Israel M. Sánchez

NEW DATA ON THE MOSCHIDAE (MAMMALIA, RUMINANTIA) FROM THE UPPER MIOCENE OF SPAIN (MN 10- MN 11)

ABSTRACT New data on the last representatives of the Spanish Miocene Moschidae (Mammalia, Ruminantia) are presented and discussed. A new species of Micromeryx, Micromeryx soriae, is described on the basis of fossil material from the locality of La Roma-2 (upper Vallesian; MN 10, Teruel province, Spain, previously assigned to Micromeryx sp.), Batallones-1, and Batallones-10 (upper Vallesian; local zone J, MN 10, Madrid province, Spain). The new species is characterized by possessing relatively hypsodont lower molars of advanced morphology and a unique type of Palaeomeryx-fold. M. soriae gives insight to the last representatives of the genus, which achieved an overall lower molar morphology that mimics to certain degree that of Hispanomeryx. several characters of the upper molars of the Miocene moschids Micromeryx and Hispanomeryx are described that are useful for characterizing both genera. These characters are used to solve the taxonomic problems of PM-659, an upper molar from Puente Minero (MN 11, Teruel province, Spain) that represents one of the last survivors of the Moschidae in the Iberian Peninsula. The systematic utility of the upper molars of Hispanomeryx and Micromeryx is demonstrated, and the use of size as the only way to distinguish between the two genera is refuted. Finally, the study of the morphological characters of PM-659 clearly supports its generic change from Micromeryx sp. to Hispanomeryx sp., thus recording the presence of Hispanomeryx in the Iberian Peninsula as late as the lower Turolian.

MICROMERYX AZANZAE SP. NOV. (RUMINANTIA: MOSCHIDAE) FROM THE MIDDLE-UPPER MIOCENE OF SPAIN, AND THE FIRST DESCRIPTION OF THE CRANIUM OF MICROMERYX

Abstract A new species of moschid, Micromeryx azanzae sp. nov., is described on the basis of new fossil material from Toril-3 localities (upper Aragonian; local zone G3, MN 7/8, Zaragoza province, Spain). The fossil localities of Toril-3A and 3B have yielded exceptional material of this taxon, including the first ever described Micromeryx skull remains, dramatically improving the knowledge of Micromeryx morphology; previously unknown regions of the skeleton described here include the tympanohyal complex and most of the postcranial skeleton, both of which will be of great importance in reconstructing the sequence of character state changes within the Moschidae. Evidence for metric sexual dimorphism in M. azanzae is also discussed. Micromeryx azanzae is the best-represented species of the Moschidae in the Spanish fossil record, not only because of the quality and quantity of the fossil material but also because the species appears at nine fossil sites and three sedimentary basins that span three million years (local zones F–H, MN 6 to early MN 9).

A new genus of tragulid ruminant from the early Miocene of Kenya

We re-describe the type material of the tiny African tragulid “Dorcatherium” moruorotensis from the early Miocene of Kenya, and erect the new genus Afrotragulus, the first African Miocene tragulid that does not belong to Dorcatherium. This new taxon is characterized by its elongated and stretched lower molars with a unique morphological dental pattern that strongly contrasts with that of Dorcatherium. We additionally include the former “Dorcatherium” parvum, also a small species from the early Miocene of Kenya, into the new genus Afrotragulus as Afrotragulus parvus, figuring it for the first time. We discuss the usefulness of body size as the main taxonomical criterion for studying the Tragulidae. And finally we comment on the taxonomical and morphological diversity of tragulids in the moment of their sudden early Miocene re-apparition and expansion in the Old World.

New data on the Moschidae (Mammalia, Ruminantia) from the upper Miocene of Spain (Mn 10–Mn 11) New data on the Moschidae (Mammalia, Ruminantia) from the upper Miocene of Spain (Mn 10–Mn 11)

ABSTRACT New data on the last representatives of the Spanish Miocene Moschidae (Mammalia, Ruminantia) are presented and discussed. A new species of Micromeryx, Micromeryx soriae, is described on the basis of fossil material from the locality of La Roma-2 (upper Vallesian; MN 10, Teruel province, Spain, previously assigned to Micromeryx sp.), Batallones-1, and Batallones-10 (upper Vallesian; local zone J, MN 10, Madrid province, Spain). The new species is characterized by possessing relatively hypsodont lower molars of advanced morphology and a unique type of Palaeomeryx-fold. M. soriae gives insight to the last representatives of the genus, which achieved an overall lower molar morphology that mimics to certain degree that of Hispanomeryx. several characters of the upper molars of the Miocene moschids Micromeryx and Hispanomeryx are described that are useful for characterizing both genera. These characters are used to solve the taxonomic problems of PM-659, an upper molar from Puente Minero (MN 11, Teruel province, Spain) that represents one of the last survivors of the Moschidae in the Iberian Peninsula. The systematic utility of the upper molars of Hispanomeryx and Micromeryx is demonstrated, and the use of size as the only way to distinguish between the two genera is refuted. Finally, the study of the morphological characters of PM-659 clearly supports its generic change from Micromeryx sp. to Hispanomeryx sp., thus recording the presence of Hispanomeryx in the Iberian Peninsula as late as the lower Turolian.

New insights on the taphonomy of the exceptional mammalian fossil sites of Cerro de los Batallones (late Miocene, Spain) based on rare earth element geochemistry

Abstract Cerro de los Batallones fossil sites are distinguished by large and diverse accumulations of Miocene vertebrate fauna. Little taphonomic research has been conducted on these assemblages so far, however. Results of Rare Earth Element (REE) analyses constrain diverse aspects of the taphonomic history undergone by the bones and constitute a starting point for subsequent taphonomic studies. Cerro de los Batallones localities were formed as cavities and seem to be composed of two types of assemblages that differ in their stratigraphic position, internal stratigraphic architecture, taxonomic composition and several taphonomic features. Despite these differences, chemically analyzed bones from the Batallones-1 upper and lower level assemblages exhibit undistinguishable REE patterns both within and between them. This, together with other taphonomic features, indicates that bones are autochthonous and that the depositional context remained constant during the sedimentation of the cavity filling. In addition, REE analyses are a key tool in unveiling the provenance of those fossil bones that could be regarded as allochthonous considering their peculiar macroscopic modifications. Negative Ce anomalies exhibited by isolated fossil bones lead to the proposal that the ponds that existed in the lower level of Batallones-1 were oxic. This inference clarifies the mode of accumulation of individuals in this assemblage: the carcasses did not accumulate massively over a brief period of time but rather they concentrated and, therefore, decayed over a relatively prolonged time span.

The First Known Asian Hispanomeryx (Mammalia, Ruminantia, Moschidae)

The first known Asian Hispanomeryx (Mammalia, Ruminantia, Moschidae) Israel M. Sanchez a , Daniel Demiguel b , Victoria Quiralte a & Jorge Morales a a Departamento de Paleobiologia, Museo Nacional de Ciencias Naturales-CSIC, C/ Jose Gutierrez Abascal 2, 28006, Madrid, Spain b Institut Catala de Paleontologia, Universitat Autonoma de Barcelona, Edifici ICP, Campus de la UAB s/n, 08193, Cerdanyola del Valles, Barcelona, Spain

Systematics and evolution of the Miocene three-horned palaeomerycid ruminants (Mammalia, Cetartiodactyla)

Palaeomerycids were strange three-horned Eurasian Miocene ruminants known through fossils from Spain to China. We here study their systematics, offering the first cladistic phylogeny of the best-known species of the group, and also reassess their phylogenetic position among ruminants, which is currently disputed. The beautifully preserved remains of a new palaeomerycid from middle Miocene deposits of Spain, Xenokeryx amidalae gen. et sp. nov., helps us to better understand palaeomerycid anatomy, especially that of the nuchal region in the skull, significantly improving our current knowledge on these enigmatic ruminants. Our results show two main lineages of palaeomerycids, one containing the genus Ampelomeryx diagnosed by a characteristic type of cranium / cranial appendages and some dental derived traits, and another one that clusters those forms more closely related to Triceromeryx than to Ampelomeryx, characterized by a more derived dentition and a set of apomorphic cranial features. Xenokeryx branches as a basal offshoot of this clade. Also, we find that Eurasian palaeomerycids are not closely related to North American dromomerycids, thus rejecting the currently more accepted view of palaeomerycids as the Eurasian part of the dromomerycid lineage. Instead of this, palaeomerycids are nested with the African Miocene pecoran Propalaeoryx and with giraffoids. On the other hand, dromomerycids are closely related to cervids. We define a clade Giraffomorpha that includes palaeomerycids and giraffids, and propose an emended diagnosis of the Palaeomerycidae based on cranial and postcranial characters, including several features of the cranium not described so far. We also define the Palaeomerycidae as the least inclusive clade of pecorans containing Triceromeryx and Ampelomeryx. Finally, we reassess the taxonomy of several palaeomerycid taxa.

First African record of the Miocene Asian mouse-deer Siamotragulus (Mammalia, Ruminantia, Tragulidae): implications for the phylogeny and evolutionary history of the advanced selenodont tragulids

New remains of the small tragulid Dorcatherium songhorensis Whitworth, 1958 from the Early Miocene fossil site of Napak XXI (Uganda) include the first significant sample of postcranial bones from this species ever described. The limb bones of this tragulid are very similar to that described in the Miocene Asian long-legged tragulids of the genus Siamotragulus Thomas et al., 1990, a type previously unknown in the African Miocene. A cladistic analysis links D. songhorensis to a Siamotragulus clade as its basal offshoot, so we propose the name Siamotragulus songhorensis (Whitworth, 1958) for this species. Also, the Siamotragulus clade belongs to a monophyletic group that includes Afrotragulus Sánchez et al., 2010 and the extant Asian genera Moschiola and Tragulus. This inclusive clade is characterized by both a derived selenodont dentition and an advanced postcranial skeleton. Additionally Siamotragulus shows some cursorial refinements reflected in its postcranial skeleton including the pecoran-like metatarsals III–IV. Siamotragulus songhorensis shows that the genus Siamotragulus was not endemic to Asia as previously thought, and that a highly diverse guild of tragulids, including different members of the advanced selenodont clade, inhabited Africa as early as the Early Miocene (19–20 Ma).

Sexual Dimorphism of the Frontal Appendages of the Early Miocene African Pecoran Prolibytherium Arambourg, 1961 (Mammalia, Ruminantia)

ISRAEL M. SANCHEZ,* ! 2 VICTORIA QUIRALTE,2 JORGE MORALES,2 BEATRIZ AZANZA,3 and MARTIN PICKFORD4; department of Zoology, Museum of Zoology, Cambridge University, Downing Street, Cambridge, CB2 3EJ, United Kingdom, ims32@cam.ac.uk; israelms@mncn.csic.es; 2Departamento de Paleobiologia, Museo Nacional de Ciencias Naturales-CSIC, C/ Jose Gutierrez Abascal, 2, 28006 Madrid, Spain; 3Departamento de Ciencias de la Tierra (Paleontologia), Facultad de Ciencias, Universidad de Zaragoza, 50009, Zaragoza, Spain; Departement Histoire de la Terre, UMR 7202 (CR2P) du CNRS, case postale 38, 8, rue Buffon, 75005 Paris, France

Comparative anatomy, phylogeny, and systematics of the Miocene giraffid Decennatherium pachecoi Crusafont, 1952 (Mammalia, Ruminantia, Pecora): State of the art

ABSTRACT Decennatherium pachecoi Crusafont, 1952, is one of two giraffid species described from the Miocene of the Iberian Peninsula. This species is recovered exclusively from Vallesian faunas (MN9-10, late Miocene, 10–11 Ma). Despite being relatively well represented in the fossil record, except for the skull and ossicones, the complete vertebral column, and part of the upper dentition, its systematics and phylogenetic position among giraffids are the subject of debate. We update our knowledge of D. pachecoi, revising all Spanish material assigned to this species, as well as previously undescribed fossils. We reassess the systematics of Decennatherium, including its potential relationship with the second Iberian giraffid, the early Turolian Birgerbohlinia Crusafont, 1952, by means of the first cladistic analysis of the Giraffidae that includes Decennatherium together with the most relevant African and Eurasian taxa, both fossil and extant. Our results link Decennatherium with a ‘samothere’ clade, whereas Birgerbohlinia is nested within a ‘sivathere’ clade, thus refuting a previously assumed direct relationship between the two Spanish forms. Finally, we discuss some other possible finds of the genus Decennatherium in Greece, Turkey, and Iran.