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The first known Asian Hispanomeryx (Mammalia,Ruminantia, Moschidae)Israel M. Sánchez a , Daniel Demiguel b , Victoria Quiralte a & Jorge Morales aa Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC, C/ JoséGutiérrez Abascal 2, 28006, Madrid, Spainb Institut Català de Paleontologia, Universitat Autònoma de Barcelona, Edifici ICP, Campusde la UAB s/n, 08193, Cerdanyola del Vallès, Barcelona, Spain

Available online: 08 Nov 2011

To cite this article: Israel M. Sánchez, Daniel Demiguel, Victoria Quiralte & Jorge Morales (2011): The first known AsianHispanomeryx (Mammalia, Ruminantia, Moschidae), Journal of Vertebrate Paleontology, 31:6, 1397-1403

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Journal of Vertebrate Paleontology 31(6):1397–1403, November 2011© 2011 by the Society of Vertebrate Paleontology

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THE FIRST KNOWN ASIAN HISPANOMERYX (MAMMALIA, RUMINANTIA, MOSCHIDAE)

ISRAEL M. SANCHEZ,*,1 DANIEL DEMIGUEL,2 VICTORIA QUIRALTE,1 and JORGE MORALES1; 1Departamento dePaleobiologıa, Museo Nacional de Ciencias Naturales-CSIC, C/ Jose Gutierrez Abascal 2, 28006, Madrid, Spain,israelms@mncn.csic.es; 2Institut Catala de Paleontologia, Universitat Autonoma de Barcelona, Edifici ICP, Campus de la UAB s/n,08193, Cerdanyola del Valles, Barcelona, Spain, daniel.demiguel@icp.cat

Hispanomeryx Morales, Moya-Sola and Soria, 1981, is amoschid pecoran from the middle-early late Miocene of Eu-rope (MN 6 to MN 11; Morales et al., 1981; Azanza, 1986;Prothero, 2007; Vislobokova, 2007; Sanchez et al., 2009, 2010).Three species of Hispanomeryx have been currently described, allof them from Spain: the Aragonian H. aragonensis Azanza, 1986,and H. daamsi Sanchez, Domingo and Morales, 2010, and theVallesian Hispanomeryx duriensis Morales, Moya-Sola, and So-ria, 1981(type species). Besides Spain, very scarce Hispanomeryxremains have been discovered in Turkey and North Caucasus(Moya-Sola, 1986; Gentry, 1990, 1994; Pickford et al., 1999). Thislack of sufficiently abundant fossils of non-Iberian Hispanomeryxensured that the evolutionary history of the genus outside theIberian Peninsula was hitherto very poorly known. In additionto this, Hispanomeryx was considered to be restricted to the occi-dental Tethys, because, contrary to Micromeryx, no remains werediscovered East of the Caucasus so far (Sanchez et al., 2010).However, the classical American Museum of Natural HistoryCentral Asiatic Expeditions (AMNH CAE) of 1928–1930 (seeWang et al., 2003, for a thorough description of the AMNH CAE)yielded good fossil remains of ruminant faunas from the latemiddle Miocene of China (Tunggur Formation; Pilgrim, 1934;Colbert, 1936a, 1936b, 1940), and among these fossils stored atthe AMNH (New York) we discovered an extraordinary sampleof previously non-described moschid faunas from the TunggurFormation that includes both Micromeryx and Hispanomeryx re-mains.

We describe here the very first known Asian Hispanomeryx,from the locality of Wolf Camp (Tunggur Formation, China)excavated by the AMNH CAE in 1930 (Andrews, 1932; Wanget al., 2003), which give new insights into the evolutionary history,paleobiogeography, and morphological diversity of the genus.Also, we present the phylogeny of Hispanomeryx including alldescribed species.

MATERIALS AND METHODS

Fossil Material

The type series of Hispanomeryx andrewsi, sp. nov., comprisesall the available fossil material from Wolf Camp (site 822) cu-rated by the AMNH attributable to the genus Hispanomeryx. Allmeasurements are presented as Supplementary Data (availableonline at www.vertpaleo.org/jvp/JVPcontents.html). We followthe terminology of tooth elements and the set of measurements ofAzanza (2000) (English version in Sanchez et al., 2008), and alsofollow Barone (1999) for nomenclature of the postcranial skele-ton.

*Corresponding author.

Institutional Abbreviations—AMNH, American Museum ofNatural History, New York, U.S.A.

Cladistic Analysis

Hispanomeryx andrewsi, sp. nov., was included in a mod-ified data set from Sanchez et al. (2010) for the phylogenyof Hispanomeryx, with the addition of eight additional char-acters (morphology of the M2, relative size of M1/M2, mor-phology of the paracone rib, relative development of the M2parastyle, morphology of the p4, morphology of the third lobeof the m3, and metatarsal plantar tuberosity; Appendices 1–2).The data matrices were compiled in MacClade 4.05 and run inPAUP 4.0b10 (Macintosh versions). All characters were opti-mized as unordered, unweighted, and ACCTRAN. The data ma-trix included seven taxa and 26 characters. We chose the basalbovid Namacerus garepiensis Morales et al., 2003 (lower middleMiocene, Namibia, Africa), as the outgroup because the Bovi-dae have been repeatedly clustered with the Moschidae as theirsister group in a series of recently published phylogenetic worksinvolving both molecular and morphological data sets (Hassaninand Douzery, 2003; Guha et al., 2007; Marcot, 2007; Sanchezet al., 2010). A bootstrapped ‘branch and bound’ search with 1000replicates produced a single most parsimonious tree (MPT) of 31steps (CI = 0.935; RI = 0.9; HI = 0.065) with four internal nodes(Fig. 1). The character-state distributions for the internal nodesinvolving Hispanomeryx, as well as the autapomorphic states forall Hispanomeryx species are presented in the Table 1.

SYSTEMATIC PALAEONTOLOGY

MAMMALIA Linnaeus, 1758CETARTIODACTYLA Montgelard, Catzeflis and Douzery,

1997RUMINANTIA Scopoli, 1777

MOSCHIDAE Gray, 1821HISPANOMERYX Morales, Moya-Sola, and Soria, 1981

Type Species—Hispanomeryx duriensis Morales, Moya-Sola,and Soria, 1981.

HISPANOMERYX ANDREWSI, sp. nov.(Figs. 2–3)

Holotype—AMNH 92287, left hemimandible with p2–m3.Paratypes—The remaining referred material from Wolf Camp

(site 822).Etymology—Dedicated to Roy Chapman Andrews.Type Locality and Horizon—Wolf Camp (site 822), Tunggur

Formation, Moergen Fauna, late middle Miocene, China (seeWang et al., 2003).

Referred Material—AMNH 92287 (left partial mandible),AMNH 92286 (left partial mandible), AMNH 92267 (right par-tial mandible), AMNH 26499 (right partial mandible fragment),

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TABLE 1. Hispanomeryx andrewsi sp. nov. Character/state distribu-tions for the internal nodes A, B, C and all Hispanomeryx species. Italicsindicate ambiguous synapomorphies.

NODE/TAXON CHARACTER: STATE

Node A 7: 1; 11: 1; 12: 1Node B 10: 1; 13: 1H. aragonensis 19: 0H. duriensis 14: 1; 23: 1H. daamsi 17: 1; 22: 1H. andrewsi 4:1; 5:1; 8: 0; 10: 2

AMNH 26499a (right partial mandible fragment), AMNH 92268(right partial mandible), AMNH 92269 (left partial mandible),AMNH 92288 (right partial mandible fragment), AMNH 92265(left partial mandible fragment), AMNH 92265a (right partialmandible fragment), AMNH 92265b (left partial mandible frag-ment), AMNH 92265c (left partial mandible fragment), AMNH92265d (right partial mandible fragment), AMNH 92265e (rightpartial mandible fragment), AMNH 92270 (left maxilla), AMNH143433 (first phalanx).

Diagnosis—Large-sized Hispanomeryx with relatively well-developed protruding metastylids, strong M2 parastyle thatprojects buccally, and elongated M2 with triangular distalprofile. Besides these characters, Hispanomeryx andrewsi dif-fers from H. duriensis in: smaller size, lack of metaconule-fold in the upper molars and presence of cristid obliqua inthe p4. Differs from H. aragonensis in: distally closed thirdlobe of the m3, larger size, and more developed crown-height(hypsodonty). Differs from H. daamsi in: larger size, lack ofmetaconule-fold in the upper molars and non-lingually turnedprehypocristid.

Description

Lower Dentition—The Wolf Camp remains represent amoschid (see Sanchez et al., 2010, for the diagnostic charactersof the Moschidae) that shows all the dental diagnostic featuresof Hispanomeryx: lower premolar row significantly reduced (seeSupplementary Data), lack of Palaeomeryx-fold, very reducedm1 with respect to m2, column-shaped lingual structures, and flatlingual walls in the lower cheek teeth. The deciduous lower cheekteeth are nearly identical to that of H. duriensis and H. daamsi(see Sanchez et al., 2010). The lower premolars are also typicalfor the genus (see Sanchez et al., 2010), with the fully developedanterolingual cristid in the p4 that lingually closes the anteriorvalley. The p4 has a cristid obliqua. The cuspids in the lowermolars are less flattened and aligned than those of H. durien-sis and H. daamsi, more similar to those of H. aragonensis. Themetastylid of the m3 is more developed than in any other His-panomeryx. This structure can be very narrow and protruding,especially in several specimens such as AMNH 92267, AMNH92268, AMNH 26499a, and AMNH 92265a. By comparison, H.daamsi and H. duriensis lack or almost lack metastylids, and H.aragonensis presents poorly developed metastylids in all threelower molars. Hispanomeryx andrewsi has a distally closed thirdlobe of the m3, confirming that the open state of this characteris an autapomorphy of H. aragonensis (see Sanchez et al., 2010).The mesial cingulid is faint, as is typical of Hispanomeryx. With amoderate wear all cuspids become united by the cristids, with theexception of the pre-hypocristid.

Upper Dentition—The deciduous upper cheek teeth are notwell preserved, but they are very similar to that of H. daamsi (seeSanchez et al., 2010). DP2 and DP3 show well-developed buc-cal structures and the DP4 appears to have all cuspids united by

the cristae with a medium degree of wear. The upper molars ofH. andrewsi lack protoconal- and metaconule-folds (H. daamsiand H. duriensis both have metaconule-fold), and also entostylesand cingula. The buccal structures in the upper molars are morecolumnar than pyramid-shaped, and the internal walls are flat.The M2 has a marked distal triangular profile. The parastyle ofthe M2 is very developed, protruding perpendicular to the buccalwall. The buccal cusps of the M1 are less aligned than that of theM2.

First Phalanx—The first phalanx is not different from that ofother representatives of the genus as H. daamsi (see Sanchezet al., 2010, for a thorough description), including the open cen-tral sulcus of the proximal articular surface, the subrectangularexternal facet of the proximal articular surface, the long furrowfor the interosseus muscle, and the elliptic insertion area for theinterdigital ligament.

DISCUSSION

Hispanomeryx was recently nested within a monophyleticMoschidae (Fig. 1) defined as the clade comprising Micromeryx,Hispanomeryx, Moschus, and ‘Moschus’ grandeavus, their morerecent common ancestor and all of its descendants (Sanchezet al., 2010). This hypothesis excluded from the MoschidaeAmerican taxa such as the ‘blastomerycines’ and some Oligo-Miocene forms such as Dremotherium, regarding true moschidsas a derived Eurasian clade of bovoid pecorans with a middleMiocene origin (Sanchez et al., 2010). Our MPT is consistentwith that of Sanchez et al. (2010), showing a well-supported His-panomeryx clade with H. aragonensis as its most basal offshoot.Hispanomeryx andrewsi is the most autapomorphic of all His-panomeryx species. Besides the distally triangular-shaped M2,the other two autapomorphic traits of H. andrewsi are some-what surprising considering the tendency of node C taxa to de-velop very bovid-like lower molars, with almost non-existentmetastylids and very poorly developed metaconid lingual ribs,and the already reduced (albeit not so much) same charactersin the basal H. aragonensis. However, H. andrewsi presents rel-atively developed and protruding metastylids, especially the m3.This feature is more prominent in some specimens than in oth-ers, but it clearly contrasts with the remaining species in thegenus. Also, the parastyle in the M2 is very developed in H. an-drewsi, contrasting with the comparatively poorly developed M2parastyle of the three other Hispanomeryx species. Given thatspecific dental structures of extinct ruminants are found to be in-volved in the chewing process and seem to act by reducing stressduring occlusion (DeMiguel et al., 2006, 2009), the dental mor-phology of H. andrewsi could indicate a non-uniform biomechan-ical behavior. In fact, differences in the structural complexity ofthe molars are particularly notable when comparing buccal andlingual regions. According to the results of the above-mentionedstudies based on finite-element analyses (FEAs) upon several dif-ferent ruminant taxa, it is likely that the relatively strongly devel-oped ribs, styles and stylids in H. andrewsi, and more specificallythose found in the paracone and the metaconid, would sustainhigh levels of stress during chewing, thus improving mechani-cal efficiency for an effective breakdown of foods. It is not sur-prising to suggest, therefore, that such unexpected plasticity inthe dentition of Hispanomeryx may have been one of the fac-tors explaining the great evolutionary success of this genus dur-ing the Miocene (Sanchez and Morales, 2006; Sanchez et al.,2010).

Although the more ancient records of Hispanomeryx are stilllocated in western Europe (Sanchez and Morales, 2006; Sanchezet al., 2010), the description of H. andrewsi expands the bio-geographic range of the genus from a strictly occidental Tethys

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FIGURE 1. Hispanomeryx andrewsi, sp. nov. A, Phylogenetic position and taxonomical composition (at genus level) of the Moschidae. Modifiedfrom Sanchez et al. (2010). B, MPT showing the phylogenetic position of H. andrewsi. Bootstrap values are given for internal nodes A, B, and C(Hispanomeryx clade). C, Calibrated phylogenetic tree of the Moschidae, with special emphasis on Hispanomeryx. Black color represents WesternEurope-Spain (only Spain for Hispanomeryx); white color, Turkey-Caucasus; and grey color, Asia.

distribution to a broad Eurasian range. As pointed out bySanchez et al. (2010), Hispanomeryx was unknown outside theIberian Peninsula after the MN 6. Wang et al. (2003) corre-late the Tunggur Formation with the European MN 7/8, so His-panomeryx andrewsi not only sheds light on the presence of thegenus in a chronostratigraphic level and biogeographic area in

which it was unknown to occur, but also demonstrates that thetwo most important Miocene moschid genera, Micromeryx andHispanomeryx, were both distributed across Eurasia, and thatthe musk deer clade was indeed already present in Asia previ-ously to its extinction from Europe around the middle Turolian(Fig. 1).

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FIGURE 2. Hispanomeryx andrewsi, sp. nov., type series. A, holotype AMNH 92287 in buccal view; B, holotype in lingual view; C, holotype inocclusal view; D, paratype AMNH 92267 in buccal view; E, AMNH 92267 in lingual view; F, AMNH 92267 in occlusal view; G, paratype AMNH92265 in buccal view; H, AMNH 92265 in lingual view; I, AMNH 92265 in occlusal view; J, paratype AMNH 92265b in buccal view; K, AMNH 92265bin lingual view; L, AMNH 92265b in occlusal view; M, paratype AMNH 92268 in buccal view; N, AMNH 92268 in lingual view; O, AMNH 92268 inocclusal view; P, paratype AMNH 92286 in buccal view; Q, AMNH 92286 in lingual view; R, AMNH 92286 in occlusal view; S, paratype AMNH 92270in occlusal view; T, AMNH 92270 in buccal view; U, paratype AMNH 92265e in buccal view; V, AMNH 92265e in lingual view; W, AMNH 92265e inocclusal view; X, paratype AMNH 143433 in external view; Y, AMNH 143433 in interdigital view; Z, AMNH 143433 in proximal view. Abbreviations:Mtsd, metastylid; Ps, parastyle. 2 cm scale applies to the dental materials; 1 cm scale applies to the first phalanx only.

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FIGURE 3. Hispanomeryx andrewsi, sp. nov., type series. A, paratype AMNH 26499 in buccal view; B, AMNH 26499 in lingual view; C, AMNH26499 in occlusal view; D, paratype AMNH 26499a in buccal view; E, AMNH 26499a in lingual view; F, AMNH 26499a in occlusal view; G, paratypeAMNH 92265d in buccal view; H, AMNH 92265d in lingual view; I, AMNH 92265d in occlusal view; J, paratype AMNH 92265a in buccal view; K,AMNH 92265a in lingual view.

CONCLUSIONS

The Miocene musk deer Hispanomeryx was hitherto unknownfrom Asian deposits. However, the new species Hispanomeryxandrewsi from the Tunggur Formation of China constitutes thefirst evidence of an Asian Hispanomeryx, invalidating the classicview of the genus as a strictly European taxon. Also, it is the firstHispanomeryx described from outside Spain and posterior to theMN 6. The cheek teeth of H. andrewsi, characterized by the au-tapomorphic acquisition of structures that were unknown to oc-cur in the remaining Hispanomeryx forms (relatively prominentribs, styles, and stylids), were able to sustain a high level of stressduring chewing. It also shows that Hispanomeryx was more mor-phologically heterogeneous than previously thought. Finally, thepresence of H. andrewsi in the middle Miocene of China demon-

strates that the Moschidae were already present in Asia well be-fore becoming extinct in Europe by the middle Turolian.

ACKNOWLEDGMENTS

We thank Ruth O’Leary (AMNH) for cataloguing and giv-ing us access to the Tunggur moschid material curated by her.Also we are grateful to JVP referees for their helpful commentsthat enhanced the quality of the original manuscript. I.M.S. ac-knowledges a CSIC JAE-Doc contract co-funded by the FSE.D.D. acknowledges an ICP postdoctoral research contract. Thisstudy is part of the research projects CGL2008-05813-CO2-01and CGL2010-21672/BTE, MICINN (Spanish Government), andthe Research Group CAM-UCM 910607.

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Submitted February 15, 2011; revisions received July 11, 2011; acceptedAugust 18, 2011.Handling editor: Jessica Theodor.

APPENDIX 1. Modified character list of Sanchez et al. (2010)used in the phylogenetic analysis.

(1) Condition of the tympanohyal vagina: reduced enclosing,affecting only to the proximal third part of the tympanohyal(0); complete enclosing, the tympanohyal vagina is not visi-ble in lateral view (1).

(2) Width of the tympanohyal vagina: relatively narrow (0); rel-atively wide (1).

(3) Morphology of the P3: short with individualized centralcone (0); semicircular and fully selenodont (1).

(4) Morphology of the M2: square shaped (0); triangular distalprofile (1).

(5) Relative size of M1/M2: subequal in length (0); M2 muchlonger than M1 (1); M2 a bit longer than M1 (2).

(6) Morphology of the paracone rib: pyramid-shaped (0);columnar (1).

(7) Relative development of the paracone rib: well developed(0); weak (1).

(8) Relative development of the parastyle in the M2: very welldeveloped, projecting bucally (0); more or less developed(1).

(9) Relative development of the metaconid lingual rib: little de-veloped (0); very poorly developed or almost non-existent(1); hypertrophied and hook-like (2).

(10) Relative development of the metastylid: little developed(0); very weak (1); developed (2).

(11) Lower premolar/lower molar row length ratio: not reducedpremolar row (0); reduced premolar row (1).

(12) m1 size vs. m2 size: m1 subequal or slightly smaller than m1(0); m1 far more smaller than m2 (1).

(13) Relative lower molar height: relatively high (0); relativelyvery high (1).

(14) Cristid obliqua in the p4: present (0); absent (1).(15) General morphology of the lower molars: relatively shorter

and broader (0); relatively narrower and elongated (1).(16) Development of the mesial cingulid: little developed (0); hy-

pertrophied (1).(17) Morphology of the prehypocristid: straight and pointing to

the center of the teeth (0); lingually turned and fused withthe pre-entocristid (1).

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(18) Morphology of the p4: the mesolingual conid joins thecristid obliqua through the posterolingual cristid (0); as in1 with very developed anterolingual cristid that encloses to-tally or almost totally the anterior valley (1).

(19) Morphology of the third lobe of the m3: distally open (0);distally closed (1).

(20) Number of cuspids of the third lobe of the m3. monocuspi-date (0); bicuspidate, with presence of well developed post-entoconulidcristid (1).

(21) Morphology of the articular facet for the semilunate in theradius: without lateral notch (0); with lateral notch (1).

(22) Morphology of the tuber calcanei: short and broad (0);transversally compressed and dorsoplantarly elongated (1).

(23) Plantar morphology of the sustentaculum tali: short sus-tentaculum, with marked plantar notch (0); expanded sus-tentaculum, with poorly developed or non-existent plantarnotch, and almost straight plantar border (1).

(24) General morphology of the astragalus: slender and elon-gated (0); short and wide, strongly cubic in shape(1).

(25) Morphology of the lateral condyle in the distal trochleaof the astragalus: very inclined lateral border with a tri-angular and well marked proximal notch (0); vertical orslightly inclined lateral border, without proximal notch(1).

(26) Metatarsal plantar tuberosity: absent (0); present (1).

APPENDIX 2. Character-taxon matrix used in the analysis.

Taxon 1 2 20 0 6

Moschus 1 0 1 0 0 0 0 0 2 1 0 0 0 0 0 1 0 1 1 1 0 0 0 0 0 1H. duriensis ? ? 0 0 2 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 0 1 1 1 1H. aragonensis ? ? 0 ? ? ? ? ? 0 0 1 1 0 0 1 0 0 1 0 1 ? ? ? 1 1 ?H. daamsi 0 1 0 0 2 1 1 1 1 1 1 1 1 0 1 0 1 1 1 1 1 1 0 1 1 1H. andrewsi ? ? ? 1 1 1 1 0 0 2 1 1 1 0 1 0 ? 1 ? 1 ? ? ? ? ? ?Namacerus 0 1 0 0 ? 1 0 1 0 0 0 0 0 0 1 0 0 0 1 0 0 ? ? 1 1 0Micromeryx 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1

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