PALEVO 1631-0683 Elsevier S1631-0683(12)00050-4 10.1016/j.crpv.2012.01.004 Research article General palaeontology, systematics and evolution (Vertebrate palaeontology) Paléontologie générale, systématique et évolution / General palaeontology, systematics and evoluion (Paléontologie des vertébrés / Vertebrate palaeontology) The southernmost Miocene occurrence of the last European herpetotheriid Amphiperatherium frequens (Metatheria, Mammalia) Découverte la plus méridionale du dernier herpétothériidé européen Amphiperatherium frequens (Metatheria, Mammalia) au Miocène Furió Marc marc.furio@icp.cat a Ruiz-Sánchez Francisco J. b Crespo Vicente D. b Freudenthal Matthijs c d Montoya Plinio b Institut Català de Paleontologia M. Crusafont, Edifici ICP, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, 08193 Barcelona, Spain Departament de Geologia, Universitat de València, Valencia, Spain Departamento de Estratigrafía y Paleontología, Universidad de Granada, Granada, Spain Netherlands Centre for Biodiversity, Naturalis, Leiden, The Netherlands 11 5 S1631-0683(12)X0005-8 371 377 © 2012 Académie des sciences. Published by Elsevier B.V. All rights reserved. 2012 Académie des sciences Full (PDF)

The present work provides for the first time a detailed description of teeth attributable to metatherians in the Miocene fossil record of Spain, and justifies their generic and specific ascription. The fossil elements found correspond to Amphiperatherium frequens, the last herpetotheriid that inhabited Europe. This is so far the southernmost occurrence of this species, thus showing that its geographic range extended further southward than previously thought.

Ce travail donne la première description détaillée d’un métathérien trouvé dans des sédiments miocènes de l’Espagne et déterminé au niveau de l’espèce. Les fossiles trouvés appartiennent à l’espèce Amphiperatherium frequens, le dernier herpétothériidé ayant peuplé l’Europe. Il s’agit de la découverte la plus méridionale pour les métathériens, étendant ainsi considérablement leur aire géographique, une avancée nouvelle dans l’étude des faunes du Miocène européen.

 Herpetotheriidae, Europe, Neogene, Aragonian, Ribesalbes-Alcora, Spain Herpetotheriidae, Europe, Neogène, Aragonien, Ribesalbes-Alcora, Espagne presented Presented by Philippe Taquet Introduction

Herpetotheriids have recently been found to be stem metatherians and the sister group of all living marsupials (Horovitz et al., 2009 and Sánchez-Villagra et al., 2007). These metatherian mammals were rather common in central Europe during the Paleogene. They reached their maximum diversity during the Early Eocene, as demonstrated by the nine different species found in the sites of Avenay and Condé-en-Brie (Crochet, 1980). From then on, the group decreased in both relative abundance and number of species within mammal assemblages. The Oligocene represented a period of regular decline for the herpetotheriids on the European continent, beginning with six evolutionary lines and declining to only two (Crochet, 1980). The last known occurrences of Amphiperatherium frequens in the Middle Miocene from France, Germany and Switzerland (Fig. 1, Table 1) may represent the extinction of this group in central Europe before 13.8 ma, according to the dates of Kälin and Kempf (2009) and Abdul Aziz et al. (2010).

The Eurasian evolutionary history of the group outside central Europe is obscure. The fossil record of Cenozoic metatherians is particularly poor in Asia (Crochet et al., 2007 and Ni et al., 2007), especially in Neogene sediments, which have provided merely three dental elements: two from Thailand (Siamoperadectes minutus, Ducrocq et al., 1992 and Mein and Gisnsburg, 1997) and one from China (S. clandestinus, Storch and Qiu, 2002). All of them have been found in localities equivalent in age to MN4 (Early Miocene).

The Cenozoic fossil record of metatherians in southwestern Europe is similarly scarce (Badiola and Cuesta, 2006 and Crochet, 1980). Miocene Herpetotheridae in Spain have been poorly documented, always restricted to indeterminable finds (Crochet, 1980 and Ziegler, 1999) or personal communications (Robles et al., 1991). In the present work, we give for the first time a detailed description of teeth attributable to a Miocene herpetotheriid from Spain, and their generic and specific ascription are thoroughly justified.

 Material and methods

Only upper dental elements have been recovered. The nomenclature for the description of the teeth and the reference lines to take the measurements follow Crochet (1980). Measurements were carried out at the Department of Geology of the University of Valencia, using a Leica MZ75 binocular microscope by means of displacement of a mechanical stage connected to Sony Magnescale measuring equipment. The fossil material is stored at the Museu de Geologia de la Universitat de València (MGUV) under catalogue numbers MGUV-25184 (MAB5-394), MGUV-25185 (MAB5-395), MGUV-25190 (MAB5-400), MGUV-25237 (MAB5-447), MGUV-25254 (MAB0B-6), MGUV-25313 (MAB4-5), MGUV-25345 (BC1-30), MGUV-25579 (MTR1-4).

Institutional abbreviations: MGUV: Museu de Geologia de la Universitat de València, Burjassot (Spain); SCSIE: Servei Central de Suport a la Investigació Experimental, Valencia (Spain); UV: Universitat de València, Valencia (Spain).

Localities: BC: Barranc de Campisano; MAB: Mas d’Antolino; MTR: Mas de Torner.

 Geographic and geologic setting

The fossil material described in the present work comes from different levels of the sections of Mas d’Antolino B (MAB0B, MAB4 and MAB5), Mas de Torner (MTR1) and Barranc de Campisano (BC1). All of them are situated in the western area of the Ribesalbes-Alcora Basin, an intramontane basin in eastern Spain (Agustí et al., 1988 and Anadón, 1983). These sections are made up of about 60 m of grey and yellow mudstones, sandstones and limestones, and they all belong to the Unit 3 Sequence defined by Anadón (1983). Twenty levels in these sections have provided micromammal fossil remains. The five beds recording the presence of herpetotheriids have provided a total of almost 200 fossil mammal teeth. The small mammal assemblages in these localities are detailed in Table 2.

The association of Megacricetodon primitivus and Ligerimys (either florancei or ellipticus) in the four localities where A. frequens is present is indicative of their Lower Miocene age (Lower Aragonian, zones B and/or C) (Daams and Freudenthal, 1988). The only site not recording Megacricetodon and Ligerimys is Mas de Torner 1. Nevertheless, its stratigraphic position with respect to the other four localities is unequivocally indicative of a similar age (Ruiz-Sánchez et al., 2010).

 Systematic paleontology and description

Infraclass: METATHERIA Huxley, 1880

Family: HERPETOTHERIIDAE Trouessart, 1879

Genus: Amphiperatherium Filhol, 1879

Amphiperatherium frequens (von Meyer, 1846) (Fig. 2)

dP3: (MAB5-400; Fig. 2A, E: L = 1.34; W = 1.65; MAB5-447; Fig. 2B, F: L = 1.41; W = 1.78; MTR1-4; Fig. 2C, G: L = 1.31; W = 1.68; BC1-30; Fig. 2D, H: L = 1.42; W = 1.74). The metacone is the highest cusp. The paracone is only slightly lower than the metacone. A small and short anterior ridge originates from the top of the paracone. The protocone is much lower than the metacone. The paraconule is discernible as a two-branch divergence of the preprotocrista in MAB5-400 and BC1-30, but it is not divided in MAB5-447 and MTR1-4. The centrocrista is less dilambdodont (more straight) than in definitive upper molars. The anterolabial corner is much protruding in occlusal view. In MAB5-400, the ectoflexus is rather straight and it displays stylar cusps A, B and C as small elevations placed at the labial edge. However, these stylar cusps are absent in MTR1-4, MAB5-447 and BC1-30 (only in the latter specimen there is a weak cusp A). In all the dP3s found, cusp D is always discernible and it is clearly the largest one of the stylar shelf. The metaconule is difficult to discern. In MAB5-400, the metaconule is represented by a faint swelling of the postprotocrista. The roots are not preserved in any specimen.

M1: (MAB5-394; Fig. 2I, L: L = 1.93; W = 2.58). The metacone is the highest cusp. Protocone and paracone have a similar height. The metaconule is a faint elevation of the posterior end of the postprotocrista. The stylar cusp B is the highest cusp of the ectoflexus. There is neither a stylar cusp C, nor a ridge connecting the stylar cusps A and B. The roots are not preserved in this specimen.

M3: (MAB5-395; Fig. 2J, M: L = 1.71). Only the labial half is preserved. The “v” shape of the labial margin suggests that this tooth is an M3. The wear affects the entire occlusal surface. The stylar cusps A, B and D are only recognized as weak rather conical elevations.

M4: (MAB0B-6; Fig. 2K, N: L = 1.46). Only the labial side is preserved. The curvature of the crista of the right half in occlusal view (Fig. 2N) suggests that it might correspond to the metacrista (posterior arm of the metacone) of an M4. Consequently, the paracone is the highest cusp, and the paracrista is the longest ridge of the preserved part. Cusp A is discernible as the anterolabial end of the paracrista. The cusp B is immediately advanced to the labial notch and it is the most prominent of the ectoflexus. Cusp E is quite well developed. Cusps C and D are extremely reduced and they are difficult to discern.

Undetermined Upper Molar or Premolar. The specimen MAB4-5 (Fig. 2O) is an undetermined upper molar or deciduous premolar, lacking the anterolabial side.

 Results

Taxonomically, the absence of a connection between the stylar cusps A and B in the upper molars justifies their ascription to the genus Amphiperatherium. The species A. frequens is distinct from A. exile by having weaker stylar cusps C and D, which sometimes disappear altogether (Crochet, 1980). The complete absence of the stylar cusp C in the M1 (Fig. 2L) and its almost complete reduction in M3 and M4 (Fig. 2M, N), support their specific ascription to A. frequens. The five characters for sub-specific identification proposed by von Koenigswald (1970) were reviewed by Crochet (1980), who concluded that actually none of them has a systematic significance. However, Ziegler (1999) considered the distinction of chrono-subspecies in the Miocene (A. f. frequens, A. f. erkertshofense or A. f. wintershofense) justified, remarking that they are only determinable when sufficient material is available. Unfortunately, the scarce material found in Ribesalbes-Alcora Basin (with only upper teeth) and the variability observed in the dP3 found (the only repeated element) do not allow a sub-specific ascription.

It is noteworthy that the discovery of four left deciduous premolars indicates the presence of at least four different juvenile to subadult individuals. This is a highly unusual occurrence of young specimens, representing half of the total dental remains of the herpetotheriid sample from all layers. No lower dental elements have been found. This is probably because Amphiperatherium is, overall, a minimal component within the micromammalian assemblages in all these localities.

The limited presence of this taxon is probably a result of several climatic and environmental factors. It is worth to mention that most of the Miocene finds of Amphiperatherium in Europe correspond to Central European countries (Fig. 1), where the environmental conditions were rather humid (Utescher et al., 2011, and references therein). Ribesalbes-Alcora belongs to the Ibero-Levantine Province, which is characterized by wetter conditions than in other places of the Iberian Peninsula (Agustí, 1978, Agustí, 1990, Agustí et al., 1984, Casanovas-Vilar and Agustí, 2007 and Sesé, 1988). This is evidenced, for instance, by the presence of Ligerimys (Daams et al., 1988 and Van der Meulen and Daams, 1992) or the flying squirrel Miopetaurista in the locality of Buñol (MN4, local zone C, Magro Basin; Daams, 1976 and Daams, 1977). Nonetheless, a lesser degree of humidity in this region with respect to Central Europe could have resulted in less abundant populations of Amphiperatherium than in Germany, Switzerland or France.

Another point to consider is the age of the sites, which coincides with the Middle Miocene Climatic Optimum (17–15 Ma, Zachos et al., 2001). Not by chance, the extinction of Amphiperatherium in central Europe also coincides with the end of this climatic event. The warm temperatures could have played an important role permitting the southern expansion of environments and populations formerly limited to Central Europe. The Ramblian and the transition to the Early Aragonian are characterized by densely forested areas in some parts of Spain (DeMiguel et al., 2010). This picture suggests that the Ribesalbes-Alcora populations were close to the southern boundary of their suitable range, and more fragile to any climatic or environmental change. In last term, a northern retraction of this boundary combined with the fragmentation of the populations could have resulted in the regional extinction of Amphiperatherium in the Iberian Peninsula.

In any case, this is the first time that the presence of the herpetotheriid A. frequens is adequately documented in the Miocene of Spain. Former references to the presence of herpetotheriids in the Neogene of the Iberian Peninsula were limited to identifications at the family level or personal communications to authors, giving no details nor figuring the material found. Consequently, this occurrence of A. frequens in the Early Aragonian (late Early Miocene) of the Ribesalbes-Alcora Basin represents the hitherto southernmost presence of this species in Europe, and the last record of a metatherian mammal in the Iberian Peninsula.

 Acknowledgements

We thank E. Navarro and A. Ibáñez (SCSIE, UV) for capable laboratory assistance. This research was supported by projects PIPH-2009 SGR 754 (AGAUR, Generalitat de Catalunya), CGL2011-28681 (Spanish Ministerio de Economía y Competitividad) and GVPRE/2008/320 (Generalitat Valenciana). The field work in Araia was supported from 2008 to 2010 by the Conselleria de Cultura of the Generalitat Valenciana. We acknowledge the useful comments on the manuscript provided by R. Ziegler, J. Prieto and two anonymous reviewers, which improved the quality of the paper.

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Map showing the countries with previously known Miocene occurrences of Amphiperatherium (grey-shaded). The new localities from Ribesalbes-Alcora Basin in Spain are indicated with an arrow.

Carte montrant les pays où la présence d’Amphiperatherium au Miocène était déjà connue (en gris). Les nouvelles localités du bassin Ribesalbes-Alcora en Espagne sont indiquées par une flèche.

Amphiperatherium frequens from MAB0B, MAB4, MAB5, MTR1 and BC1, Ribesalbes-Alcora basin, Spain. A. MAB5-400, left dP3 (labial view). B. MAB5-447, left dP3 (labial view); C, MTR1-4, left dP3 (labial view). D. BC1-30, left dP3 (labial view). E. MAB5-400, left dP3 (occlusal view). F. MAB5-447, left dP3 (occlusal view). G. MTR1-4, left dP3 (occlusal view). H. BC1-30, left dP3 (occlusal view). I. MAB5-394, right M1 (labial view). J. MAB5-395, right M3 (labial view). K. MAB0B-6, left M4 (labial view). L. MAB5-394, right M1 (occlusal view). M. MAB5-395, right M3 (occlusal view). N. MAB0B-6, left M4 (occlusal view). O. MAB4-5, undetermined upper molar or premolar (occlusal view). Scale bar equals 1 mm.

Amphiperatherium frequens de MAB0B, MAB4, MAB5, MTR1 et BC1, bassin de Ribesalbes-Alcora, Espagne. A. MAB5-400, dP3 gauche (vue labiale). B. MAB5-447, dP3 gauche (vue labiale). C. MTR1-4, dP3 gauche (vue labiale). D. BC1-30, dP3 gauche (vue labiale). E. MAB5-400, dP3 gauche (vue occlusale). F. MAB5-447, dP3 gauche (vue occlusale). G. MTR1-4, dP3 gauche (vue occlusale). H. BC1-30, dP3 gauche (vue occlusale). I. MAB5-394, M1 droite (vue labiale). J. MAB5-395, M3 droite (vue labiale). K. MAB0B-6, M4 gauche (vue labiale). L. MAB5-394, M1 droite (vue occlusale). M. MAB5-395, M3 droite (vue occlusale). N. MAB0B-6, M4 gauche (vue occlusale). O. MAB4-5, molaire ou prémolaire supérieure indéterminée (vue occlusale). Barre d’échelle : 1 mm.

Biostratigraphic chart with the localities with fossil remains of Amphiperatherium. In MN1 to MN4 from France and Germany, the order established by Ziegler (1999) prevails when no other indication of relative position exists.

Cadre biostratigraphique, avec les localités qui ont fourni des restes fossiles d’Amphiperatherium. De MN1 à MN4 en France et en Allemagne, l’ordre établi par Ziegler (1999) est suivi quand aucune autre indication sur la position relative n’existe.

France Switzerland Germany Austria Czechia Spain Datation Aguilar et al., 1999 and Aguilar et al., 2003 Kälin (1997) Heissig (1997) Daxner-Höck et al. (1998) Fejfar (1990) Ruiz-Sánchez et al. (2010) This paper Heissig (1989) Kälin and Kempf (2009) Ziegler (2006) Harzhauser et al. (2002) Kempf et al. (1997) Abdul Aziz et al. (2010) Mein (1989) MN7+8 Ergeten b Greuterschberg b MN6 Collet-Redon a Schwamendingen Gallenbach 2b Zeglingen Gisseltshausen Rümikon Unterzolling Sagentobel Laimering 3 Uzwil-Nutzenbuech Chatzloch MN5 Vermes 1 Altenstadt c Teiritzberg Hambach 6C Obergänsendorf Eitensheim Oggenhausen Oggenhof Affalterbach Massendorf Sandelzhausen Puttenhausen Niederaichbach Münchsmünster Langenmoosen MN4 Vieux-Collonges Eiboden Forsthart Oberdorf 3+4 Dolnice Mas d’Antolino 5 Port-La-Nouvelle Hubertingen Rembach Mas d’Antolino 4 La Romieu Glovelier Rauscheröd 1b Mas d’Antolino 0B Erkertshofen 1 Barranc de Campisano 1 Erkertshofen 2 Mas de Torner 1 Petersbuch 2 MN3 Beaulieu Goldinger Tobel 8 Schnaitheim Maigen Estrepouy Bierkeller St.Vincent-de-Lamontjoie Wintershof-West Serre de Verge Stubersheim 3 MN2 Balizac Schaffhausen Ulm-Uniklinik Bouzigues La Mèbre 698 Haslach La Brète La Chaux 7 Eggingen Laugnac Budenheim Montaigu Frankfurt-Nordbassin MN1 Saulcet Ulm-Westtangente Plaissan Hochstadt St. Gérand-Ie-Puy Ravolzhausen Pyrimont-Challonges Lautern 2 Les Cévennes Weisenau La Paillade Weissenburg 6 Paulhiac Relais des Cathares N.F.M.-Montpellier

The age of this locality is still under discussion ranging from MN5 to MN7 (Aguilar and Clauzon, 1979 and Heissig, 1989).

These localities are considered MN8 sensu Kälin and Kempf (2009).

Amphiperatherium was not found in new samplings from Altenstadt (Prieto et al., 2009). Data from Aguilar and Clauzon, 1979, Aguilar et al., 2003, Böttcher et al., 2009, Daxner-Höck et al., 1998, Engesser and Mödden, 1997, Engesser et al., 1993, Fejfar, 1990, Kälin, 1997, Kälin and Kempf, 2009, Mein, 1989, Prieto, 2011a, Prieto, 2011b, Ziegler, 1999 and Ziegler, 2000.

Small mammal assemblage in the five localities from the Ribesalbes-Alcora Basin with fossil teeth of Amphiperatherium.

Assemblage de micromammifères des cinq localités du bassin de Ribesalbes-Alcora, avec des dents fossiles d’Amphiperatherium.

MAB0B MAB4 MAB5 MTR1 BC1 Lithology Marls Marls Marls Marls Marls n mammal teeth 8 6 150 10 22 Amphiperatherium frequens x x x x x Plesiodimylus sp. x x Erinaceidae indet. x Talpidae indet. x Insectivora indet. x x x Sciuridae indet. x x Democricetodon/Fahlbuschia sp. x x x Megacricetodon cf. primitivus x x x x Glirudinus cf. modestus x Microdyromys sp. x Simplomys cf. julii x x Pseudodryomys cf. ibericus x Myomiminae indet. x Gliridae indet. x x x Ligerimys aff. florancei x x Ligerimys ellipticus x x