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Agencia Estatal de Investigacion-European Regional Development Fund of the European Union CGL2016-76431-P CGL2017-82654-P Generalitat de Catalunya (CERCA Program) Centre National de la Recherche Scientifique (CNRS) Generalitat de Catalunya 2017 SGR 086 2017 SGR 116

Three species of Deinotherium sensu stricto (Proboscidea, Deinotheriidae), i.e., excluding Prodeinotherium, generally considered to have nonoverlapping chronostratigraphic distributions, are currently recognized from the Miocene of Europe: Deinotherium levius (late Astaracian/Aragonian, MN7+8), Deinotherium giganteum (type species; Vallesian, MN9–MN10), and Deinotherium proavum (a senior synonym of Deinotherium gigantissimum; Turolian, MN11–MN13). Here we describe a sample of 26 cheek teeth from four roughly coeval localities of Ronda Oest de Sabadell (ROS), in the Vallès-Penedès Basin, northeastern Iberian Peninsula, dated to the latest Vallesian (∼9.4–9.1 Ma, MN10) on biostratigraphic grounds. The remains from ROS-D3 represent all the permanent upper and lower dentition and can be unambiguously assigned to D. proavum, the largest deinothere from Europe, based on their large size (well above the range of D. giganteum). Remains from the other localities (ROS-D2, ROS-D5, and ROS-D8) are smaller and generally overlap in size with both D. proavum and D. giganteum. However, an assignment to the former species is more likely given that these species are not known to co-occur and that their dental size ranges were already known to overlap partially. The remains of D. proavum from ROS represent the oldest record of this species, the occurrence of which in the late Vallesian had already been recorded in the slightly younger (∼9.1 Ma) locality of Sinap 49, Turkey. Given that D. proavum is recorded before the Vallesian/Turolian boundary, and that smaller individuals of this species overlap in size with D. giganteum from the Vallesian, caution is required when making biostratigraphic correlations based on late Miocene deinothere remains from Europe.

We describe new dental remains of the genus Iberictis (Carnivora: Mustelidae) from the late early Miocene of the Iberian Peninsula. The new fossils of Iberictis azanzae from Artesilla (16.5–16.3 Ma, MN4; Calatayud-Teruel Basin, Zaragoza, Spain) add important morphological information about this species. Material from another species, Iberictis buloti, is described from els Casots (16.5–16.3 Ma, MN4; Vallès-Penedès Basin, Barcelona, Spain). This material constitutes the most complete sample of Iberictis and the first record of I. buloti in the Iberian Peninsula. Our analyses confirm the taxonomic validity of both species, and confirm the more plesiomorphic status of I. buloti compared to I. azanzae. Re-examination of large mustelid Miocene genera (Dehmictis, Ekorus, Eomellivora, Hoplictis, Iberictis, Ischyrictis, and Plesiogulo) and their inclusion for the first time in a cladistic analysis indicate that Iberictis is the sister taxon of Plesiogulo, and that these genera constitute the sister group of the extant wolverine (Gulo gulo). Our analysis thus confirms a close relationship between the early Miocene Iberictis, the late Miocene Plesiogulo, and the Plio-Pleistocene Gulo. Iberictis is the oldest member of Gulonini, the total clade of wolverines, thereby tracking the fossil record of this clade back to the early Miocene. We further propose a new systematic arrangement for the aforementioned large Miocene mustelids into the subfamilies Guloninae, Mellivorinae, and Mustelinae., This research received support by A.V. from the SYNTHESYS3 Project http://www.synthesys.info/(SYNTHESYS; AT-TAF-5457), which is financed by European Community Research Infrastructure Action under the FP7 Capacities Programme, and the European Union’s Seventh Framework Programme (FP7/2007-2013) under grant agreement n° 226506 (SYNTHESYS; SE-TAF-3637). A.V. also received support by an American Museum of Natural History Collection Study Grant Program 2014. The support ofthe DST-NFR Centre of Excellence in Palaeosciences (CoE-Pal) towardthis research for A. V. (COE2018-09POST) is hereby acknowledged. This study was alsosupported by CGL2015-68333-P (MINECO/FEDER-UE), Agencia Estatal de Investigación–European Regional Development Fund of the European Union (CGL2016-76431-P, AEI/FEDER, EU), the Generalitat de Catalunya (CERCA Programme), and the research group UCM910607.

Villa A, Abella J, Alba DM, Almécija S, Bolet A, Koufos GD, et al., Monitor lizards (genus Varanus) inhabited Europe at least from the early Miocene to the Pleistocene. Their fossil record is limited to about 40 localities that have provided mostly isolated vertebrae. Due to the poor diagnostic value of these fossils, it was recently claimed that all the European species described prior to the 21st century are not taxonomically valid and a new species, Varanus amnhophilis, was erected on the basis of fragmentary material including cranial elements, from the late Miocene of Samos (Greece). We re-examined the type material of Varanus marathonensis Weithofer, 1888, based on material from the late Miocene of Pikermi (Greece), and concluded that it is a valid, diagnosable species. Previously unpublished Iberian material from the Aragonian (middle Miocene) of Abocador de Can Mata (Vallès-Penedès Basin, Barcelona) and the Vallesian (late Miocene) of Batallones (Madrid Basin) is clearly referable to the same species on a morphological basis, further enabling to provide an emended diagnosis for this species. Varanus amnhophilis appears to be a junior subjective synonym of V. marathonensis. On the basis of the most complete fossil Varanus skeleton ever described, it has been possible to further resolve the internal phylogeny of this genus by cladistically analyzing 80 taxa coded for 495 morphological and 5729 molecular characters. Varanus marathonensis was a large-sized species distributed at relatively low latitudes in both southwestern and southeastern Europe from at least MN7+8 to MN12. Our cladistic analysis nests V. marathonensis into an eastern clade of Varanus instead of the African clade comprising Varanus griseus, to which it had been related in the past. At least two different Varanus lineages were present in Europe during the Neogene, represented by Varanus mokrensis (early Miocene) and V. marathonensis (middle to late Miocene), respectively., This research has been supported by the Spanish Agencia Estatal de Investigación (Ministerio de Economía, Industria y Competitividad)–European Regional Development Fund of the European Union (CGL2016-76431-P and CGL2017-82654-P, AEI/FEDER EU), by the Generalitat de Catalunya (CERCA Programme) and by the Italian MIUR (PRIN 2009MSSS9L_002). The study of the Batallones material was carried out thanks to the SYNTHESYS Project funded by European Community Research Infrastructure Action under the FP7 ‘Structuring the European Research Area’ Programme (ES-TAF-1975 to MD). Access to the collections of Natural History Museum Wien was also possible thanks to a SYNTHESYS grant (AT-TAF-4591) to AV. AB is a Newton International Fellow (NF160464), funded by the Royal Society. Fieldwork at ACM was funded by CESPA Gestión de Residuos, S.A.U. The Cerro the los Batallones project was funded by the Ministerio de Economía y Competitividad de España (CGL2015-68333-P), the Research Groups CSIC 641538 and CAM-UCM 910607, and the Dirección General de Patrimonio (Comunidad Autónoma de Madrid).

Flying squirrels are the only group of gliding mammals with a remarkable diversity and wide geographical range. However, their evolutionary story is not well known. Thus far, identification of extinct flying squirrels has been exclusively based on dental features, which, contrary to certain postcranial characters, are not unique to them. Therefore, fossils attributed to this clade may indeed belong to other squirrel groups. Here we report the oldest fossil skeleton of a flying squirrel (11.6 Ma) that displays the gliding-related diagnostic features shared by extant forms and allows for a recalibration of the divergence time between tree and flying squirrels. Our phylogenetic analyses combining morphological and molecular data generally support older dates than previous molecular estimates (~23 Ma), being congruent with the inclusion of some of the earliest fossils (~36 Ma) into this clade. They also show that flying squirrels experienced little morphological change for almost 12 million years., eLife digest Mammals can walk, hop, swim and fly; a few, like marsupial sugar gliders or colugos, can even glide. With 52 species scattered across the Northern hemisphere, flying squirrels are by far the most successful group that adopted this way of going airborne. To drift from tree to tree, these small animals pack their own ‘parachute’: a membrane draping between their lower limbs and the long cartilage rods that extend from their wrists. Tiny specialized wrist bones, which are unique to flying squirrels, help to support the cartilaginous extensions. The origin of flying squirrels is a point of contention: while most genetic studies point towards the group splitting from tree squirrels about 23 million years ago, the oldest remains – mostly cheek teeth – suggest the animals were already soaring through forests 36 million years ago. However, recent studies show that the dental features used to distinguish between gliding and non-gliding squirrels may actually be shared by the two groups. In 2002, the digging of a dump site in Barcelona unearthed a peculiar skeleton: first a tail and two thigh bones, big enough that the researchers thought it could be the fossil of a small primate. In fact, and much to the disappointment of paleoprimatologists, further excavating revealed that it was a rodent. As the specimen – nearly an entire skeleton – was being prepared, paleontologists insisted that all the ‘dirt’ attached to the bones had to be carefully screen-washed. From the mud emerged the minuscule specialized wrist bones: the primate-turned-rodent was in fact Miopetaurista neogrivensis, an extinct flying squirrel. Here, Casanovas-Vilar et al. describe the 11.6 million years old fossil, the oldest ever found. The wrist bones reveal that the animal belongs to the group of flying squirrels that have large sizes. Evolutionary analyses that combined molecular and paleontological data demonstrated that flying squirrels evolved from tree squirrels as far back as 31 to 25 million years ago, and possibly even earlier. In addition, the results show that Miopetaurista is closely related to Petaurista, a modern group of giant flying squirrels. In fact, their skeletons are so similar that the large species that currently inhabit the tropical and subtropical forests of Asia could be considered living fossils. Molecular and paleontological data are often at odds, but this fossil shows that they can be reconciled and combined to retrace history. Discovering older fossils, or even transitional forms, could help to retrace how flying squirrels took a leap from the rest of their evolutionary tree.

New remains of felid jaws and teeth are described from several localities of the local stratigraphic series of Abocador de Can Mata (ca. 11.9 to 11.6 Ma, Middle Miocene; Valles-Penedes Basin, Catalonia, Spain). Three different taxa are identified: Styriofelis turnauensis , Pseudaelurus romieviensis and Pseudaelurus quadridentatus . The described remains of P. romieviensis enable extending considerably the chronological range of this species in the Iberian Peninsula, in agreement with its record in the rest of Europe. Moreover, it is shown for the first time that P. romieviensis may possess a p2. The presence of this tooth therefore does not constitue a valid diagnostic feature to distinguish P. romieviensis from P. quadridentatus .

The Bovidae are one of the most fascinating cetartiodactyl families, not only because of their complicated taxonomic structure, which includes a large plethora of living (cattle, antelopes, goats) ...

Anchitherium nievei sp. nov. is erected on the basis of dentognathic remains from the late Aragonian (MN7+8, ca. 12.3–12.1 to 11.7–11.6 Ma, Middle Miocene) of three localities from the Abocador de Can Mata local stratigraphic series (els Hostalets de Pierola, Valles-Penedes Basin, Catalonia, Spain). This taxon differs from endemic species of Anchitherium from the inner Iberian basins, most closely resembling A. aurelianense, A. hippoides and A. steinheimense by its marked brachyodonty and small dental size, although differing by dental proportions and several details of occlusal morphology. From a paleoenvironmental viewpoint, the presence of this taxon fits well with previous inferences that indicated a relatively closed and humid forested habitat, more similar to that of similarly aged French localities.

During the last decade, new discoveries in several Iberian basins, together with the description of previously unpublished finds, have significantly increased the recorded paleodiversity of fossil Primates (Mammalia: Euarchonta) in the Iberian Peninsula. Here we provide an updated compendium of the primate fossil record in Iberia during the Cenozoic and further summarize the changes in primate paleodiversity through time, which are then analyzed in the light of changing climatic conditions. Thanks to favorable climatic conditions, the highest diversity of Iberian primates was reached during the Eocene, thus reflecting the radiation of both adapoids and omomyoids; only a single plesiadapiform genus is in contrast recorded in the Iberian Peninsula. Near the Eocene-Oligocene boundary, paleoclimatic changes led to a primate diversity crisis and other faunal changes, although two Iberian omomyoids survived the Grande Coupure. From the Middle Miocene onwards, catarrhine primates are recorded in the Iberian Peninsula. During the Middle and Late Miocene, they are represented by pliopithecoids and hominoids, restricted to NE Iberia. The Miocene hominoids from Iberia are of utmost significance for understanding the Eurasian hominoid radiation and its role in the origins of the great-ape-and-human clade. Following the local extinction of these taxa during the early Late Miocene, due to progressively increased seasonality and concomitant changes in plant communities, cercopithecoids are also recorded in the Iberian Peninsula from the latest Miocene through the Plio-Pleistocene, although they finally became locally extinct, whereas hominoids are again represented by fossil humans during the Pleistocene. Durante la última década, nuevos descubrimientos en varias cuencas ibéricas, junto con la descripción de hallazgos previos inéditos, han incrementado significativamente la paleodiversidad de Primates fósiles (Mammalia: Euarchonta) registrada en la Península Ibérica. Proporcionamos aquí un compendio actualizado del registro fósil de los primates en Iberia durante el Cenozoico, y resumimos además los cambios en paleodiversidad de los primates a lo largo del tiempo a la luz de las condiciones climáticas cambiantes. Gracias a condiciones climáticas favorables, la diversidad más alta de primates ibéricos se produjo durante el Eoceno, reflejando así la radiación tanto de los adapoideos como de los omomioideos; en cambio, sólo un único género de plesiadapiformes se registra en la Península Ibérica. Hacia el límite Eoceno-Oligoceno, los cambios paleoclimáticos condujeron a una crisis de diversidad de los primates y otros cambios faunísticos, aunque dos omomioideos ibéricos sobrevivieron a la Grande Coupure. Del Mioceno Medio en adelante, los primates catarrinos se registran en la Península Ibérica. Durante el Mioceno Medio y Superior, están representados por pliopitecoideos y hominoideos, restringidos al NE de Iberia. Los hominoideos del Mioceno de Iberia son de gran importancia para comprender la radiación de los hominoideos eurasiáticos y su papel en los orígenes del clado de los grandes antropomorfos y los humanos. A continuación de la extinción local de estos taxones durante el Mioceno Superior inicial, debido al incremento progresivo de la estacionalidad así como a cambios en las comunidades vegetales, los cercopitecoideos también se registran en la Península Ibérica a partir del Mioceno más terminal en adelante. Los cercopitecoideos se registran en la Península Ibérica durante todo el Plio-Pleistoceno, aunque finalmente también se extinguieron localmente, mientras que los hominoideos vuelven a estar representados otra vez por los humanos fósiles durante el Pleistoceno.

The evolution of blanid amphisbaenians (Mediterranean worm lizards) is mainly inferred based on molecular studies, despite their fossils are common in Cenozoic European localities. This is because the fossil record exclusively consists in isolated elements of limited taxonomic value. We describe the only known fossil amphisbaenian skull from Europe – attributed to Blanus mendezi sp. nov. (Amphisbaenia, Blanidae) – which represents the most informative fossil blanid material ever described. This specimen, from the Middle Miocene of Abocador de Can Mata (11.6 Ma, MN7+8) in the Vallès-Penedès Basin (Catalonia, NE Iberian Peninsula), unambiguously asserts the presence of Blanus in the Miocene of Europe. This reinforces the referral to this genus of the previously-known, much more incomplete and poorly-diagnostic material from other localities of the European Neogene. Our analysis – integrating the available molecular, paleontological and biogeographic data – suggests that the new species postdates the divergence between the two main (Eastern and Western Mediterranean) extant clades of blanids, and probably precedes the split between the Iberian and North-Western African subclades. This supports previous paleobiogeographic scenarios for blanid evolution and provides a significant minimum divergence time for calibrating molecular analyses of blanid phylogeny.

Here, we report on an isolated pliopithecid M3/ (IPS35028) from locality ACM/C3-B2 (12.0 Ma, MN7) of the late Middle Miocene stratigraphic series of Abocador de Can Mata (ACM, Valles-Penedes Basin, NE Iberian Peninsula). This tooth is about 0.2 million years older than the remains of Pliopithecus canmatensis (11.8-11.7 ma), recorded from several localities from the ACM series. The unusual occlusal features of IPS35028, together with the lack of homologous material for several pliopithecid species, preclude a precise taxonomic attribution of the C3-B2 specimen, which does not fit the morphology of any known pliopithecid M3/. In particular, although an attribution to P. canmatensis would seem reasonable on the basis of size, identical geographic provenance, and similar age, the morphology of IPS35028 appears too primitive compared to the M1/ and M2/ of the former taxon. Instead, the C3-B2 pliopithecid displays several primitive features shared with the dionysopithecine Dionysopithecus and the pliopithecine Pliopithecus piveteaui. It therefore seems more likely that IPS35028 represents a previously unknown pliopithecid taxon, although a formal taxonomic recognition of its probable distinct status is not advisable, given the scarcity of the currently available material. Alternatively, this taxon might be more closely related to small-bodied African catarrhines (such as dendropithecids). However, the morphology of the ACM specimen is not particularly similar to that of the M3/ of these African taxa. Hence, based on age and geographic provenance, an attribution of IPS35028 to the Pliopithecidae is favored here.

Pliopithecus (Pliopithecus) canmatensis sp. nov. is described from several Late Aragonian localities from Abocador de Can Mata (ACM) in els Hostalets de Pierola (Valles-Penedes Basin, Catalonia, Spain), spanning from ∼11.7 to 11.6 Ma (C5r.3r subchron), and being correlated to the MN8 (reference locality La Grive L3). The ACM remains display a pliopithecine dental morphology with well-developed pliopithecine triangles on M/2 and M/3. This, together with other occlusal details, negates an attribution to the subgenus Epipliopithecus. Although slightly smaller, the ACM remains are most similar in size to comparable elements of P. piveteaui and P. antiquus. Several occlusal details (such as the greater development of the buccal cingulid in lower molars) and dental proportions (M/3 much longer than M/2), however, indicate greater similarities with P. antiquus from Sansan and La Grive. The ACM remains, however, differ from P. antiquus in dental proportions as well as occlusal morphology of the lower molars (including the less peripheral position of the protoconid and more medial position of the hypoconulid, the more mesial position of the buccal cuspids as compared to the lingual ones, the narrower but distinct mesial fovea, the higher trigonid, and the more extensive buccal cingulid, among others). These differences justify a taxonomic distinction at the species level of the ACM pliopithecid remains with respect to P. antiquus. Previous pliopithecid findings from the Valles-Penedes Basin, previously attributed to P. antiquus, are neither attributable to the latter species nor to the newly erected one. Am J Phys Anthropol, 2010. © 2009 Wiley-Liss, Inc.