Inner Mongolia. Museum of Natural History (Washington, DIAGNOSIS: m2 differs from that of B. D.C.); UW, University of Wyoming Geolo- brandti in having a weaker paracristid and gical Museum (Laramie, Wyoming). a paraconid that is more lingual and less mesial in position. SYSTEMATIC PALEONTOLOGY ETYMOLOGY: Ulaan, Mongolian, red. The only known specimen was recovered from PRIMATESLINNAEUS,1758 a prominent Bumbanian red bed in the Erlian Basin. OMOMYIDAETROUESSART,1879 Baataromomys, new genus DESCRIPTION TYPESPECIES: Baataromomys ulaanus,new The single isolated right lower molar species. (fig. 1) can be confidently identified as m2 DIAGNOSIS: Smaller than all other omo- because its trigonid is mesiodistally com- myids except Teilhardina, Anemorhysis, Teto- pressed and lingually closed. The tooth as noides, Trogolemur, and Sphacorhysis. m2 a whole is remarkably bunodont and low- differs from that of Trogolemur, Sphacorhysis, crowned, and most of its individual cusps Anemorhysis, and Tetonoides inhavinga smal- and crests are relatively blunt and rounded. lerparaconidthatislessfullyconnatewiththe The molar has a relatively square occlusal metaconid; a lower, less projecting entoconid; outline, being 1.79 mm long, 1.47 mm wide amuchweakercrestconnectingthe metaconid at the trigonid, and 1.52 mm wide at the with the entoconid; a weaker buccal cingulid; talonid. In terms of its size and proportions, and in lacking a mesially expanded and shelf- m2 of Baataromomys ulaanus falls between like paracristid. m2 differs from that of samples of Eurasian Teilhardina and North Teilhardina asiatica, T. belgica, and North American ‘‘Teilhardina’’ (fig. 2). 4 AMERICAN MUSEUMNOVITATES NO. 3571 The trigonid as a whole is slightly inclined mesially. The weakly developed paracristid descends the mesial face of the protoconid beforeturninglinguallytounitethelattercusp with the paraconid. The tip of the protoconid and the buccal arm of the protocristid are worn to expose the dentine, forming a small depression that tapers distolingually. The metaconid is roughly the same size as the protoconid and is situated almost directly lingual to the latter cusp. Its distolingual corner forms a blunt and insignificant ridge, the postmetacristid, that descends the lingual face of the postvallid. The mesial surface of the metaconid is smooth, bearing no trace of a premetacristid. The lingual arm of the protocristid projects buccally from the meta- conid, meeting the buccal arm of the proto- cristid near the midline of the tooth. A transverse line of dentine joins the tip of the metaconid with the lingual arm of the proto- cristid. The wear facets on the protocristid suggestthat thecrest wasveryblunt,and that its lingual and buccal arms probably did not form a continuous shearing crest. The para- conid is very small, occurring at the mesiolin- gual corner of the trigonid. A tiny pit of dentine is exposed at the apex of the para- conid, indicating that the cusp was low even when it was unworn. No crest connects the paraconid and metaconid. The topography of the internal part of the trigonid is unusual in that the region between the protoconid and metaconid is actually elevated above the level of the protocristid and paracristid. Thetalonidislongerandslightlywiderthan the trigonid. The hypoconid is the dominant cusp, but it and the cristid obliqua remain relativelylowandblunt.Thelingualsurfaceof the hypoconid is smooth and slopes down to thetalonidbasin.Itsbuccalanddistalsurfaces are continuous, forming a rounded distobuc- cal corner. The cristid obliqua joins the distal wall of the trigonid beneath the protoconid, resulting in a shallow hypoflexid. The buccal Fig. 1. Baataromomys ulaanus, n. gen. et sp., holotype, IVPP V14614, isolated right m2. A, arm of the postcristid is low and blunt, Occlusal view; B, lingual view; C, buccal view. connecting the hypoconulid with the cristid Scalebarindicates 1.00 mm. obliqua at about a right angle. The tip of the hypoconidisworntoexposeasmallroundpit ofdentine.Thecristidobliquaisslightlyworn, exposing a tiny line of dentine. The entoconid forms the distolingual corner of the talonid. 2007 NI ETAL.: ASIAN CENOZOICEUPRIMATE 5 Fig. 2. Size of m2 in the holotypes of Baataromomys ulaanus, n. gen. et sp. and B. brandti, shown in relation to those of Eurasian Teilhardina and North American ‘‘Teilhardina’’. Measurements of North American ‘‘Teilhardina’’ are taken from Bown and Rose (1987). Vertical and horizontal axes are inmillimeters. This cusp is much smaller and lower than the leaving the talonid basin open lingually. The hypoconid, so that the deepest point of the lingual arm of the postcristid is insignificant talonid basin lieslingualtothe midline of this and fuses seamlessly with the transversely structure. The mesiobuccal surface of the elongated hypoconulid. The hypoconulid is entoconid is smooth but concave, forming worn, exposing the dentine. No significant the distolingual border of the talonid basin. cusp remains, but it must have been relatively The entocristid is very low and blunt. It does large in its unworn condition, based on the not connect with the weak postmetacristid, exposedislandofdentine.Nosulcusisformed 6 AMERICAN MUSEUMNOVITATES NO. 3571 between the entoconid and hypoconulid, nor Donrussellia. For example, both have very between the hypoconulid and hypoconid. A broad and shallow talonid basins, with hypo- thinbutdistinctcingulidsurroundsthebuccal conid and entoconid cusps that are not very halfofthetooth.Itoriginatesfromthebaseof elevated. In buccal view, the hypoconid and the paracristid at its mesiobuccal side, con- cristid obliqua on m2 in Baataromomys are tinues distobuccally, envelops the hypoconid, very low, like those of Donrussellia and in and terminates just below the buccal side of contrast to those of Teilhardina. Like the hypoconulid. Donrussellia, Baataromomys retains a small but distinct hypoconulid on m2. COMPARISONS Altanius orlovi differs markedly from Baataromomys ulaanus in many aspects of Baataromomys differs enough from other m2 morphology. In Altanius, m2 is smaller, basal omomyids that it is also necessary to relativelynarrower,andmuchhighercrowned compare it with basal adapiforms, such as than that of Baataromomys. The trigonid of Donrussellia, as well as the enigmatic primate m2inAltaniusismuchhigherthanitstalonid, Altanius (fig. 3). bears three high and conical cusps, and has Thetrigonidofm2inB.ulaanusdiffersmost a more triangular occlusal outline. The para- notably from that of Donrussellia gallica and conid is slightly larger than the metaconid on D.provincialisinthepositionofthemetaconid m2 in Altanius, and these cusps are fused at and the orientation of the protocristid. In their bases to form a lingual pair of trigonid Donrussellia, the metaconid of m2 is more cuspsthatarehigherthantheprotoconid.The distal in position, which contributes to the protoconid and hypoconid of Altanius are relative isolation of this cusp. The relatively morepointedandbearmoresignificantbuccal distallocationofthemetaconidinDonrussellia ridges, whereas the hypoconulid is much also results in a more oblique, rather than weaker. In Altanius, the cristid obliqua of m2 transverse, orientation for the protocristid. In is connected to the postvallid below the contrast, the metaconid of m2 in protoconid-metaconid notch, resulting a very Baataromomys is located only slightly distal deep hypoflexid. In contrast, the cristid to the protoconid, so that the protocristid is obliqua of Baataromomys is more buccally nearly transverse. In addition to the wider oriented, thereby forming a very shallow separationbetween the paraconidand metaco- hypoflexid. The talonid basin of Altanius is nid on m2 in Donrussellia compared with much deeper and narrower than that of Baataromomys, the paraconid in the former Baataromomys, and its lingual side is closed genus is relatively larger and its base is not by a relatively large entoconid and a high fusedwiththebaseofthemetaconid.Inlingual entocristid.These important differences inm2 view,thetrigonidofm2inDonrusselliaismore morphology rule out any possibility of close deeply notched as a result. These lower molar affinitiesbetweenAltaniusandBaataromomys. trigonid features distinguish Donrussellia from Teilhardina asiatica is the oldest undoubted all omomyids, so the fact that Baataromomys euprimate known from Asia (Ni et al., 2004). differs from Donrussellia in these respects is ItcloselyresemblesEuropeanT.belgicabutis unremarkable. On the other hand, there are morphologically even more primitive. T. severalwaysinwhichBaataromomysresembles asiatica and T. belgica share some similarities R Fig. 3. Schematic drawings of m2 morphology in Baataromomys and a comparative sample of early Eoceneeuprimates.Allspecimensarescaledtothesamemesiodistallengthandaredepictedinocclusal(left column), buccal (middle column), and lingual (right column) views. Taxa and specimens illustrated are as follows:Baataromomysulaanus,holotype,IVPPV14614(A–C);Baataromomysbrandti,holotype,UM99031 (D–F); Teilhardina asiatica, holotype, IVPP V12357 (G–I); ‘‘Teilhardina’’ americana, holotype (reversed), UW 6896 (J–L); Anemorhysis sublettensis, holotype (reversed), USNM19205 (M–O); Tetonoides pearcei, holotype,USNM22426(P–R);Steiniusvespertinus,USGS25027(S–U);Donrusselliaprovincialis,holotype (reversed), MNHNRI-170 (V–X).Each scale barindicates 1.00mm. 2007 NI ETAL.: ASIAN CENOZOICEUPRIMATE 7 8 AMERICAN MUSEUMNOVITATES NO. 3571 in m2 morphology with B. ulaanus, including Baataromomysbrandti, originally referredto a relatively buccal position of the cristid Teilhardina, is the earliest haplorhine primate obliqua, a shallow hypoflexid, and a lingually knownfromNorthAmerica(Gingerich,1993). open talonid basin. On the other hand, Theholotypeofthisspeciesisanisolatedright differences in m2 morphology between these m2 from the earliest Wasatchian (Wa-0) of two species of Teilhardina and B. ulaanus are northwestern Wyoming. This species shares significant. In contrast to the bunodont manysimilaritieswithB.ulaanus.Inbothtaxa, condition in B. ulaanus, the m2 in T. asiatica the trigonid of m2 is mesiodistally narrow; the andT.belgicaexhibitsrelativelysharpercusps paraconid, metaconid, and protoconid are andmoretrenchantcrests.Inocclusaloutline, rounded; the paraconid is not twinned with thetrigonidofm2inT.asiaticaandT.belgica the metaconid; the talonid basin is broad and is more triangular than is the case in B. shallow;crests(includingthepostcristid,cristid ulaanus. While being smaller than the proto- obliqua, entocristid, and postmetacristid) are conid and metaconid, the paraconid of m2 in lowandblunt;andthehypoconulidisrelatively T. asiatica and T. belgica is not as reduced as prominent.B.brandtiisdistinguishedfromthe that of B. ulaanus. In lingual view, the new taxon in having a slightly stronger para- paraconid on m2 protrudes mesially in T. cristid and a paraconid that is located less asiatica and T. belgica, but this cusp is fused lingually. with the base of the metaconid in B. ulaanus. ‘‘Teilhardina’’ americana is a relatively The talonid of m2 in B. ulaanus differs from primitive omomyid. Szalay (1982) argued that that of T. asiatica and T. belgica in being ‘‘T.’’americanaistooderivedtobereferredto broader, in having a less elevated hypoconid Teilhardina. Instead, Szalay (1982) synony- and cristid obliqua (especially apparent in mized ‘‘T.’’ americana with Anemorhysis te- buccalview),andinhavingamoreprominent nuiculus. Bown and Rose (1987) maintained hypoconulid. a species-level distinction between ‘‘T.’’ amer- Kohatius is another early Eocene omomyid icana and A. tenuiculus, but they acknowl- genus from Asia. Documented specimens, edged the close affinities between them, re- includingarightm1,asmallfragmentofaleft ferring both species to Teilhardina. Compared p4, a right mandible fragment preserving p4 with that of B. ulaanus, m2 in ‘‘T.’’ americana andthealveoliform1–m2,andanisolatedleft has taller cusps and sharper crests. The p4, are from the Kuldana Formation of paraconid of m2 in ‘‘T.’’ americana is much northern Pakistan (Russell and Gingerich, larger and protrudes farther mesially than is 1980, 1987; Thewissen et al., 1997). These the case in B. ulaanus, yielding a more tri- specimens may represent as many as three angular occlusal outline for the trigonid. differentspecies(RussellandGingerich, 1980, Together, the paraconid and metaconid on 1987; Thewissen et al., 1997). Because m2 m2 in ‘‘T.’’ americana form a massive, bi- remains unknown for Kohatius, direct com- cusped structure on the lingual side of the parisons with Baataromomys are not possible. trigonid, while the reduced paraconid of B. However, m1 of Kohatius has taller cusps and ulaanusseemstobemerelyavestigialappend- stronger crests than do those on m2 of B. ageofthemetaconid.Asubtlebutcontinuous ulaanus. Additionally in Kohatius, the para- crestconnects theapices oftheparaconidand conid and metaconid are widely separated on metaconid on m2 in ‘‘T.’’ americana. This m1, while these cusps (along with the trigonid structure becomes much better developed in as a whole) are notable for being closely some younger and more derived North compressed on m2 in B. ulaanus. Finally, the American omomyids, such as Tetonius, m1 talonid in Kohatius bears a tall and Tetonoides, and Trogolemur. In B. ulaanus, projecting entoconid and a relatively tren- no equivalent crest unites the paraconid and chant entocristid, in contrast to the more metaconid on m2. The talonid of m2 in ‘‘T.’’ bunodont conditions that prevail on m2 in B. americana is much deeper and more fully ulaanus. These morphological distinctions are closed than that of B. ulaanus. On m2 in ‘‘T.’’ too great to be attributed to different molar americana all of the crests surrounding the loci alone. talonid basin, as well as the hypoconid and 2007 NI ETAL.: ASIAN CENOZOICEUPRIMATE 9 entoconid, are stronger and more elevated relatively simple cusp pattern, Baataromomys than is the case in B. ulaanus, but the lacksanycompellingfeaturesincommonwith hypoconulid remains much less significant. Teilhardina. Because both of the aforemen- ‘‘T.’’ americana also has a stronger buccal tioned characters are almost certainly primi- cingulid on m2 than does B. ulaanus. tive for Omomyidae, there is no reason to Steiniushasbeenrankednearthebaseofthe presume that Baataromomys is more closely omomyid evolutionary radiation based on its relatedtoTeilhardinathanitistoSteinius,for primitive dental anatomy, despite the fact that example. it occurs stratigraphically well above the Several of the distinctive molar characters earliest North American omomyids (Rose and of Baataromomys diverge from conditions Bown,1991;Roseetal.,1994).Bothspeciesof found in other omomyids, thereby inviting Steinius are much larger than B. ulaanus, but comparisons with other higher level taxa of these taxa share many features of molar primates. For example, m2 of Baataromomys morphology in common. The most significant has bunodont and slightly inflated cusps, difference between Steinius and Baataromomys trigonidandtalonidof similar width, a cristid relates to the larger size and more mesial obliqua that reaches the postvallid near the positionoftheparaconidonm2intheformer. base of the protoconid, a shallow hypoflexid, On the other hand, both taxa possess very a moderately developed hypoconulid, and broad talonids and relatively low hypoconids a weak buccal cingulid. These features have andobliquecristids.Cuspsandcrestsaremore often been considered as derived characters trenchant inSteinius, whereas the entirecrown amongprimates,andKayandWilliams(1994) of m2 is more bunodont in Baataromomys. included them in their description of the Given the large difference in size, the morpho- morphotype of the hypothetical anthropoid logical similarities between these taxa are ancestor. The potential significance of these surprising. characters in Baataromomys is difficult to assess at present, given the very limited DISCUSSION anatomicalinformationthatiscurrentlyavail- able for this taxon. Baataromomys may AlthoughBaataromomysulaanusiscurrent- eventually help to clarify the phylogenetic ly documented by a single m2, several dental relationships among basal omomyids and features shared by B. ulaanus and B. brandti other euprimate clades, but more nearly suggest that they are closely related. These complete specimens of this taxon will be characters include a mesiodistally compressed required to test this possibility. trigonid, bunodont and rounded protoconid Along with Teilhardina asiatica, Baataro- and metaconid cusps, a low talonid with momys ulaanus documents at least modest a broad basin, and a prominent hypoconulid. diversity in the Asian euprimate radiation at Despite their widely divergent geographic or near the beginning of the Eocene. Such provenances, we think that the combination precociouseuprimatediversityinAsiaishardly of these traits indicates that Asian B. ulaanus surprising. Indeed, early Asian euprimates and North American B. brandti are more must remain significantly undersampled if the closely related to each other than either of phylogenetic evidence pointing toward an them is to other omomyids. We therefore Asian origin for this clade is accurate. On the transferNorthAmerican‘‘Teilhardina’’brand- other hand, if B. ulaanus and B. brandti prove ti to the genus Baataromomys. to be as closely related as current evidence B. brandti was previously interpreted as suggests, then Baataromomys would join nu- a potential common ancestor for European merous other mammalian taxa indicating Teilhardina belgica and North American dispersal across the Beringian region near the ‘‘Teilhardina’’ americana (Gingerich, 1993). Paleocene–Eoceneboundary(e.g.,Beard,1998; Based on our morphological comparisons, B. Beard and Dawson, 1999). Finally, the geo- brandti shares more features in common with graphic occurrence of Baataromomys ulaanus B. ulaanus than it does with any species of suggests that forested habitats suitable for Teilhardina. Aside from its small size and small euprimates must have been present, at 10 AMERICAN MUSEUMNOVITATES NO. 3571 least sporadically, on the Mongolian Plateau Dashzeveg,D.,andM.C.McKenna.1977.Tarsioid during the early Cenozoic. primate from the early Tertiary of the Mongolian People’s Republic. Acta Palaeonto- logicaPolonica22:119–137. ACKNOWLEDGMENTS Eizirik, E., W.J. Murphy, M.S. Springer, and S.J. O’Brien.2004.Molecularphylogenyanddating We thank Drs. Li Chuankui, Eric Delson, ofearlyprimatedivergences.InC.F.Rossand Alfred L. Rosenberger, Richard F. Kay, and R.F. Kay (editors), Anthropoid origins: new an anonymous reviewer for instructive dis- visions: 45–64. New York: Kluwer Academic/ cussion and suggestions, Sun Chengkai, Jin Plenum Publishers. Xun, and Bai Bin for assistance with field- Gingerich, P.D. 1986. Early Eocene Cantius torre- si—oldest primate of modern aspect from work, and Mark A. Klingler for help with NorthAmerica. Nature 320:319–321. illustrations. Financial support from the Gingerich, P.D. 1993. Early Eocene Teilhardina American Museum of Natural History and brandti: oldest omomyid primate from North the Carnegie Museum of Natural History is America. Contributions from the Museum of gratefully acknowledged. Funding was pro- Paleontology,UniversityofMichigan28:321– vided by grants from the National Natural 326. Science Foundation of China (40202003 to Gingerich, P.D., D. Dashzeveg, and D.E. Russell. X.N. and 40532010 to Y.W.) and U.S. 1991.Dentitionandsystematicrelationshipsof National Science Foundation (BCS 0309800 Altaniusorlovi(Mammalia,Primates)fromthe early Eocene of Mongolia. Geobios 24: to C.B. and J.M., and EAR-0120727 to 637–646. J.M.). Godinot, M. 1994. 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