The genus Hispanomeryx (Mammalia, Ruminantia, Moschidae) and its bearing on musk deer phylogeny and systematics
2010; Wiley; Volume: 53; Issue: 5 Linguagem: Inglês
10.1111/j.1475-4983.2010.00992.x
ISSN1475-4983
AutoresIsrael M. Sánchez, M. Soledad Domingo, Jorge Morales,
Tópico(s)Bat Biology and Ecology Studies
ResumoPalaeontologyVolume 53, Issue 5 p. 1023-1047 Free Access The genus Hispanomeryx (Mammalia, Ruminantia, Moschidae) and its bearing on musk deer phylogeny and systematics ISRAEL M. SÁNCHEZ, ISRAEL M. SÁNCHEZ Department of Zoology, Museum of Zoology, University of Cambridge, Downing Street, Cambridge, CB2 3EJ, UK; e-mail israelms@mncn.csic.es Museo Nacional de Ciencias Naturales-CSIC, C/José Gutiérrez Abascal, 2. 28006 Madrid, Spain; e-mails mcnsd748@mncn.csic.es; mcnm166@mncn.csic.esSearch for more papers by this authorM. SOLEDAD DOMINGO, M. SOLEDAD DOMINGO Museo Nacional de Ciencias Naturales-CSIC, C/José Gutiérrez Abascal, 2. 28006 Madrid, Spain; e-mails mcnsd748@mncn.csic.es; mcnm166@mncn.csic.esSearch for more papers by this authorJORGE MORALES, JORGE MORALES Museo Nacional de Ciencias Naturales-CSIC, C/José Gutiérrez Abascal, 2. 28006 Madrid, Spain; e-mails mcnsd748@mncn.csic.es; mcnm166@mncn.csic.esSearch for more papers by this author ISRAEL M. SÁNCHEZ, ISRAEL M. SÁNCHEZ Department of Zoology, Museum of Zoology, University of Cambridge, Downing Street, Cambridge, CB2 3EJ, UK; e-mail israelms@mncn.csic.es Museo Nacional de Ciencias Naturales-CSIC, C/José Gutiérrez Abascal, 2. 28006 Madrid, Spain; e-mails mcnsd748@mncn.csic.es; mcnm166@mncn.csic.esSearch for more papers by this authorM. SOLEDAD DOMINGO, M. SOLEDAD DOMINGO Museo Nacional de Ciencias Naturales-CSIC, C/José Gutiérrez Abascal, 2. 28006 Madrid, Spain; e-mails mcnsd748@mncn.csic.es; mcnm166@mncn.csic.esSearch for more papers by this authorJORGE MORALES, JORGE MORALES Museo Nacional de Ciencias Naturales-CSIC, C/José Gutiérrez Abascal, 2. 28006 Madrid, Spain; e-mails mcnsd748@mncn.csic.es; mcnm166@mncn.csic.esSearch for more papers by this author First published: 16 September 2010 https://doi.org/10.1111/j.1475-4983.2010.00992.xCitations: 31AboutSectionsPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinked InRedditWechat Abstract Abstract: We update the systematics and comparative anatomy of the genus Hispanomeryx Morales, Moyà-Solà and Soria, 1981 through the description of a new and abundant fossil material from the middle Miocene localities of Toril-3, Manchones-1 and Manchones-2, Zaragoza Province, Spain. Hispanomeryx was only known by dental remains, mainly mandibles and lower teeth, and very scarce postcranial material; the fossil sample studied here includes cranial, mandibular, dental and postcranial remains, and it allows us to describe in depth, for the first time, the anatomy of the genus. We also erect the new species Hispanomeryx daamsi. The material is good and abundant enough as to include Hispanomeryx in a cladistic analysis performed to explore its phylogenetic relationships within the Pecora. This analysis includes Hispanomeryx in a monophyletic Moschidae (musk deer) composed by Micromeryx, Hispanomeryx, 'Moschus' grandeavus and extant Moschus, and recovers a clade composed by moschids and bovids; this is the first time that a Moschidae-Bovidae sister-group relationship is backed-up by morphological data. A direct sister-group relationship between Hispanomeryx and the Bovidae is thus rejected. Several taxa previously assigned to the 'Moschidae' are rejected as true moschids. Finally, the cladistic phylogenetic analysis of Hispanomeryx demonstrates its monophyly and shows a basal species, H. aragonensis Azanza, 1986, and a clade formed by H. duriensis Morales et al., 1981 and H. daamsi sp. nov, characterized by the presence of more derived lower molars than those of H. aragonensis. Hispanomeryx Morales, Moyà-Solà and Soria, 1981 is a small pecoran ruminant from the middle–late Miocene of Eurasia. Its remains come basically from Spain (Morales et al. 1981, 1992, 2000, 2004; Azanza 1986; Moyà-Solà 1986; Azanza et al. 1999, 2004; Álvarez Sierra et al. 2003; Peláez-Campomanes et al. 2003; Sánchez and Morales 2004, 2006; Sánchez 2006; Prothero 2007; Sánchez et al. 2009), although it has also been identified in Turkey and North Caucasus (Moyà-Solà 1986; Gentry 1990, 1994; Pickford et al. 1999). Hispanomeryx duriensis Morales et al., 1981 (type species, ranging MN 9-MN 10) and H. aragonensis Azanza, 1986 (MN 7/8) are the only two described species in the genus. Both taxa were diagnosed on the basis of their lower dentition. Very few postcranial elements of H. duriensis have been described (see Morales et al. 1981), and the very scarce postcranial remains of H. aragonensis were totally unknown until recently (Sánchez 2006). Similarly, some forms such as the Turolian Hispanomeryx sp. from Puente Minero (MN 11; Sánchez et al. 2009), or those aforementioned remains from outside the Iberian Peninsula, consist solely on dental material. Thus, our knowledge on the general anatomy of Hispanomeryx was severely restricted so far. First classified as a moschid (musk deer), with close affinities to extant Moschus spp. Linnaeus, 1758 (Morales et al. 1981; Azanza 1986), the relatively hypsodont bovid-like lower molars of Hispanomeryx together with its distally open metatarsal sulcus, made the systematics and phylogenetic position of the genus highly controversial. The status of Hispanomeryx ranged from moschid to giraffoid, bovoid or bovid, and even a family Hispanomerycidae was proposed to accommodate the genus (see Morales et al. 1981; Moyà-Solà 1986, 1987; Janis and Scott 1987; Gentry 1990, 2000; Morales et al. 1995; McKenna and Bell 1997; Gentry et al. 1999; Hassanin and Douzery 2003; Prothero 2007; Prothero and Foss 2007; Solounias 2007; Vislobokova and Lavrov 2009). This instability reflects the difficulties to assess the character state distribution of Hispanomeryx in the context of pecoran evolution, mainly due to the scarcity of published fossil remains, but also due to its apparently contradictory morphological traits as perceived in the classical notion of moschids as morphologically heterogeneous cervoid pecorans. Given the relatively poor known fossil record and the few works published on the subject, Hispanomeryx has remained largely unknown, being almost unnoticed, if compared with other taxa, in the majority of works regarding the Pecora (e.g. Janis and Scott 1987; Gentry et al. 1999; Prothero 2007; Solounias 2007; Prothero and Foss 2007). Even so, its high potential relevance for the understanding of the phylogenetic relationships of moschids was fully recognized (Hassanin and Douzery 2003). The fossil record of Hispanomeryx has been greatly improved after the excavation and study of the fossil sites of Toril-3 (MN 7/8, local zone G3, Zaragoza Province, Spain). These localities have yielded an abundant collection of vertebrate fossils that comprises lissamphibians, chelonians, squamates, aves and mammals (Azanza et al. 1999, 2004; Álvarez Sierra et al. 2003; Sánchez and Morales 2008). Among mammals, fossils of Hispanomeryx and Micromeryx are unusually abundant, and include cranial, dental and postcranial remains. This richness makes possible a deep comparative study of the anatomy of Hispanomeryx at an extent not reached before. This paper encompasses the first in-depth comparative description of the anatomy of Hispanomeryx on the basis of the Toril-3 remains, the description of H. daamsi sp. nov., and the systematic revision of the genus, including the phylogenetic relationships of its members. Also, an additional cladistic analysis is performed that allow us, in the first place, to resolve the affinities of Hispanomeryx and assess its key importance for the diagnosis and definition of the Moschidae, and in second place, to explore the phylogenetic relationships of moschids within the Pecora. This fact is of the outmost importance to understand the definition and diagnosis of the Bovoidea. Locality and geological setting The fossil sites of Toril-3 (Toril-3A and 3B; upper Aragonian, local zone G3, MN 7/8) are located to the northeast of the town of Daroca, Zaragoza Province, Spain, and belong to the Villafeliche-Daroca sector of the Calatayud-Montalbán Basin (Text-fig. 1). The sites of Toril-3 are situated in the most basal part of a succession that contains the transition from the upper Aragonian to the Vallesian (Azanza et al. 1999, 2004; Daams et al. 1999; Alcaláet al. 2000; Álvarez Sierra et al. 2003). Both localities belong to the same horizon, and the distinction between them (–A and –B) only reflects the fact that they are separated by less of 60 m of farmland. Toril-3 is the type locality of the bovid Samotragus pilgrimi Azanza, Nieto and Morales, 1998 and the moschid Micromeryx azanzae Sánchez and Morales, 2008. Figure TEXT-FIG. 1. Open in figure viewerPowerPoint Map of the tertiary Spanish continental basins, with remarks on the situation of the Toril-3 fossil sites near the town of Daroca. CMB, Calatayud-Montalbán Basin; DB, Duero Basin; EB, Ebro Basin; MB, Madrid Basin. Materials and methods Material The new species of Hispanomeryx described in this work is based upon a selection of fossils from Toril-3A and Toril-3B. The complete sample was not used because the fossil site of Toril-3A is still being excavated and providing new moschid remains, which are not prepared for study. This material represents the most complete and abundant set of Hispanomeryx fossils yet described. Postcranial data of H. aragonensis come from still unpublished Spanish fossil material (Sánchez 2006). Moschus data come from specimens curated by the AMNH (New York, USA), the Museum of Zoology of the University of Cambridge (Cambridge, UK), and the Museo Anatómico de la Universidad de Valladolid (Valladolid, Spain). Data of Blastomeryx'elegans'Matthew and Cook, 1909 (Blastomeryx gemmifer sensu Prothero, 2007) come from the Agate and Trinity River collections curated by the AMNH (New York, USA). Data of basal Cervidae come from Spanish unpublished material (IMS, pers. obs.) and published data on Procervulus Gaudry, 1877 (Rössner 1995); also, anatomical data have been recovered from osteological material of extant Muntiacus Rafinesque, 1815 curated by the Museo Anatómico de la Universidad de Valladolid (Valladolid, Spain). Data regarding giraffids and tragulids come from the collections of comparative anatomy of the MNCN-CSIC (Madrid, Spain) and the Museum of Zoology of the University of Cambridge (Cambridge, UK). Data of Dremotherium Geoffroy Saint-Hilaire, 1833 come from the type locality of Saint-Gérand-le-Puy, France (Sigogneau 1968) and from the fossil sample of Cetina de Aragón, Spain (Ginsburg et al. 1994; IMS, pers. obs.). Data of Amphitragulus Pomel, 1853 come from the type locality of Saint-Gérand-le-Puy, France and Quercy (France). Data regarding Orangemeryx Morales, Soria and Pickford, 1999, Namibiomeryx Morales, Soria and Pickford, 1995, Namacerus Morales, Soria, Pickford and Nieto, 2003, and Sperrgebietomeryx Morales, Soria and Pickford, 1999 come from the direct study of the original material from the Sperrgebiet (Namibia; see Morales et al. 1995, 1999, 2003a, 2003b, 2008). The data on Moschus grandeavus Schlosser, 1924 have been extracted from the descriptions and figured material depicted in Vislobokova and Lavrov (2009). Finally, several morphological data come from Janis and Scott (1987) and Webb and Taylor (1980). Methods Measurements. All measurements (detailed in Supporting Information, Tables S1–S3) were taken with digital calipers. We follow the set of measurements proposed by Sánchez and Morales (2008). Nomenclature. We use the terminology of Barone (1999) for anatomic nomenclature of the cranial and postcranial skeleton, and that of Azanza (2000) for nomenclature of the dentition. Phylogenetic analyses. The data matrices were compiled in MacClade 4.05 and run in PAUP 4.0b10 (Macintosh versions). Also, we used MacClade 4.05 to produce the scaffold constraints. Institutional and anatomical abbreviations. AMNH, American Museum of Natural History, New York, USA; M-1, Manchones-1; M-2, Manchones-2; MC III-IV, metacarpal III-IV; MNCN-CSIC, Museo Nacional de Ciencias Naturales-CSIC, Madrid, Spain; MPZ, Museo Paleontológico de la Universidad de Zaragoza, Zaragoza, Spain; MT III-IV, metatarsal III-IV; TOR3A, Toril-3A; TOR3B, Toril-3B. Systematic paleontology MAMMALIA Linnaeus, 1758CETARTIODACTYLA Montgelard, Catzeflis and Douzery, 1997RUMINANTIA Scopoli, 1777PECORA sensu Webb and Taylor, 1980BOVOIDEA Gray, 1821 Emended diagnosis. Pecoran ruminants characterized by well developed postentocristid that fuses with the posthypocristid, distally closing the lower molars; expanded postprotocrista; subcentral tympanohyal vagina; reduced lateral enclosing of the tympanohyal vagina; laterally inflated tympanic bulla, with developed caudal expansion; postglenoid process with medial expansion; absence of lacrimal fossa; shortened P2; aligned cusps in the lower molars; non-stepped lateral border of the dorso-distal process of the tibia; and distally open metatarsal sulcus. MOSCHIDAE Gray, 1821 Emended diagnosis. Bovoid ruminants characterized by a foramen ovale separated from the infratemporal fossa by a low and small crest; highly derived p4 with a mesolingual conid that joins with the cristid obliqua through the posterolingual cristid and projects a very developed anterolingual cristid that closes, totally or almost totally, the anterior valley; m3 with a bi-cuspidate third lobe displaying a well developed post-entoconulidcristid; presence of posterior metatarsal tuberosity; and long and wide capitular facet in the radius. HISPANOMERYX Morales, Moyà-Solà and Soria, 1981 1968 Gazella sp. Crusafont et al. Type species. Hispanomeryx duriensis Morales et al., 1981. Emended diagnosis. Moschid of relative medium to large size characterized by a reduced lower premolar tooth row if compared to Micromeryx and Moschus; upper molars with flat and aligned external cusps, straight and column-like buccal structures; lower molars with highly aligned cusps and very reduced lingual structures; very reduced, almost non-existent metastylid; very weak cingulids; absence of Palaeomeryx-fold; m1 much smaller than m2; processus condilaris lacking lateral ramus; wide subcentral tympanohyal vagina, with small and proximally located lamina vaginalis; laterally enclosed temporal canal; contact between the postglenoid process and the external acoustic meatus; presence of a lateral notch in the semilunate facet of the radius; short and compact astragalus; rectangular external proximal facet in the first phalanx; absence of post-articular plateau in the second phalanx; short and high third phalanx, with developed extensor process. Other referred species. Hispanomeryx aragonensis Azanza, 1986, Hispanomeryx daamsi sp. nov. Hispanomeryx duriensis Morales, Moyà-Solà and Soria, 1981 Emended diagnosis. Large sized Hispanomeryx with absence of cristid obliqua in the p4. The lower molars are high-crowned, with very poorly developed lingual structures. Ocurrence. The type locality of H. duriensis is El Lugarejo (Duero Basin, local zone H, MN 9, Vallesian, Ávila Province, Spain; Text-fig. 2). The species has also been found in the locality of Los Valles de Fuentidueña (Duero Basin, local zone H, MN 9, Vallesian, Segovia Province, Spain), and a form assigned to Hispanomeryx sp. cf. H. duriensis has been found in the sites of Cerro de los Batallones (Madrid Basin, local zone J, MN 10, Vallesian, Madrid Province, Spain; Morales et al., 1981; Sánchez and Morales 2006). Figure TEXT-FIG. 2. Open in figure viewerPowerPoint Chronostratigraphic position of the fossil localities where Hispanomeryx daamsi, H. duriensis, H. aragonensis and Hispanomeryx sp. are found. The star symbol represents the type locality and the circle symbol marks the remaining sites where a given species is found. Black colour represents H. daamsi; white colour, H. aragonensis; light grey colour, H. duriensis; and dark grey colour, Hispanomeryx sp. The cf. forms are not distinguished from the nominal species in this figure. List of sites: BAT, Batallones; EL, El lugarejo; ES, Escobosa de Calatañazor; M1, M2, Manchones-1 and -2; LA, Los Andurriales; LC-1B, La Ciesma-1B; LVF, Los Valles de Fuentidueña; PA3, Paracuellos-3; PM, Puente Minero; SO, Solera; TOR3, Toril-3. Hispanomeryx aragonensis Azanza, 1986 Emended diagnosis. Relatively small Hispanomeryx, it is a primitive species characterized by the distally open third lobe in the m3. Ocurrence. The type locality of H. aragonensis is La Ciesma-1B (Ebro Basin, local zone G3, MN 7/8, upper Aragonian, Zaragoza Province, Spain; Text-fig. 2). The species probably also appear in the sites of Escobosa de Calatañazor (Duero Basin, local zone G3, MN 7/8, upper Aragonian, Soria Province, Spain), Los Andurriales (Calatayud-Montalbán Basin, local zone G3, MN 7/8, upper Aragonian, Zaragoza Province, Spain) and Solera (Calatayud-Montalbán Basin, local zone G3, MN 7/8, upper Aragonian, Zaragoza Province, Spain), although this material is still non-described. A few non-described fossils assigned preliminary to Hispanomeryx sp. cf. H. aragonensis (see Sánchez and Morales 2006) have been found in Paracuellos-3 (Madrid Basin, local zone G1, MN 6, upper Aragonian, Madrid Province, Spain). Hispanomeryx daamsi sp. nov. TEXT-FIG. 3. - TEXT-FIG. 9. Figure TEXT-FIG. 3. Open in figure viewerPowerPoint Hispanomeryx daamsi Sánchez, Domingo and Morales sp. nov. Selected mandibular remains from Toril-3 and Manchones-1. A–C, holotype, MNCN-33300, right hemimandible with p4-m3. A, buccal, B, lingual, and C, occlusal views. D–E, paratype, MPZ 2008/177, complete right hemimandible with p3-m3. D, buccal, and E, occlusal views. F, close detail of m2-m3 from the holotype. G, paratype, MNCN-33301, complete right hemimandible with p2 (fragmented)-m3 in lingual view. H, paratype, TOR3B-28, right hemimandibular fragment with p2-p3 showing the diastema area in buccal view. I–J, paratype, MPZ 2008/179, right hemimandible with p4-m3. I, buccal, and J, lingual views. K, paratype, MNCN-33305, right juvenile hemimandible with dp2-m1 in buccal view. L, MA1-5070, right juvenile hemimandible with dp2-dp4 and emerging m1 in buccal view; this specimen corresponds to an extremely young individual. M–N, MA1-5069, left hemimandible with p3-m3. M, occlusal, and N, lingual views. O, MA1-5068, right hemimandible with p3-m3 in buccal view. PrHyp, pre-hypocristid. Figure TEXT-FIG. 4. Open in figure viewerPowerPoint Hispanomeryx daamsi Sánchez, Domingo and Morales sp. nov. Cranial remains from Toril-3. A–C, paratype, MNCN-5001, fragmented skull. A, lateral (left), B, ventral, and C, dorsal views. D–E, paratype, MPZ 2008/172, fragmented skull with right upper molar row. D, ventral, and E, dorsal views. FrB, frontal bone; FrNs, fronto-nasal suture; MaxB, maxillar bone; OccC, occipital condyles; Or, orbit; PalB, palatine bone; ParB, parietal bone; SgC, sagittal crest (cast on the matrix); TmpB, temporal bone; TyBll, tympanic bulla. Figure TEXT-FIG. 5. Open in figure viewerPowerPoint Hispanomeryx daamsi Sánchez, Domingo and Morales sp. nov. Detail of the left bulla of MNCN-5001 in ventro-lateral view. ExAdM, external auditory meatus; OccB, occipital bone; PGp, postglenoid process; TyhVg, tympanohyal vagina; TyBll, tympanic bulla; LmVg, lamina vaginalis. Figure TEXT-FIG. 6. Open in figure viewerPowerPoint Hispanomeryx daamsi Sánchez, Domingo and Morales sp. nov. Selected maxillary remains with upper cheek teeth from Toril-3 and Manchones-1. A–B, paratype, MPZ 2008/173, complete maxillary roof with P2-M3 (both rows). A, ventral, and B, lateral (left maxillar) views. C, paratype, MPZ 2008/175, left juvenile maxillar fragment with dP2-M1 in linguo-occlusal view. D, M1–M2 detail of the right molar row of MPZ 2008/173, with remarks on some of the discussed structures. E, paratype, MPZ 2008/194, right juvenile maxillar fragment with dP2-M1 in linguo-occlusal view. F, MA1-5073, right maxillar fragment with P4–M3 in linguo-occlusal view. MesSt, mesostyle; Mtc-F, metaconule-fold; ParSt, parastyle; PPrt, post-protocrista. Figure TEXT-FIG. 7. Open in figure viewerPowerPoint Hispanomeryx daamsi Sánchez, Domingo and Morales sp. nov. Selected fore limb remains from Toril-3. A, paratype, MPZ 2008/202, distal fragment of humerus in dorsal view. B, paratype, MNCN-33353c, distal fragment of humerus in dorsal view. C, paratype, TOR3A-68CL, distal fragment of humerus in dorsal view. D–E, paratype, MPZ 2008/204, right radius. D, dorsal, and E, palmar views. F, paratype, MPZ 2008/209, right semilunate in proximal view. G, paratype, MPZ 2008/210, left semilunate in proximal view. H–I, paratype, MPZ 2008/207, left scaphoid. H, lateral, and I, medial views. J–K, paratype, MPZ 2008/212, right magnotrapezoid. J, proximal, and K, distal views. L–M, paratype, MPZ 2008/211, left magnotrapezoid. L, proximal, and M, distal views. N, paratype, MPZ 2008/222, right metacarpal III–IV in proximal view. O–P, paratype, MPZ 2008/220, right juvenile metacarpal III–IV. O, dorsal, and P, plantar views. Q–R, paratype, MPZ 2008/221, left juvenile MC III–IV. Q, dorsal, and R, plantar views. 1 cm scale applies to F–M (carpals), 2 cm scale applies to the rest. Figure TEXT-FIG. 8. Open in figure viewerPowerPoint Hispanomeryx daamsi Sánchez, Domingo and Morales sp. nov. Selected hind limb remains and phalanges from Toril-3. A–B, paratype, MPZ 2008/217, left tibia. A, dorsal, and B, plantar views. C–D, paratype, MPZ 2008/216, right calcaneum. C, medial, and D, plantar views. E–F, paratype, MPZ 2008/215, right astragalus. E, dorsal, and F, plantar views. G–H, paratype, MPZ 2008/214, left astragalus. G, dorsal, and H, plantar views. I–J, paratype, MNCN-33350a, left navicular-cuboid. I, proximal, and J, distal views. K–L, paratype, MPZ 2008/213, left navicular-cuboid. K, proximal, and L, distal views. M, paratype TOR3B-33330, juvenile MT III–IV, dorsal view. N, paratype, TOR3B-33331, MT III–IV, dorsal view. O, paratype, TOR3B-33334, MT III–IV, proximal view. P–Q, paratype, MNCN-33358d, first phalanx. P, external, and Q, interdigital views. R–S, paratype, MPZ 2008/224, first phalanx. R, external, and S, interdigital views. T–U, paratype, MPZ 2008/232, second phalanx. T, interdigital, and U, external views. V–W, paratype, MPZ 2008/233, second phalanx. V, interdigital, and W, external views. X–Y, paratype, MNCN-33357d, third phalanx. X, interdigital, and Y, external views. Z–AA, paratype, MPZ 2008/245, third phalanx. Z, interdigital, and AA, external views. 1 cm scale applies to P–AA (phalanges), 2 cm scale applies to the rest. Figure TEXT-FIG. 9. Open in figure viewerPowerPoint Hispanomeryx daamsi Sánchez, Domingo and Morales sp. nov. Selected postcranial remains from Manchones-1 and -2. A, MA1-287, right navicular-cuboid in proximal view. B, MA2-61, distal fragment of metatarsal III–IV in dorsal view. C, MA1-248, second phalanx in interdigital view. D, MA1-5224, third phalanx in interdigital view. 1999 Hispanomeryx aragonensis Azanza et al. 2003 Hispanomeryx aragonensisÁlvarez-Sierra et al., pl. 2. 2004 Hispanomeryx aragonensis Azanza et al., pl. 2. 2006 Hispanomeryx sp. nov. 1 Sánchez and Morales, fig. 6, pls 1–2. Derivation of name. Dedicated to the memory of Dr. Remmert Daams. Holotype. MNCN-33300, right hemimandible with p4-m3 (Text-fig. 3A–C). Paratypes. The remaining referred material from Toril-3. Material. Skull: MNCN-5001, MPZ 2008/172; maxillary bone with teeth: MPZ 2008/173 (complete roof with the two maxillae), MPZ 2008/174, MPZ 2008/175, MPZ 2008/176, MNCN-33314, MA1-5073; mandibles and mandibular fragments: MPZ 2008/177, MPZ 2008/178, MPZ 2008/179, MPZ 2008/180, MPZ 2008/181, MPZ 2008/182, MPZ 2008/183, MPZ 2008/184, MPZ 2008/185, MPZ 2008/186, MPZ 2008/187, MPZ 2008/188, MPZ 2008/189, MPZ 2008/190, MPZ 2008/191, MPZ 2008/192, MPZ 2008/197, MNCN-33306, MNCN-33304, MNCN-33301, MNCN-33305, MNCN-33302, TOR3B-28, MA1-5068, MA1-5069, MA1-5070, MA1-5071, MA1-5072; dP2: MNCN-33309; P2: MNCN-33324d; P4: MNCN-33324c, MNCN-33324b; M1: MPZ 2008/195, MNCN-33324a; M2: MPZ 2008/196; m1: MPZ 2008/198, MPZ 2008/199; MPZ 2008/200; MPZ 2008/201, TOR3B-99; m2: TOR3A-4, MNCN-33313b; m3: TOR3A-3, MNCN-33313a; humerus: MPZ 2008/202, TOR3A-68CL, MNCN-33353c, MNCN-33352a; radius: MPZ 2008/203, MPZ 2008/204, MPZ 2008/205, MPZ 2008/206, MNCN-33378; scaphoid: MPZ 2008/207; semilunate: MPZ 2008/208, MPZ 2008/209, MPZ 2008/210; magnotrapezoid: MPZ 2008/211, MPZ 2008/212; navicular-cuboid: MPZ 2008/213, MNCN-33350, MA1-287; astragalus: MPZ 2008/215, MPZ 2008/216, MA1-233; calcaneum: MPZ 2008/216; tibia: MPZ 2008/217, MPZ 2008/218, MPZ 2008/219, MNCN-33370a, MNCN-33370b, MNCN-33370d; metacarpal III-IV: MPZ 2008/220, MPZ 2008/221, MPZ 2008/222, MA1-200; metatarsal III-IV: MPZ 2008/223, MNCN-33331, MNCN-33330, TOR3B-44, MNCN-33334, TOR3B-42, MNCN-33332, MA1-201, MA2-61; first phalanx: MPZ 2008/224, MPZ 2008/225, MPZ 2008/226, MPZ 2008/227, MPZ 2008/228, MPZ 2008/229, MPZ 2008/230, MPZ 2008/231, TOR3A-1000CL, TOR3A-1002CL, MNCN-33358a, MNCN-33358b, MNCN-33358c, MNCN-33358d; second phalanx: MPZ 2008/232, MPZ 2008/233, MPZ 2008/234, MPZ 2008/235, MPZ 2008/236, MPZ 2008/237, MPZ 2008/238, MPZ 2008/239, MPZ 2008/240, MPZ 2008/241, MPZ 2008/242, MNCN-33364b, MNCN-33364c, MNCN-33364d, MNCN-33364e, MNCN-33364f, MNCN-33364j, MNCN-33364i, MA1-248; third phalanx: MPZ 2008/243, MPZ 2008/244, MPZ 2008/245, MPZ 2008/246, MPZ 2008/247, MPZ 2008/248, MPZ 2008/249, MPZ 2008/250, MPZ 2008/251, MPZ 2008/252, MPZ 2008/253, MPZ 2008/254, MPZ 2008/255, MNCN-3356b, MNCN-3356d, MNCN-3357a, MNCN-3357b, MNCN-3357c, MNCN-3357d, MNCN-3357e, MA1-5224. This list of material includes both the type series (Toril-3) and the fossils from Manchones-1 and -2. The specimens MPZ (Toril-3) are curated by the Museo Paleontológico de la Universidad de Zaragoza-Gobierno de Aragón (Zaragoza, Spain), and the specimens TOR3 (Toril-3) and MNCN (Toril-3) are curated by the MNCN-CSIC (Madrid, Spain). The specimens MA1 and MA2 (Manchones-1 and -2) are also curated by the MNCN-CSIC (Madrid, Spain). Diagnosis. Medium-sized Hispanomeryx characterized by a lingually-turned prehypocristid which fuses with the pre-entocristid, enlarging the center valley. Differential diagnosis. H. daamsi differs from H. duriensis in the presence of cristid obliqua in the p4, and in having a narrow and elongated tuber calcanei and a smaller body size. Differs from H. aragonensis in having larger body size, lower molars with more reduced lingual stylids and ribs, and distally closed third lobe of the m3. Differs from both species in the presence of lingually-turned pre-hypocristid. Ocurrence. The type locality of Hispanomeryx daamsi is Toril-3 (Calatayud-Montalbán Basin, local zone G3, MN 7/8, upper Aragonian, located near the village of Daroca, Zaragoza province, Spain; Text-fig. 2). The species has also been found in the fossil sites of Manchones-1 and -2 (Calatayud-Montalbán Basin, local zone G2, MN 6, upper Aragonian). Stratigraphical range. Hispanomeryx daamsi ranges through c. 0.6 myr, from the local zone G2 to the local zone G3 (late MN 6 to early MN 7/8) (Sánchez and Morales 2006). Description Remarks. We describe the Hispanomeryx fossils from Toril-3 and Manchones-1 and -2, emphasizing the anatomical comparisons with extant Moschus, the other two species of Hispanomeryx and, when needed, basal bovids; although we point out the comparative anatomy of Hispanomeryx and Micromeryx several times, a detailed comparative study of their anatomical traits is out of the scope of this work due to its extension and the pivotal importance of this subject for the study of these two taxa, so it will be presented in another paper (Sánchez and Morales in manuscript). The dental and postcranial fossils from Manchones are morphologically identical to their Toril-3 counterparts, so they are all described as a whole. Skull. This is the first in-depth description of the cranium of Hispanomeryx (Text-figs 4–5), although Moyà-Solà (1986) described some cranial features of the genus on the basis of two skulls that he did not figure (see Moyà-Solà, 1986, p. 269). Our two available skulls from Toril-3 (MNCN-5001 and MPZ 2008/172) have preservation problems due to diagenetic damage. MNCN-5001 retains the frontals and the proximal part of the nasals, as well as the braincase area including the ear region, but lacks the rostrum and the lacrimal and zygomatic bones; MPZ 2008/172 preserves the braincase, orbits and part of the right maxilla (including P4-M3) with remains of the palatal roof, although the specimen is heavily dorso-ventrally compressed. We describe MNCN-5001, adding data from MPZ 2008/172 if necessary, since MNCN-5001 preserves more elements and far more three-dimensional integrity. The dorsal profile of the skull describes a marked supraorbital convexity, similar to that of Moschus. The dorsal border of the orbit is preserved from the frontolacrimal suture to the constriction area over the frontoyugal suture, which shows a marked triangular morphology as in the extant musk deer. The orb
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