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Highlights The oldest human evidence in western Europe is currently documented at 1.4 Ma. Understanding the climatic variability that these hominins encountered is relevant. Paleoclimatic data are provided for an archaeopaleontological sequence. The Early Pleistocene hominins were able to cope with changing climatic conditions. Abstract The oldest evidence of a human presence in western Europe is currently documented at the late Early Pleistocene sites of Barranco León and Fuente Nueva 3 (Guadix-Baza Basin, SE Spain), dated respectively to 1.4 and 1.2 Ma. Understanding the climatic variability that these hominins encountered and coped with is of outstanding importance for placing their activities in an ecological context and understanding their capacity to adapt to changing conditions. Glacial-interglacial variability during this period was considerably less intense than during later phases of the Pleistocene. To date, however, no quantitative estimates are available for this climate variability in Early Pleistocene hominin sites, and no ‘cold’ reconstructions directly associated with the early hominin occupations of Europe have been performed either. Here stratigraphically constrained quantitative climatic reconstructions are provided for the sites of Barranco León and Fuente Nueva 3, using an improvement on the Mutual Ecogeographic Range method by projecting the niche envelope of the extant representatives ofOphisaurussp. We ascertain differences in temperature and rainfall between the different layers of the two sites, in line with previous reconstructions that revealed warm, humid periods (‘interglacial') as well as more temperate but drier periods (‘glacial'), consistent with Early Pleistocene climate cyclicity. Our new estimates suggest that late Early Pleistocene hominins, though conditioned to some degree by climatic factors, were able to deal with changing climatic and environmental conditions (‘interglacial' and ‘glacial') in the southwestern extremity of the European continent.

This volume honours Professor Jorge Morales – one of the world´s leading experts in fossil carnivorans, a great scholar, and inspiration of many to follow him in the discipline of palaeontology – to mark his recent retirement. In July 2021, Prof. Morales retired after 50 years working and building up a reputation not only as a leading scientist but also as a teacher, facilitator of science, and mentor. This festschrift celebrates his lifelong, outstanding scientific and cultural achievements to mammal palaeontology. It contains contributions by colleagues, former students, and friends that come from far and wide. Articles pay homage to the jubilarian´s research interests, covering the systematics, palaeoecology, biochronology, palaeobiogeography, taphonomy, and geoheritage. On the basis of the written acknowledgements, it is easy to recognise the affection and deep esteem Prof. Morales induces in everyone. If we think about him, our professional and personal lives would not be the same without his support, mentorship, friendship, and good humour. It is a proud privilege to thank him for his career in terms of production and excellence which without doubt placed Spanish palaeontology among the global elite. Prof. Morales is retired, but he remains active in research, and as inspiration to all of us., This publication is part of projects PID2020-116220GB-I00 and PGC2018-094122-B- I00 funded by the Ministerio de Ciencia e Innovación/Agencia Estatal de Investigación/10.13039/501100011033/, with funding also by the Government of Aragon (Research Group ref. E33_20R).

Venta Micena (Orce, Guadix-Baza Basin, Spain) is an Early Pleistocene locality renowned for the richness and quality of its palaeontological record. VM is spread over an area of 2.5 km2, where several exposed fossil outcrops are visible amidst its gorges and ravines. The best known of these sites, VM3, has been interpreted as a hyaena den. In addition, a new site, named VM4, has recently been the focus of fieldwork and taphonomic studies. The publication by Luz on et al. (2021) pointed out that VM4 presents a more complex history than VM3. First, two different sub-levels were identified: VM4-I and VM4-II. Secondly, the preliminary taphonomic analysis showed conspicuous differences with regard to VM3. Nevertheless, such interpretation has been challenged by Palmqvist et al. (2022) who proposed that VM3 and VM4 are both the result of a single depositional process, entailing the selective transport of skeletal parts by the giant extinct hyaena Pachycrocuta brevirostris back to its den. Using well-preserved faunal elements whose depositional context and provenance are reliable, in this paper we show that: 1) there are two clearly defined sub-levels in VM4 with some shared taphonomic characteristics as well as some notable differences; 2) VM3 and VM4 exhibit enough divergence to support differences in site formation processes; 3) The interpretation of both VM4-I and VM4-II is more consistent with their characterisation as open-air sites in which multiple agents and depositional processes contributed to its formation, rather than with hyaena dens. Nevertheless, excavations are still in progress at VM4 and therefore any results and interpretations ought to be considered as provisional., Consejeía de Cultura de la Junta de Andalucia through the General Research Project Primeras ocupaciones humanas y contexto paleoecologico a partir de los depositos Plio-pleistocenos de la cuenca Guadix-Baza BC.03.032/17, Inside the Artefacts & Ecofacts" Excellence Unit (University of Granada) HUM-607 PID 2021.125098NB.I00 MCIN/AEI/10.13039/501100011033/FEDER, Una manera de hacer Europa ProyExcel_00274 2021 SGR 01238, Agencia de Gestio D'Ajuts Universitaris de Recerca Agaur (AGAUR) Generalitat de Catalunya, National Health and Medical Research Council (NHMRC) of Australia PRE2020-094482 CEX2019-000945-M-20-1, Spanish Government, Archaeometrical Studies. Inside the Artefacts & Ecofacts" Excellence Unit (University of Granada

Venta Micena is an area containing several palaeontological sites marking the beginning of the Calabrian stage (Early Pleistocene). The richness of the fossil accumulation including species of Asian, African and European origin, makes Venta Micena a key site for the the palaeoecological and palaeoenvironmental study of southern Europe during the Early Pleistocene. Thus, research has been focused on Venta Micena 3, which was originally interpreted as a single palaeosurface associated with a marshy context, in which most of the fauna was accumulated by Pachycrocuta brevirostris. Recent excavations have unearthed a new site, Venta Micena 4, located in the same stratigraphic unit (Unit C) and in close proximity to Venta Micena 3. Here we show the frst analyses regarding the taphonomic and spatial nature of this new site, defning two stratigraphic boundaries corresponding to two diferent depositional events. Furthermore, the taphonomic analyses of fossil remains seem to indicate a diferent accumulative agent than Pachycrocuta, thus adding more complexity to the palaeobiological interpretation of the Venta Micena area. These results contribute to the discussion of traditional interpretations made from Venta Micena 3., Regional Government of Andalusia, with both contracts associated to General Research Project BC.03.032/17, Spanish Public Employment Service (Spanish Ministry of Labour and Social Economy), Spanish Ministry of Science, Innovation and Universities, with an FPI Predoctoral Grant (Reg. PRE2019-089411) associated to project RTI2018-099850-B-IOO and the University of Salamanca, Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA) has received fnancial support from the Spanish Ministry of Science and Innovation through the “María de Maeztu” program for Units of Excellence (CEX2019-000945-M), Projects 2017SGR-859 (Gov. of Catalonia, AGAUR), and CGL2016-80000-P (Spanish Ministry of Economy and Competitiveness), Research Group HUM-607 (Junta de Andalucía).

The fossil record of lynxes provides clear evidence of a large range across the North Hemisphere during the Pliocene and Pleistocene. However, their origin, systematics and evolutionary relationships are still fraught with difficulties and controversy. Here we report a complete hemimandible of a medium-sized felid from the Early Pleistocene (MN17, middle Villafranchian, 2.05 Ma) site of La Puebla de Valverde (Teruel, Spain). Based on comparative and multivariate analyses of the lower dentition of 458 individuals of medium-sized Lynx, Caracal and Leptailurus, we confidently ascribe the remains to Lynx aff. issiodorensis. Although the dental proportions are somewhat different from those of the Eurasian L. issiodorensis (smaller canines and more elongated p4/m1), Lynx aff. issiodorensis shows affinities with the Issoire lynx from the contemporaneous site of Saint Vallier (France), sharing a similar morphology of the mandible, reduced canines, and long m1. We further test the hypothesis that examines the presence of the African/Asian Caracal in the European Plio/Pleistocene for C. depereti and C. issiodorensis, and discard the attribution of L. issiodorensis into Caracal. This mandible extends the record of the genus and contributes to update our understanding of the Lynx lineage and its variability within the European fossil record.

Altres ajuts: CERCA Programme/Generalitat de Catalunya Although the suid assemblages from the Miocene of the Vallès-Penedès Basin (NE Iberian Peninsula) are reasonably well known, taxonomic studies devoted to them have lagged behind in recent decades. We describe the unpublished suid dentognathic remains from the earliest Vallesian (MN9) of Creu de Conill 20 (CCN20; 11.18 Ma), which represents the First Appearance Datum of hipparionin equids in western Europe. The sample includes 118 specimens, mostly isolated teeth, and a few maxillary and mandibular fragments. More than three-quarters of the specimens are assigned to the suine Propotamochoerus palaeochoerus, which is characteristic of MN9, albeit the described remains are slightly larger than average for the species. The rest of the sample belongs to a large tetraconodontine that is assigned to Parachleuastochoerus valentini, recorded elsewhere from MN7+8 to MN9, except for two specimens attributed to the small suid cf. Albanohyus sp. Our results support a synchronous dispersal of Hippotherium and P. palaeochoerus into Western Europe at ~ 11.2 Ma, suggesting that the latter is a suitable biochronological marker of the Vallesian. In turn, the remains of Pa. valentini refine our knowledge of the dental morphology of this species and strengthen the view that this species (unlike Conohyus doati and Conohyus melendezi) is not a junior synonym of Conohyus simorrensis. The lack of Listriodon splendens and Versoporcus sp. from CCN20, together with the scarcity of Albanohyus, contrasts with their abundance in the roughly coeval site of Castell de Barberà, hinting at local paleoenvironmental differences.

The Abocador de Can Mata (ACM) composite stratigraphic sequence (els Hostalets de Pierola, Vallès-Penedès Basin, NE Iberian Peninsula) has yielded a diverse primate assemblage from the late Aragonian (Middle to Late Miocene). Detailed litho-, bio-, and magnetostratigraphic control has enabled an accurate dating of these fossil remains. Comparable data, however, were lacking for the nearby locality of Can Mata 1 (CM1), which yielded a dryopithecine canine of a female individual. Given the lack of hipparionin equids and giraffids, CM1 has been correlated to the latest Aragonian (Mammal Neogene [MN] zone MN7+8). Here we revise the age of CM1 based on fieldwork and associated paleomagnetic samplings undertaken in 2018–2021. Our results extend the ACM composite sequence upward and indicate that CM1 correlates to the earliest Vallesian (MN9). The updated ACM sequence has a thickness of ∼300 m and comprises 12 magnetozones correlated to subchrons C5Ar.1r to C5n.2n (∼12.6–11.1 Ma; latest MN6 to earliest MN9, late Aragonian to earliest Vallesian). CM1 is correlated to C5r.1r (11.146–11.056 Ma), with an interpolated age of 11.11 Ma, thus postdating the dispersal of hipparionin horses into the Vallès-Penedès Basin—which is correlated to the previous subchron C5r.1n, with an interpolated age of 11.18 Ma, and by definition marks the beginning of the Vallesian. CM1 also minimally postdates the earliest record of giraffids at ACM—representing their earliest well-dated occurrence in the basin—being correlated to C5r.1n with an interpolated age of 11.11 Ma. We conclude that CM1 has an earliest Vallesian (MN9) age of ∼11.1 Ma, intermediate between the Aragonian dryopithecins and the Vallesian hispanopithecins. Ongoing paleontological surveillance at ACM thus offers the prospect to yield additional earliest Vallesian ape remains, which are essential to clarify their taxonomic allocation as well as to confirm whether hispanopithecins evolved locally from dryopithecins rather than immigrating from elsewhere during MN9., This publication is the result of R+D+I projects PID2020-117289GB-I00, PID2020-116908GB-I00, PID2020-116220GB-I00, PID2020-117118GB-I00, and PID2019-106440GB-C21, funded by the Ministerio de Ciencia e Innovación/Agencia Estatal de Investigación/10.13039/501100011033/. The research has been funded also by the Generalitat de Catalunya, including: CERCA Programme; Agència de Gestió d'Ajuts Universitaris i de Recerca (Consolidated Research Groups 2017 SGR 086, 2017 SGR 116, and 2017 SGR 596); and Departament de Cultura (CLT009/18/00071 and ARQ001SOL-136-2022).

We would like to thank all the curators, collection manager, and scientists who helped us allowed us to study and provided access to material: C. Argot and G. Billet (Muséum national d’Histoire naturelle, Paris), D. Geraads (Museum national d’Histoire Naturelle) for providing the Sivatherium specimen, E. Robert (Université Claude Bernard, Lyon 1), D. Berthet (Musée des Confluences Lyon), M. Orliac (Université Montpellier 2) for providing the bony labyrinth of Bachitherium, F. Duranthon and Y. Laurent (Muséum d’histoire naturelle, Toulouse), S. Legal, P. Coster, C. Balm, O. Maridet, O. Lapauze and J. Tissier (Parc Naturel Régional du Luberon and excavation team Murs Project), A. de Perthuie for accessing his private collection, Christiane Zeitler, R. Ziegler and E. Heizmann (Staatliches Museum für Naturkunde Stuttgart), P. Brewer, A. Garbout, and F. Ahmed (Natural History Museum, London), U. Göhlich and G. Daxner-Höck (Naturhistorisches Museum Wien), A. Van der Geer (Netherland Centre for Biodiversity Leiden), R. C. Hulbert Jr. (University of Florida, Gainesville), A. M. García Forner and P. Montoya (Museu de Geologia de la Universitat de València, Burjassot) for providing the petrosals of Birgerbohlinia, M. Pina (University of Manchester) for scanning support, and J. Morales (Museo Nacional de Ciencias Naturales, Madrid), J. Galkin, R. O’Leary, M. Hill Chase, C. Grohé and A. Gishlick (AMNH New York, USA), M. Celik (University of Queensland), and M. Scheidegger. We are also grateful to all the persons and institutions who scanned for us. Tandra Fairbanks (NMB) is thanked for her help with the English. B.M. and L.C. are grateful to the Swiss National Science Foundation for supporting this research through the projects 200021_178853 and 200021_159854/1 on the ear region evolution in ruminants. B.M. P300P2_161065 and P3P3P2_161066 on the evolution of the early ruminants. F.B. acknowledges support from a Gerstner Scholarship at the American Museum of Natural History. G.R. thanks the German Research Foundation project RO 1197/3-1. G.M., B.M., and L.C. want to thank the Museum National d’Histoire Naturelle de Paris for financing the Ast-RX-2013-051 Project. D.D.M. acknowledges R+D+I project ref. PID2020-116220GB-I00 from the Ministerio de Ciencia e Innovación/Agencia Estatal de Investigación/10.13039/501100011033/. I.S. acknowledges support by the Spanish Ministry of Science and Innovation (projects ref. PID2020-117289GB-I00 and PID2020-116220GB-I00), and the Generalitat de Catalunya (CERCA Programme). M.K. is funded by Japan Society for the Promotion of Science (KAKENHI Grant No. 19K04060). M.R. acknowledges MINECO project CGL2011-25754 for providing funding for the analysis. S.W. thanks the Strategic Priority Research Program of Chinese Academy of Sciences (XDB26000000) and the National Natural Science Foundation of China (41872001). G.S. and B.Mü. acknowledge financial support from the Swiss National Science Foundation in the frame of the R’equip initiative (316030_133802). Publisher Copyright: © 2022, The Author(s). Extrinsic and intrinsic factors impact diversity. On deep-time scales, the extrinsic impact of climate and geology are crucial, but poorly understood. Here, we use the inner ear morphology of ruminant artiodactyls to test for a deep-time correlation between a low adaptive anatomical structure and both extrinsic and intrinsic variables. We apply geometric morphometric analyses in a phylogenetic frame to X-ray computed tomographic data from 191 ruminant species. Contrasting results across ruminant clades show that neutral evolutionary processes over time may strongly influence the evolution of inner ear morphology. Extant, ecologically diversified clades increase their evolutionary rate with decreasing Cenozoic global temperatures. Evolutionary rate peaks with the colonization of new continents. Simultaneously, ecologically restricted clades show declining or unchanged rates. These results suggest that both climate and paleogeography produced heterogeneous environments, which likely facilitated Cervidae and Bovidae diversification and exemplifies the effect of extrinsic and intrinsic factors on evolution in ruminants. publishersversion published

The Guadix-Baza Basin (GBB) in Andalucía, Spain, comprises palaeontological and archaeological sites dating from the Early Pliocene to the Middle Pleistocene, including some of the earliest sites with evidence for the presence of early humans (Homo sp.) in Europe. Thus, the history of climate and environments in this basin contributes significantly to our understanding of the conditions under which early humans spread into Europe during the Early Pleistocene. Here we present estimates of precipitation and primary productivity in the GBB from the Pliocene to the Middle Pleistocene based on dental ecometrics in fossil communities of large herbivorous mammals, and perform an ecometrics-based distribution modelling to analyse the environmental conditions of Early and Middle Pleistocene human sites in Europe. Our results show that Early Pleistocene humans generally occupied on average relatively diverse habitats with ecotones, such as woodlands and savannas, but avoided very open and harsh (cool or dry) environments. During the Middle Pleistocene in Europe, humans occupied a comparatively much broader range of environments than during the Early Pleistocene, but were on average more concentrated in environments where the dental ecometric of mammals indicate wooded palaeoenvironments. In the earliest human occupation sites of the GBB, Barranco Leon and Fuente Nueva 3, the mean annual precipitation and net primary production estimates indicate climatic conditions close to modern Mediterranean sclerophyllous woodland environments, but with slightly higher primary productivity,indicating some similarity with East African woodlands. On the other hand, the environments did not resemble African grassland savannas. The browse-dominated diets of ungulates from Barranco Leon and Fuente Nueva 3 further suggest palaeoenvironments where grasses were a minor component of the vegetation. In the slightly older site of Venta Micena that has no evidence for the presence of hominins, dental ecometric estimates indicate climate and environments similar to Mediterranean “forest steppe” environments existing in the surroundings of Baza today. Grasses were prevalent in the diet of some taxa, especially equids, in Venta Micena, but most of the species show browse-dominated diets even there.

The two main primate groups recorded throughout the European Miocene, hominoids and pliopithecoids, seldom co-occur. Due to both their rarity and insufficiently understood palaeoecology, it is currently unclear whether the infrequent co-occurrence of these groups is due to sampling bias or reflects different ecological preferences. Here we rely on the densely sampled primate-bearing sequence of Abocador de Can Mata (ACM) in Spain to test whether turnovers in primate assemblages are correlated with palaeoenvironmental changes. We reconstruct dietary evolution through time (ca. 12.6-11.4 Ma), and hence climate and habitat, using tooth-wear patterns and carbon and oxygen isotope compositions of enamel of the ubiquitous musk-deer Micromeryx.Our results reveal that primate species composition is strongly correlated with distinct environmental phases. Large-bodied hominoids (dryopithecines) are recorded in humid, densely-forested environments on the lowermost portion of the ACM sequence. In contrast, pliopithecoids inhabited less humid, patchy ecosystems, being replaced by dryopithecines and the small-bodied Pliobates toward the top of the series in gallery forests embedded in mosaic environments.These results support the view that pliopithecoid primates preferred less humid habitats than hominoids, and reveal that differences in behavioural ecology were the main factor underpinning their rare co-occurrence during the European Miocene. Our findings further support that ACM hominoids, like Miocene apes as a whole, inhabited more seasonal environments than extant apes. Finally, this study highlights the importance of high-resolution, local investigations to complement larger-scale analyses and illustrates that continuous and densely sampled fossiliferous sequences are essential for deciphering the complex interplay between biotic and abiotic factors that shaped past diversity.

The study of skeletochronology and bone tissue as a record of information on ontogenetic stages and events is widely used for improving the knowledge about life histories (LHs) of extinct and extant vertebrates. Compared with dinosaurs and extant reptiles, mammalian bone histology has received little attention. Here, we calibrate for the first time bone and dental age with histological bone characteristics and LH stages in ontogenetic series of red deer. We rely on known LHs of different aged individuals of captive Cervus elaphus hippelaphus from Austria to correlate epiphyseal closure, dental eruption pattern, bone growth marks and bone tissue patterns in femora and tibiae, and of wild Cervus elaphus hispanicus from Spain. Our data show that females (of both subspecies) attain skeletal maturity earlier than males. At this moment, epiphyseal closure (in femora and tibiae) and dental eruption are complete and long bones start to deposit an external fundamental system. The results also show that the attainment of reproductive maturity in red deer occurs slightly before skeletal maturity.

Geoconservation and geoethics are two emergent domains in geosciences. During the last decade, both topics have increasingly gained the attention of geoscientists and the society, but the main geoethical dilemmas related to the conservation and management of geoheritage are not clearly identified yet. This work aims at providing an overview on the meaning and scope of geoethics and how it intersects geoheritage and the practice of geoconservation. Some case studies—many of which are under current debate and have a high potential as geoeducational resources—are presented for addressing ethical, social and cultural settings as well as dilemmas affecting geoheritage. We find that there are particular cases (mostly concerning the trade of fossils, and in particular the growing concern about activities that rely on amber from Myanmar) for which a clear dichotomy of views makes them much more problematic and complex. These cases deserve more suitable legal frameworks that help implement more balanced ethical standards and practice guidelines for geoconservation, guarantee human rights and needs in relation to that heritage and contribute to the advancement of geosciences. Particular attention is given to palaeontological heritage, as fossils are among the most threatened elements of the Earth’s diversity and are in need of more effective and statutory protection measures. In the context of geoethics applied to palaeontological heritage, and given the need of a clear understanding of what ethics in palaeontology means, a new concept—palaeontoethics—is proposed and formally defined., This work was supported by the Government of Aragon (Reference Group E33_17R, co-financed with FEDER 2014-2020 `Building Europe from Aragon', and Reference Group E33_20R), and the Spanish Ministry of Economy and Competitiveness and FEDER funds (PGC2018-093890-B-I00 and PID2019-105537RBI00). The study has been performed within the framework of the GOAL (Geoethics Outcomes and Awareness Learning) project (ref. 2017-1-PTO1-KA203-035790).

Small to medium-sized mustelids from the last 18 million years represent a heterogeneous group of carnivorans with a wide-ranging record in the northern hemisphere. They were first referred to the genera Mustela and Martes, but lately ascribed to the latter, and hence considered as the longest-lived genus within Mustelidae. However, a great many of these forms have been based upon fragmentary material and Martes has conformed progressively to a wastebasket nomen for species of uncertain relationships. Here, we describe dentognathic material of a small-sized mustelid from three middle Miocene (MN7 + 8, latest Aragonian) localities of the Iberian Peninsula that constitutes a new genus and species. Aragonictis araid, gen. et sp. nov. represents a distinct taxon if compared with early/middle Miocene forms ascribed to “Martes” spp., especially the similar-sized early Miocene Circamustela? laevidens and the middle Miocene “Martes” caedoti and “Martes” delphinensis. The finding of particular features in A. araid (low p2–3, loss or reduction of the p4 accessory cuspid with its main cuspid centrally located, presence of a sharp, beveled and lingually open m1 talonid, and reduction of M1 lingual platform) indicates affinities with the late Miocene Circamustela in the range of hypercarnivory. Our reassessment of “Martes” indicates possible evidence of cladogenesis for Miocene mustelidae with, at least, two different events being recognized in Europe—the latter during MN7 + 8 to MN9 with presence of Aragonictis and Circamustela. The finding of A. araid further confirms the presence of more densely forested environments than expected in inner Iberia during the latest middle Miocene., This study was supported by the Government of Aragon (Group ref. E33_20R), the Research Group UCM 910607, the Spanish Ministry of Economy and Competitiveness and FEDER funds (Research Projects PGC2018-094122-B-100 and PID2020-116220GB-100), and the Spanish Ministry of Science, Innovation, and Universities (“Juan de la Cierva Formación”, ref. FJC2018-036669-I).

Although the initial formulation of modern concepts of adaptive radiation arose from consideration of the fossil data, rigorous attempts to identify this phenomenon in the fossil record are largely uncommon. Here I focus on direct evidence of the diet (through tooth-wear patterns) and ecologically-relevant traits of one of the most renowned fossil vertebrates-the Miocene ruminant Hoplitomeryx from the island of Gargano-to deepen our understanding of the most likely causal forces under which adaptive radiations emerge on islands. Results show how accelerated accumulation of species and early-bursts of ecological diversification occur after invading an island, and provide insights on the interplay between diet and demographic (population-density), ecological (competition/food requirements) and abiotic (climate-instability) factors, identified as drivers of adaptive diversification. A pronounced event of overpopulation and a phase of aridity determined most of the rate and magnitude of radiation, and pushed species to expand diets from soft-leafy foods to tougher-harder items. Unexpectedly, results show that herbivorous mammals are restricted to browsing habits on small-islands, even if bursts of ecological diversification and dietary divergence occur. This study deepens our understanding of the mechanisms promoting adaptive radiations, and forces us to reevaluate the role of diet in the origins and evolution of islands mammals.

Background The two main primate groups recorded throughout the European Miocene, hominoids and pliopithecoids, seldom co-occur. Due to both their rarity and insufficiently understood palaeoecology, it is currently unclear whether the infrequent co-occurrence of these groups is due to sampling bias or reflects different ecological preferences. Here we rely on the densely sampled primate-bearing sequence of Abocador de Can Mata (ACM) in Spain to test whether turnovers in primate assemblages are correlated with palaeoenvironmental changes. We reconstruct dietary evolution through time (ca. 12.6–11.4 Ma), and hence climate and habitat, using tooth-wear patterns and carbon and oxygen isotope compositions of enamel of the ubiquitous musk-deer Micromeryx. Results Our results reveal that primate species composition is strongly correlated with distinct environmental phases. Large-bodied hominoids (dryopithecines) are recorded in humid, densely-forested environments on the lowermost portion of the ACM sequence. In contrast, pliopithecoids inhabited less humid, patchy ecosystems, being replaced by dryopithecines and the small-bodied Pliobates toward the top of the series in gallery forests embedded in mosaic environments. Conclusions These results support the view that pliopithecoid primates preferred less humid habitats than hominoids, and reveal that differences in behavioural ecology were the main factor underpinning their rare co-occurrence during the European Miocene. Our findings further support that ACM hominoids, like Miocene apes as a whole, inhabited more seasonal environments than extant apes. Finally, this study highlights the importance of high-resolution, local investigations to complement larger-scale analyses and illustrates that continuous and densely sampled fossiliferous sequences are essential for deciphering the complex interplay between biotic and abiotic factors that shaped past diversity.

The geosciences have experimented recently the urgent necessity to count on practitioners who possess an ethical conscience and the desire to act responsibly and serve the society. This is especially necessary in the case of our paleontological heritage. Fossils are natural objects resulting from natural processes that connect us with our natural environments and, also importantly, with our origins and past. The paleontological heritage is therefore strongly linked with our natural, social and cultural heritage, and cannot be interpreted or studied without this synergetic perspective. Transmitting paleontological knowledge to geoscientists (including educators) and authorities, especially based on the findings from the fossil record, must be pivotal in order to ensure adequate protection and conservation of the paleontological heritage, promote responsible research practices and attract attention by society.An ethical and correct management of the paleontological heritage often raise key ethical concerns. There are a range of useful examples concerning i) the increasing use of technological advances and an ambitious development of infrastructures (e.g., mining activities and exploitation of georesources, railroad, highway and residential projects, etc.) often initiated, funded, and influenced by government agencies or public and/or private organizations; ii) individual actions to collect the most spectacular, relevant fossils related to both commercial or collecting, or simple vandalism; and iii) the increasing use of fossils in paleontological research, didactic and touristic activities and exhibitions—and its profound impact on sites and fossils, that relate to our paleontological heritage and can thus foster personal growth, enrich citizens´ knowledge and promote (and improve) interaction between society and this field of geosciences.More specifically, in Spain there are examples and case studies in the field of geoscience that illustrate how to gain awareness and knowledge about geoethical conflicts between works of infrastructures, and the geoconservation of paleontological sites and the discovery of new fossils, with benefits for society, the administration, and the scientific community.Identifying and considering ethical issues and dilemmas associated to these topics in paleontology are important for both ethical (adhering to general principles or conceptions of what is right and wrong) and practical reasons. As such, geoscientists who are more aware of their ethical and social responsibilities concerning our paleontological heritage will be better able to put their knowledge at the service of society and to help increase public trust in geosciences.

The intra-montane Guadix-Baza Basin is one of the few continental basins in Europe that hosts a well-dated set of fossiliferous sites spanning from the latest Miocene to the late Middle Pleistocene. The Cúllar de Baza 1 (CB-1) represents a key site to investigate the effects of the Early-Middle Pleistocene Transition, considered a fundamental transformation in the Earth’s climate state. Our review and update of the large mammal assemblage, and particularly equids, is of paramount relevance to understand the systematic affinities and the evolution of the Early and Middle Pleistocene European horses. We confirm the occurrence of two different taxa, the medium sized Equus altidens and the larger E. suessenbornensis. Moreover, we illustrate that CB-1 is essential for the biochronological studies of the latest Early Pleistocene/Middle Pleistocene transition (Epivillafranchian/Galerian ELMA); in particular with regard to the Last Occurrences of the Etruscan rhino Stephanorhinus etruscus and the large deer Megaloceros savini and the First Occurrence of the water-rat Arvicola mosbachensis in the Iberian peninsula. Finally, a development of a mosaic environment characterised the CB-1 site contrasting with the conditions reported for other Iberian late Early and Middle Pleistocene localities., This research received support from the SYNTHESYS Project ES-TAF-5621 http://www.synthesys.info/ which is financed by European Community Research Infrastructure Action under the FP7 «Capacities » Programme at Museo Nacional de Ciencias Naturales (CSIC) and from Project CGL 2016-79334-P (Dirección General de Ciencia y Tecnología, Spain).

The Valles-Penedes Basin (Catalonia, Spain) is classical area for the study of Miocene land mammal faunas. Nevertheless, the early Miocene part of the record has deserved little attention as compared to younger intervals. Most notably, the small mammals of this age have not been described in detail, consequently hampering the correlation of the Valles-Penedes record with other chronological schemes. In this work we describe the rich and diverse small mammal fauna from Les Cases de la Valenciana site (Gelida, Alt Penedes) which includes marsupials, eulipotyphlans, lagomorphs and rodents. On the basis of the presence of the cricetids Megacricetodon and Democricetodon this site is correlated with European Neogene zone MN4, yielding an age of 17–16 Ma. However, the rodent assemblage is comparable to that of chronologically close localities of the Calatayud-Montalban Basin (Aragon, Spain), indicating that the same biochronological scheme can be applied to both areas. In this way, the coexistence of the eomyids Ligerimys ellipticus and Ligerimys florancei coupled with the presence of Megacricetodon primitivus indicates a correlation with local biozone Ca of the Calatayud-Montalban Basin, ranging from 16.3 to 16 Ma. Finally, the correlation of other early Miocene sites of the Valles-Penedes Basin is discussed and refined.

Climatic oscillations at the Gelasian/Calabrian transition modified terrestrial palaeoenvironmental settings in the European region. A gradual drop in global temperatures beginning about 2.7 Ma led to drier conditions and to a reduction in, and subsequent disappearance of, sub-tropical vegetation in the central Mediterranean area by ca 1.2 Ma. Large ungulates are sensitive to vegetation changes and faced with harsher environmental settings may shift their feeding strategies to exploit available food resources in different ecosystems. In fossil assemblages such dietary adaptations are reflected by tooth morphology (a phylogenetic signal) and tooth wear degree (a direct signal of the species' diet). In this paper, we investigate how large herbivores responded to palaeoenvironmental changes that occurred at the passage between the Gelasian and Calabrian ages in the Italian Peninsula, analysing the dental wear patterns and hypsodonty of the early Pleistocene fossil ungulates assemblage of Olivola (Aulla, Central Italy). We found that while ungulate feeding behaviours during the Gelasian spanned from browsers to grazers, in the locality of Olivola this group of herbivores display a narrower range of diet types with many taxa adopting a mixed feeding behaviour. Cervids in particular, whose fossils are often associated only with wooded environments, as a response to the reduction of covered sub-tropical vegetation, shifted from a strict browsing diet in mostly closed habitats to a more abrasive one taking advantage of the spread of open landscapes. We also provide new data on the feeding behaviour of the rare fossil caprines, Procamptoceras and Gallogoral meneghinii, suggesting a grass-rich mixed diet for the former and a certain degree of dietary plasticity for the latter. Our research thus contributes to the better understanding of how ungulates adapted in the past to exploit different resource types during pivotal climatic changes and how environments changed in Central Italy on the onset of colder and more arid conditions.

The Early-Middle Pleistocene Transition (EMPT) (ca. 1.4–0.4 Ma) represents a fundamental transformation in the Earth's climate state, starting at 1.4 Ma with a progressive increase in the amplitude of climatic oscillations and the establishment of strong asymmetry in global ice volume cycles. The progressive shift from a 41kyr–100kyr orbital rhythm was followed by the first major build-up of global ice volume during MIS 24-22, the so-called “0.9 Ma event”. The Vallparadis Section (Valles-Penedes Basin, NE Iberian Peninsula) is one of the few Pleistocene series in Europe that spans the onset of the transition (from 1.2 to 0.6 Ma), thus representing a pivotal array of localities to investigate the effect of glacial dynamics on environmental conditions in Southern Europe. Here we inspect the effects of the EMPT on terrestrial ecosystems by examining the dietary adaptations (through dental meso- and microwear patterns) of fossil ungulates from the Vallparadis Section dated before and after the “0.9 Ma event”. Results show a steady presence of open grasslands before MIS 22 and more humid conditions at MIS 21. Both before and after MIS 22, a consistent presence of ungulates with long-term patterns that point to a grazing or grass-rich mixed feeding behaviour is observed, while noticeably, short-term patterns point to increased seasonality right after the “0.9 Ma event” glacial period. This increment of seasonality may have had an important effect on the Mediterranean habitats leading to recurring changes in the quality of plant resources available to large herbivores, which in response periodically adopted more mixed feeding behaviours widening their dietary breadth to consume also sub-optimal food items during adverse seasons.

We describe new specimens of the Miocene moschid Hispanomeryx, from the early Vallesian sites of Castell de Barberà (CB) and Ecoparc de Can Mata (ECM), Vallès-Penedès Basin, representing the first Iberian record of Hispanomeryx outside the inner Miocene basins. Fossils from ECM constitute Hispanomeryx lacetanus, sp. nov., the first Hispanomeryx to be described with a ‘Palaeomeryx fold (external postprotocristid).’ Fossils from CB constitute the first unambiguous record of Hispanomeryx aragonensis outside its type locality (La Ciesma-1B, Ebro Basin, Spain) and allow us to describe for the first time the upper molars of this species. Our phylogenetic analysis of Hispanomeryx places H. andrewsi from China as the sister group of a clade containing two European lineages. One of the latter places H. daamsi in a clade with the sister species H. duriensis and H. lacetanus, sp. nov., and the other groups together the two forms of H. aragonensis (type series from La Ciesma-1B plus the new material from CB). Emended diagnoses for the genus Hispanomeryx and all previously described species are presented. The new Hispanomeryx material from the Vallès-Penedès Basin constitutes the first unambiguous forest-dwelling Hispanomeryx ever recorded in Iberia, demonstrating that the previous lack of Hispanomeryx fossils in the Vallès-Penedès Basin was due to insufficient sampling rather than to paleoecological absence. It is therefore apparent that Hispanomeryx was as environmentally plastic as its relative Micromeryx, inhabiting similar habitats that varied from dry and open to humid and forested., This work was supported by the Spanish Agencia Estatal de Investigación (Ministerio de Economía, Industria y Competitividad) European Regional Development Fund of the European Union (CGL2015-68333-P, MINECO/FEDER-UE; CGL2016-76431-P; and CGL2016-76431-P, AEI/FEDER EU), the Generalitat de Catalunya (CERCA Programme, 2017 SGR 86 GRC, and 2017 SGR 116 GRC), the Departament de Cultura of the Generalitat de Catalunya (field work grant 2014/100609), and the Leakey Foundation.

EgeUn###, Lophocyonids are one of the more enigmatic groups of Carnivora in the European Miocene fossil record. Lophocyonids are clearly distinguished from other Feliformia by their peculiar lophodont dental morphology. For this reason, the systematic relationships of the family have been controversial. Here we describe and interpret dental remains from the early Miocene of Sabuncubeli, Turkey, which we attribute to a new genus and species: Izmirictis cani. The phylogenetic analysis allows us to include Izmirictis within a monophyletic group, Lophocyonidae new rank, characterized by the molarization of the anterior premolars (P3 and p4), the lophodont adaptation of the molar dentition and the complex morphology of the incisors. The phylogenetic analysis shows a close relationship between Izmirictis and primitive hyaenids (represented herein by Protictitherium). The divergence between Lophocyonidae and Hyaenidae is estimated by biochronological data to have occurred during the early Miocene (MN2). Dental microwear analysis, although limited by poor dental enamel preservation, indicates that the pronounced lophodonty in Izmirictis cani could be connected to a herbivorous feeding habit., Spanish Research Project (MINECO/FEDER-UE) [CGL2015-68333-P]; AEI/FEDER EU [CGL2016-76431-P]; EGE University Research Grants [2004/BIL/020, 2005/TTM/001, 2016/TTM/001, 2016/TTM/002, 2015/Fen/17]; TUBITAK Research GrantTurkiye Bilimsel ve Teknolojik Arastirma Kurumu (TUBITAK) [YDABAG 102Y045]; DST-NFR Centre of Excellence in Palaeosciences (CoE-Pal) [COE2018-09POST]; Research Group CSIC [641538]; Research Group CAM-UCM [910607], We thank Hans de Bruijn (Utrecht University) and Oldrich Fejfar (Charles University in Prague) for providing casts of some lophocyonid specimens from the Aliveri and Rothensteim localities, respectively. Thanks to Susana Fraile (Museo Nacional de Ciencias Naturales, CSIC), Marta March (Institut Catala de Paleontologia Miquel Crusafont), Pip Brewer (Natural History Museum, London) and Christine Argot (Museum national d'Histoire naturelle) for access to comparative material in their care. The Spanish Research Project CGL2015-68333-P (MINECO/FEDER-UE) and the Research Groups CSIC 641538 and CAM-UCM 910607 funded this research. We also thank the AEI/FEDER EU (CGL2016-76431-P) for support. Serdar Mayda and Tanyu Kaya were funded by EGE University Research Grants 2004/BIL/020, 2005/TTM/001, 2016/TTM/001 and 2016/TTM/002, and 2015/Fen/17 and the TUBITAK Research Grant YDABAG 102Y045. The support of the DST-NFR Centre of Excellence in Palaeosciences (CoE-Pal) towards this research for Alberto Valenciano (COE2018-09POST) is hereby acknowledged. Opinions expressed and conclusions arrived at are those of the authors and are not necessarily to be attributed to the CoE. Special thanks to Martin Pickford for improving the English.We are also indebted to the Editors-in-Chief P. Barret and P. D. Taylor and two anonymous reviewer for their useful comments and suggestions, which improved the original manuscript.

Deer (Cervidae) have a long evolutionary history dating back to the Early Miocene, around 19 million years ago. The best known fossils to document this history belong to European taxa, which all bear cranial appendages more or less similar to today's deer antlers. Despite the good fossil record, relationships of the earliest stem deer and earliest crown deer are much debated. This hampers precise calibration against the independent evidence of the fossil record in molecular clock analyses. While much has been written on the Early and Middle Miocene deer, only two phylogenetic analyses have been performed on these taxa to date mostly based on cranial appendage characters. Because the petrosal bone and bony labyrinth have been shown to be relevant for phylogeny in ruminants, we describe for the first time these elements for four iconic early cervids from Europe (Procervulus dichotomus, Heteroprox larteti, Dicrocerus elegans and Euprox furcatus) and include them in a phylogenetic analysis based on the ear region exclusively. The analysis recovered E. furcatus in a sister position to the living red deer (Cervus elaphus). Further, it placed D. elegans in a sister position to Euprox+Cervus and a clade Procervulinae that includes P. dichotomus and H. larteti, in sister position to all other deer. The inclusion of E. furcatus in crown Cervidae, which was previously suggested based on antler morphology, cannot be ruled out here but needs a more comprehensive comparison to other crown deer to be confirmed. J. Morphol. 277:1329-1338, 2016. (c) 2016 Wiley-Blackwell.

Deer (Cervidae) have a long evolutionary history dating back to the Early Miocene, around 19 million years ago. The best known fossils to document this history belong to European taxa, which all bear cranial appendages more or less similar to today's deer antlers. Despite the good fossil record, relationships of the earliest stem deer and earliest crown deer are much debated. This hampers precise calibration against the independent evidence of the fossil record in molecular clock analyses. While much has been written on the Early and Middle Miocene deer, only two phylogenetic analyses have been performed on these taxa to date mostly based on cranial appendage characters. Because the petrosal bone and bony labyrinth have been shown to be relevant for phylogeny in ruminants, we describe for the first time these elements for four iconic early cervids from Europe (Procervulus dichotomus, Heteroprox larteti, Dicrocerus elegans and Euprox furcatus) and include them in a phylogenetic analysis based on the ear region exclusively. The analysis recovered E. furcatus in a sister position to the living red deer (Cervus elaphus). Further, it placed D. elegans in a sister position to Euprox + Cervus and a clade Procervulinae that includes P. dichotomus and H. larteti, in sister position to all other deer. The inclusion of E. furcatus in crown Cervidae, which was previously suggested based on antler morphology, cannot be ruled out here but needs a more comprehensive comparison to other crown deer to be confirmed. J. Morphol. 277:1329-1338, 2016. © 2016 Wiley Periodicals, Inc.

Deer are an iconic group of large mammals that originated in the Early Miocene of Eurasia (ca. 19 Ma). While there is some consensus on key relationships among their members, on the basis of molecular- or morphology-based analyses, or combined approaches, many questions remain, and the bony labyrinth has shown considerable potential for the phylogenetics of this and other groups. Here we examine its shape in 29 species of living and fossil deer using 3D geometric morphometrics and cladistics. We clarify several issues of the origin and evolution of cervids. Our results give new age estimates at different nodes of the tree and provide for the first time a clear distinction of stem and crown Cervidae. We unambiguously attribute the fossil Euprox furcatus (13.8 Ma) to crown Cervidae, pushing back the origin of crown deer to (at least) 4 Ma. Furthermore, we show that Capreolinae are more variable in bony labyrinth shape than Cervinae and confirm for the first time the monophyly of the Old World Capreolinae (including the Chinese water deer Hydropotes) based on morphological characters only. Finally, we provide evidence to support the sister group relationship of Megaloceros giganteus with the fallow deer Dama.

Resource competition and niche partitioning among the exceptionally high number of sympatric ungulates of the Early Pleistocene site of Coste San Giacomo (Central Italy) is here examined through the study of their dietary proclivities and body size. The main aim of this study is to investigate the niche differentiation mechanisms that let the fossil ungulates coexist in the same region. We also provide information about the complementarity of two different methodologies that observe diet variation at a different time scales (inner and outer mesowear) in the study of dental wear patterns of fossil ungulates. Results from analyses of dental wear degree and body masses predictions show that a wide range of feeding behaviours were adopted by the taxonomical groups (i.e., cervids, bovids and equids) in order to avoid competition. Among larger ungulates diet ranges from strict browsing (Eucladoceros sp., Gazellospira torticornis), to mixed feeding (Gallogoral meneghinii, Leptobos sp.) to pure grazing (Equus stenonis), whereas smaller taxa are more selective feeders (Axis cf. lyra, Croizetoceros cf. ramosus) with only one exception (Gazella borbonica). When taxa with the same feeding behaviour occurred in the same habitat, competition was minimised by differences in body size.

Mastication of dietary items with different mechanical properties leaves distinctive microscopic marks on the surface of tooth enamel. The inspection of such marks (dental microwear analysis) is informative about the dietary habitus in fossil as well as in modern species. Dental microwear analysis relies on the morphology, abundance, direction, and distribution of these microscopic marks. We present a new freely available software implementation, MicroWeaR, that, compared to traditional dental microwear tools, allows more rapid, observer error free, and inexpensive quantification and classification of all the microscopic marks (also including for the first time different subtypes of scars). Classification parameters and graphical rendering of the output are fully settable by the user. MicroWeaR includes functions to (a) sample the marks, (b) classify features into categories as pits or scratches and then into their respective subcategories (large pits, coarse scratches, etc.), (c) generate an output table with summary information, and (d) obtain a visual surface-map where marks are highlighted. We provide a tutorial to reproduce the steps required to perform microwear analysis and to test tool functionalities. Then, we present two case studies to illustrate how MicroWeaR works. The first regards a Miocene great ape obtained from through environmental scanning electron microscope, and other a Pleistocene cervid acquired by a stereomicroscope. © 2018 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

The Middle Pleistocene site of Fontana Ranuccio (Anagni Basin, Central Italy) has yielded nearly 25,000 remains belonging to large and small vertebrates, including four isolated teeth of Homo sp., and a rich lithic assemblage containing bone tools. Here we provide new dental wear analyses of the ungulate community to improve the palaeoenvironmental reconstruction for the site. Analyses indicate a mostly browsing diet for the cervids and a grazing diet for the perissodactyls Stephanorhinus sp. and Equus cf. E. mosbachensis, suggesting a range of habitats that spanned from forests to open grasslands. A discrepancy between mesowear and microwear results is observed in the case of the bovid, Bos primigenius; this can be tentatively explained as resulting from a temporal switch towards a sub-optimal diet, possibly reflecting the effects of marked seasonality. Findings shed new light on the structure of the 400 kyr ecosystems that existed during the early occupation of Homo in Europe.

Quaternary glacial/interglacial alternations, influenced by orbital obliquity cycles with a 41-ka long periodicity, started in the northern hemisphere around 2.6 Ma ago. Such alternations affected the terrestrial ecosystems, especially those of the Mediterranean region, with changes in the floristic communities and the dispersal and radiation of a number of large mammal open dwellers. Analyses of tooth wear patterns of ungulates from the Early Pleistocene site of Coste San Giacomo allow for a more objective reconstruction on the paleoenvironments and the climate in the Italian Peninsula during this epoch. Our results show that this area was composed by a mosaic of biomes, in particular by steppe and woodlands/wetlands. Evidence of such heterogeneity is provided by the wide spectrum of feeding behaviours found among the numerous ungulate herbivores here recorded, with cervids (Axis cf. lyra, Croizetoceros cf. ramosus and Eucladoceros sp.) exhibiting browser diets, most of the bovids (Gazella borbonica and Leptobos sp. and Gallogoral meneghinii) being intermediate feeders and the equid Equus stenonis showing a strict grazer behaviour. These results provide new insights for a timing of changing ecosystems in Southern Europe and reveal the environmental legacy of this global climatic shift, which is essential for understanding the early occupation of Homo in Europe. Thus, our data provide new evidence that such an environmental heterogeneity and a wide spectrum of available food resources could have been the main factors favouring the settlement of early species of Homo in this area.

Despite its long history of scientific study, the causes underlying the extinction of the insular hominoid Oreopithecus bambolii are still a matter of ongoing debate. While some authors consider intense tectonism and invading species the cause of its extinction ca. 6.7 Ma, others propose climatic change as the main contributing factor. We rely on long-term patterns of tooth wear and hypsodonty of the Baccinello and Fiume Santo herbivore-faunas to reconstruct changes in habitat prior to, during and after the extinction. While a mosaic of habitats was represented in Baccinello V1 (as shown by a record of browsers, mixed feeders and species engaged in grazing), more closed forests (higher proportion of browsers, shortage of mixed feeders and lack of grazers) characterised Baccinello V2. Finally, there was a partial loss of canopy cover and development of open-patches and low-abrasive grasses in Baccinello V3 (as denoted by new records of taxa involved in grazing)—although still dominated by a forested habitat (since browse was a component in all diets). Our results provide evidence for two perceptible shifts in climate, one between 8.1 and 7.1 Ma and other ca. 6.7 Ma, though this latter was not drastic enough to lead to intensive forest loss, substantially alter the local vegetation or affect Oreopithecus lifestyle—especially if considering the growing evidence of its versatile diet. Although the disappearance of Oreopithecus is complex, our data reject the hypothesis of environmental change as the main factor in the extinction of Oreopithecus and Maremma fauna. When our results are analysed together with other evidence, faunal interaction and predation by invading species from mainland Europe seems to be the most parsimonious explanation for this extinction event. This contrasts with European hominoid extinctions that were associated with major climatic shifts that led to environmental uniformity and restriction of the preferred habitats of Miocene apes.

New tragulid dental remains from the late Early Miocene (16.5–16.3 Ma, MN4) locality of Els Casots (Valles-Penedes Basin, Catalonia, Spain) are described. This sample fits well, both in size and occlusal morphology, with the material of Dorcatherium crassum from the type locality (Sansan, France; MN6). We therefore attribute the tragulid from Els Casots to this species, also in agreement with its known chronostratigraphic range throughout Europe (MN4–MN6) and the previous records of this species from other MN4 localities of the Valles-Penedes Basin. The described remains – which include the postcanine lower deciduous and permanent dentition, as well as several upper cheek teeth – constitute the most complete sample of D. crassum from the Iberian Peninsula. The presence of this species at Els Casots is consistent with the lacustrine depositional environment inferred from sedimentological evidence and associated fauna, and further confirms the nearby presence of densely forested environments with a humid climate with low seasonality.

The mammalian fossil record of Spain is long and taxonomically well resolved, offering the most complete record of faunal change for the Neogene of Europe. We evaluated changes in diversification, composition, trophic structure, and size structure of large mammals over the middle and late Miocene with methods applied to this record for the first time, including ordination of fossil localities to improve temporal resolution and estimation of confidence intervals on taxa temporal ranges. By contrast, analysis within the traditional Mammal Neogene (MN) biochronology obscures important aspects of diversification. We used inferred temporal ranges of species and evaluated per capita rates of origination, extinction, diversification, and turnover over 0.5-Myr time intervals.Three periods of significant faunal change occurred between 12.0 and 5.5 Ma: (1) From 12.0 to 10.5 Ma, elevated origination rates led to an increase in diversity without significant change in ecological structure. Immigrants and geographic-range shifts of species to lower latitudes during an interval of global cooling contributed to these faunal changes. (2) From 9.5 to 7.5 Ma, high extinction rates followed by high origination rates coincided with significant changes in taxonomic composition and ecological structure. These changes represent the Vallesian Crisis, with replacement of a fauna of forest affinities (with frugivores and browsers) by a fauna of open woodlands (with grazers and mixed feeders). (3) From 6.5 to 5.5 Ma, high extinction rates reduced diversity without substantial changes in ecological structure, and large mammal faunas became highly endemic across the northern Mediterranean region. This interval includes the Messinian Salinity Crisis, the desiccation of the Mediterranean basin. Extinction may have been caused by geographic isolation and aridification, with evolution of endemic lineages giving rise to new species in the early Pliocene. These distinct macroevolutionary patterns of faunal change correspond to different geographic scales of inferred climatic and tectonic drivers.

Pliopithecinae and Crouzeliinae (Primates: Pliopithecidae) are distinguished dentally by the sharper crests, more compressed cusps, larger foveae, and narrower molars of the latter. Traditionally, such differences were qualitatively related to increased folivory in crouzeliines. This was subsequently disproved by microwear and shearing crest analyses, indicating that all pliopithecids were soft-fruit eaters, except for the more folivorous crouzeliine Barberapithecus. This seems however at odds with the occlusal morphology of the latter, intermediate between those of Pliopithecus and the more derived crouzeliine Anapithecus. To further assess dietary evolution in this group, we report results of dental microwear for two Iberian pliopithecids: Pliopithecus canmatensis, from several Abocador de Can Mata localities (11.8–11.7 Ma, MN8), and Barberapithecus huerzeleri from Castell de Barbera (ca. 11.2–10.5 Ma, MN8, or MN9). Contrary to previously published results, our analyses suggest that all pliopithecids, including Barberapithecus, had a frugivorous diet with a significant sclerocarpic component—apparently more marked in some pliopithecines (such as P. canmatensis) than in the crouzeliine Anapithecus. If our interpretation is correct, it would mean that, within the framework of a frugivorous diet with some hard-object feeding, crouzeliine dental evolution would have been driven by selection pressures towards increased soft-fruit consumption instead of folivory. Dental differences between pliopithecids and hominoids with a significant sclerocarpic component (i.e., orangutans) might be related to phylogenetic constraints, different food-processing methods and/or fracture toughness of hard foods consumed. Although additional research would be required, results suggest that dietary niche partitioning played a significant role in the radiation of European pliopithecids. Am J Phys Anthropol 151:573–582, 2013. © 2013 Wiley Periodicals, Inc.

[EN] The land mammal record of the Vallès-Penedès Basin (Catalonia, NE Spain) ranges from the early Miocene (Ramblian) to the late Miocene (Turolian), that is from about 20 to 7 Ma. Here we present an updated review of the mammal succession focusing on biochronology as well as on environmental and faunal changes. Based on faunal similarities with central Europe, we interpret this basin as a transitional zone between the forested environments of northern regions and the more arid landscapes of the inner Iberian Peninsula. The quality of the Vallès-Penedès record and its chronostratigraphic control is clearly better for the late Aragonian and the Vallesian (between 12.6–9.0 Ma), especially for small mammals. Therefore, we analyze small mammal diversity dynamics during this interval. Contrary to previous analyses, which found an abrupt extinction event coinciding with the early/late Vallesian boundary (the Vallesian Crisis), our results show that this pattern is due to uneven sampling. Instead, taxonomic richness slowly decreased since the late Vallesian as a result of a series of extinctions that mostly affected forest-dwelling taxa., [FR] Le registre fossile des mammifères du bassin de Vallès-Penedès (Catalogne, Nord-Est de l’Espagne) s’étend du Miocène inférieur (Ramblien) jusqu’au Miocène supérieur (Turolien), soit environ de 20 à 7 Ma. Nous présentons ici une révision actualisée des successions de mammifères, en prêtant attention aux aspects biochronologiques, environnementaux et faunistiques. Sur la base des similarités observées avec les faunes d’Europe centrale, nous interprétons ce bassin, d’une part, comme une zone transitionnelle entre les environnements boisés des régions nordiques et, d’autre part, comme le paysage le plus aride de la partie intérieure de la péninsule Ibérique. La qualité du registre fossile du Vallès-Penedès, ainsi que son contrôle chronostratigraphique, sont nettement supérieurs pour la partie Aragonien supérieur et Vallésien (entre 12,6 et 9,0 Ma), particulièrement pour les petits mammifères. Pour cette raison, nous analysons ici la dynamique de la diversité des petits mammifères pour cette période. À l’inverse des analyses précédentes, qui observaient une extinction brutale à la limite Vallésien inférieur/supérieur, nos résultats montrent que ce modèle résulte en fait d’un d’échantillonnage inégal. Nous proposons plutôt une diminution lente de la richesse taxonomique à partir du Vallésien supérieur, conséquence d’une série d’extinctions touchant, en priorité, les espèces forestières., This work has been supported by the Spanish Ministerio de Economía y Competitividad (CGL2011-28681, CGL2010-1749 and RYC-2013-12470 to ICV), the Agència de Gestió d’Ajuts Universitaris i de Recerca (2014 SGR 416 and 2014 SGR 467) and the Departament de Cultura (2014/100584 and 2014/100609) of the Generalitat de Catalunya. IG-P acknowledges a JAE-Doc contract (CSIC) co-funded by the European Social Fund and Research group UCM-910607 “Evolution of Cenozoic Mammals and Continental Paleoenvironments”. AM acknowledges the support of an Erasmus Staff Training mobility grant by Erasmus+ (Life Long Learning Programme). We deeply appreciate the comments and constructive critics by the two reviewers of this manuscript, J.C. Barry and R.L. Bernor, as well as by the guest editor, L.J. Flynn. We thank Jérôme Prieto (Ludwig Maximilians University Munich) for his help with the French translation of the abstract and figure captions.

Mammals of numerous lineages have evolved high‐crowned (hypsodont) teeth particularly during the last 20 million years. This major phenotypic change is one of the most widely studied evolutionary phenomena in a broad range of disciplines, though the mechanisms underlying its transformation remain unresolved. Here, we present the first Finite Element Analysis (FEA) to investigate the alternative hypothesis that there is a biomechanical link between increased hypsodonty and a more effective mastication in deer. Our FE experiments compared patterns of stress and strain within and between different fossil and living species under different loading conditions, and found that more hypsodont teeth are suited for restricting stresses to those areas where chewing loading occurs. This mechanical improvement is consequence of specific and pronounced variations in tooth geometry and morphology of the occlusal surface that are strongly related to crown growth in the vertical plane. We demonstrate that hypsodonty enables selenodont‐teeth to adopt a mechanically improved design that increases the pressure whilst shearing foods. As ruminants are physiologically limited by both the quantity of food consumed and the time spent in the mastication and digestion, hypsodonty is highly advantageous when feeding on mechanically resistant, tough and fibrous foods. Consequently, it allows grass‐eaters to spend less time chewing, thereby increasing the volume of food ingested and/or providing more time for digestion. This study provides a promising line of evidences in support of biomechanical effectiveness, in addition to or instead of increased wear resistance, as a factor in explaining the evolutionary origins of the hypsodont phenotype., his study was funded by the Spanish Ministerio de Economía y Competitividad (projects CGL2010‐216723, CGL2010‐19116 and CGL2011‐25754, and JCI‐2011‐11697 to D.D.M.), the Research Group BSCH‐UCM910607 and the government of Aragón (E05).

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) ...

Dietary behaviour and competition for resources are investigated for the small-sized ruminants Andegameryx Ginsburg, 1971 and Procervulus Gaudry, 1877 representatives of two largely distinct states of diversification of pecorans. Results obtained from dental microwear and mesowear methodologies are concordant with a mixed feeder strategy for the taxa from the Early Miocene environments of the Iberian Chain (Central Spain). Further, the Spanish taxa investigated had less abrasive diets than their relatives from others similarly aged localities in Europe. This fact raises an important evolutionary uncertainty concerning the traditional characterization of first pecorans as specialized browsers. Instead, data strongly corroborate the recently proposed notion that some Pecora ruminant lineages were able to consume a mixture of browse and grass, and that were originally facultative mixed feeders. However, there is a large degree of variation of their dental wear features. Species were not dependent on...

Family Pliobatidae fam. nov. Type genus: Pliobates gen. nov., whose diagnosis is as for its type (and only) species, described below.

Holotype: IPS58443, a partial skeleton with an associated skull (Fig. 1 and movie S1), housed at the ICP. It is composed of 70 bones and bone fragments (table S1) found in close spatial association, which, given the lack of repeated elements, are attributed to a single adult female individual (based on the small canine alveolus), with an estimated body mass of 4 to 5 kg (tables S7 and S8). It includes large portions of the cranium with postcanine maxillary teeth (Table 1), a mandibular fragment, a partial left forelimb (nearly complete humerus, radius, partial ulna, carpals, and bones of the manual rays), more fragmentary elements of the right forelimb, and bones from the hind limb. Type locality: ACM/C8-A4 (els Hostalets de Pierola, Catalonia, Spain), in the ACM stratigraphic series (Vallès-Penedès Basin, northeast Iberian Peninsula). Age, stratigraphic position, and distribution: Only known from the type locality, which has an estimated age of 11.6 Ma (middle/late Miocene boundary) and is thus somewhat younger than all other ACM hominoid- and pliopithecoid-bearing localities (9, 31, 33, 77, 78), the latter of which have been dated to 11.7 to 11.9 Ma [updated from (77, 78)]. Etymology: Genus name from the Latin plio- (itself from the Greek, meaning “greater in extent”) and from the Greek bates (meaning “the one that walks or haunts”). The name is a contraction of the genus names Pliopithecus (“more ape”) and Hylobates (“the one that walks in the woods or in the trees”), in allusion to the small body size and the mosaic of primitive (stem catarrhine–like) and derived (crown hominoid) features displayed by the new taxon. The species epithet is the genitive of the female substantive “Catalonia,” the Latin name of Catalunya (in which the type locality is situated). Small-bodied catarrhine primate (estimated female BM of 4 to 5 kg). Dental formula 2.1.2.3. Female upper canines moderately compressed. Upper cheek teeth low-crowned and with subpyramidal, moderately peripheral, and inflated cusps. Upper premolars relatively broad and ovoid, P4 smaller than P3, both with heteromorphic cusps, a markedly convex lingual contour and a distinct lingual cingulum (more developed in the P4), a distinct transverse crest separating the restricted mesial fovea from the extensive trigon basin, and the postparacrista forming an abrupt angle with the distal marginal ridge. Upper molars only moderately broader than long, with markedly convex lingual profiles; buccal cusps quite peripheral and buccal cingula discontinuous; lingual cingula relatively well developed, shelf-like, and C-shaped, but not surrounding the hypocone (which is distinct and more peripheral than the protocone); mesial fovea restricted, with an obliquely directed preprotocrista, and trigon basin extensive, being separated by a continuous crista obliqua from the slightly smaller distal fovea, which displays no hypocone-metacone crest. M2 slightly larger than the M1, and M3 shorter and trapezoidal (due to the oblique buccal margin, with a centrally situated metacone and a rudimentary hypocone). Face small but with a distinct snout, the anterior portion of the nasals being almost parallel to the palate. Maxillary sinus large and frontal sinus present but small. Nasal aperture narrow. Nasoalveolar clivus short, with an open palatine fenestra. Anteriorly slightly narrow palate with somewhat convergent upper tooth rows. Zygomatic root moderately high. Orbits subcircular, large, and frontated, with telescopic orbital rims located over the P4. Estimated cranial capacity (69 to 75 cm3) indicating a monkey-like degree of encephalization. External auditory meatus tubular but short and not completely ossified, with a V-shaped end and its anterior portion fused with the postglenoid process. Carotid foramen perforating the bulla posterodistally, and carotid canal horizontally and anteriorly oriented. Spinosum and postglenoid foramina absent. Jugular foramen large and ventrally visible. Humerus without entepicondylar foramen and capitular tail, with a well-developed capitulum, and a narrow and deep zona conoidea.Radial head rounded and not very tilted, with a markedly beveled surface for articulation with the humeral zona conoidea, the articular surface for the ulnar radial notch extending along a large portion of the radial head, and a laterally facing bicipital tuberosity. Distal radioulnar joint fully diarthrodial, with an expanded and two-faceted semilunar articulation on the ulnar head, and a partially developed ulnar fovea. Ulnar styloid process with reduced girth and not articulating with the short pisiform. Triquetrum small and with a reduced articular surface for the ulnar styloid process. Hamate relatively long proximodistally, with a steep triquetrum facet, a relatively large head and a distally projecting hamulus. Capitate with a relatively small and oblong head and a divided facet for the second metacarpal on its radial side. The new taxon differs from pliopithecoids and dendropithecids in its lack of a humeral capitular tail, its hominoid-like proximal radial morphology, its expanded ulnar head with a two-faceted semilunar articulation, and its partially developed ulnar fovea. It further differs from these taxa and proconsulids in its more hominoid-like carpal morphology (including the lack of a pisiform facet for the styloid process, a capitate facet for the second metacarpal divided by a deep ligamentary notch, and a distally projecting hamulus in the hamate), and particularly from pliopithecoids in its overall larger muzzle, more horizontal nasals anteriorly, some details of the upper molars, and (at least compared with Epipliopithecus) the lack of an entepicondylar foramen in the humerus. It also differs from all of the above-mentioned taxa in its fused ectotympanic and postglenoid process, and from these taxa and hominids in its horizontal and anteriorly oriented carotid canal. Last, it differs from crown hominoids (hylobatids and hominids) in its incompletely ossified ectotympanic and in its more primitive dentition and forelimb morphology (particularly in the humeroulnar articulation). Although the lack of lower dentition precludes comparisons with some taxa, the upper cheek teeth of Pliobates (Fig. 1 D and Table 1) generally resemble those of other small-bodied Miocene catarrhines in both occlusal morphology and proportions (figs. S1 and S2). In contrast, they display a more primitive morphology than those of extant hominoids, including the similarly sized gibbons. Hylobatids possess more elongated cheek teeth with more peripheralized cusps, less developed cingula, and a much more extensive central fovea. Compared with Miocene small-bodied catarrhines from Eurasia and Africa (fig. S1), the upper molars of Pliobates more closely resemble those of the dendropithecid Micropithecus (21, 34, 35, 79) in several features, such as the markedly convex lingual profiles and moderately developed buccal cingula (albeit to a lesser extent than in Micropithecus), the C-shaped lingual cingulum that is mostly restricted to the protocone (not surrounding the hypocone), the well-developed and lingually situated hypocone, and the relatively narrow M1 and M2. Nevertheless, Pliobates differs in several features from Micropithecus, which has more restricted buccal cingula, a hypocone-metacone crest, and a relatively longer and less trapezoidal M3. The dentition of Pliobates more clearly differs from Epipliopithecus and other pliopithecoids (fig. S1), including from Barberapithecus [also recorded at the Vallès-Penedès Basin (36)] and Pliopithecus [previously recorded at ACM (33)] in several features, such as the more convex lingual profile, the more peripheral buccal cusps, the less developed cingula of the molars, the narrower M1 and M2, and the M3 occlusal morphology and proportions (fig. S2). With regard to microwear features, the M1 displays a pitting percentage of 30.0%, a pit breadth of 5.67 μm, and a striation breadth of 1.98 μm. Based on pitting incidence (Fig. 2, A and B), which is the most useful metric for distinguishing among dietary categories (70), Pliobates closely resembles extant frugivores (Pan troglodytes) and eclectic feeders (Papio cynocephalus) that largely rely on ripe fruit. In contrast, the pitting incidence of Pliobates is higher than in extant folivores and much lower than in extant hard-object feeders (including orangutans). Compared with other extinct catarrhines from Western Europe, the pitting percentage of Pliobates is somewhat lower than in most pliopithecoids and hominoids, for which some degree of sclerocarpy has been inferred (66, 68). This low pitting incidence is consistent with the pit- and striation-breadth measurements (which show no sign of extreme folivory or specialized hard-object feeding) and most closely approaches that of the fossil hominoids Anoiapithecus brevirostris and Hispanopithecus laietanus, previously interpreted as soft frugivores (68). These results are confirmed by a multivariate analysis that simultaneously examined the three microwear variables (Fig. 2 C and tables S9 to S11), in which Pliobates falls closer to the extant frugivorous–mixed-feeder centroid for the first and second canonical axes and is classified as a frugivore. Dental microwear analyses therefore indicate a mainly frugivorous diet for Pliobates, compatible with a high consumption of ripe fruit and a low sclerocarpic component. Dental BM estimates for the female holotype of Pliobates (table S7) range from 2.9 to 4.8 kg, with an average BM estimate of 3.9 kg and an uncertainty degree (based on the combined 95% confidence intervals for each dental locus) of 2.5 to 5.7 kg. Postcranial BM estimates (table S8) are on average 4.8 kg (range: 4.0 to 5.6 kg), based on catarrhine regressions, and 4.3 kg (range: 2.6 to 6.3 kg), based on hominoid regressions (estimates for each postcranial estimator and their confidence intervals are given in table S8). Given that the size of Pliobates is in the lower range for extant hominoids, the catarrhine regressions probably yield more accurate estimates, although the hominoid-based estimates are closer to the dental ones. Overall, the body mass of the holotype of Pliobates cataloniae can be estimated at ~4 to 5 kg (much lower than that estimated for Epipliopithecus vindobonensis, ~11 to 12 kg; table S8). A 3D virtual reconstruction of the cranium, based on the preserved specimens, is shown in fig. S3, whereas the final reconstruction (including mirrored portions) is shown in Fig. 3 and movie S1. Based on this reconstruction, the cranium of Pliobates differs from the primitive catarrhine condition (22, 37, 60) by being short, wide, and high. However, the tubular ectotympanic is short and incompletely ossified—i.e., less developed than in Saadanius and extant crown catarrhines (20–22). The maxillary sinus is extensive, as in stem catarrhines and hominoids (22, 80), and there is also a small frontal sinus, as in stem hominoids but unlike in stem catarrhines, cercopithecoids, hylobatids, and pongines (22, 37, 80). The face is short and displays anteriorly situated orbits, as in hylobatids, colobines, and some extinct small-bodied catarrhines such as Epipliopithecus, Micropithecus, and Lomorupithecus (19, 21, 38, 76, 79). However, Pliobates differs from these taxa (and more closely resembles hylobatids) by displaying a more well-defined muzzle (especially compared with Epipliopithecus) with long and more horizontal nasals, a higher zygomatic root (moderately high as in hylobatids, but less so than in hominids), an interorbital pillar nearly orthogonal to the frontal squama (as in hylobatids and chimpanzees), a high degree of orbital convergence and frontation (as in all extant hominoids), and thin and anteriorly projecting (telescopic) orbital rims [to a greater extent than in Epipliopithecus (38), and thus most closely resembling hylobatids and, as far as it can be ascertained with incomplete preservation, Micropithecus (79)]. Pliobates also displays derived hominoid features in the basicranium (Fig. 4 A), including the absence of a postglenoid foramen with a large and ventrally visible jugular foramen (as in all extant hominoids), the foramen ovale situated anteriorly and laterally to the Eusta- chian aperture (as in hylobatids and African and African apes), and the horizontal and anapes), the fusion between the auditory meatus teriorly directed carotid canal in the petrosal bone and the postglenoid process (as in hylobatids (as in hylobatids). Braincase measurements yield an average cranial capacity estimate of 69.0 cm3 (range: 41.4 to 110.7 cm3; table S12), which is close to the estimates of 60.1 and 65.3 cm3 delivered by the two most reliable estimators (57) and only slightly lower than the estimate of 75.1 cm3 obtained from foramen magnum area (table S12). According to our estimates of body mass (4.5 kg) and cranial capacity [72 cm3 (average of cranial and foramen magnum estimates)], Pliobates would display a monkey-like degree of encephalization extensively overlapping with extant cercopithecoids (fig. S4 and table S13), being much more encephalized than the stem catarrhine Aegyptopithecus, slightly more so than the stem cercopithecoid Victoriapithecus, and only slightly less so than hylobatids and the extinct hominoids Proconsul and Oreopithecus. All these taxa, like cercopithecoids, are less encephalized than the extinct hominoid Hispanopithecus (Rudapithecus) and the extant great apes. Although humans are outliers in brain size–body size allometric regressions, great apes further display an allometric grade shift compared with hylobatids (and Pliobates), which are only slightly more encephalized on average than cercopithecoids (58). The humerus (Fig. 5) resembles that of extant crown catarrhines, proconsulids, and dendropithecids by lacking (unlike Epipliopithecus) an entepicondylar foramen (17, 20, 21, 38, 79, 81, 82). Pliobates more closely resembles extant hominoids in the laterally facing bicipital tuberosity in the radius (81, 83) (Fig. 5), as well as in the configuration of the humeroradial articulation (81, 82, 84) (Fig. 6), including: in the humerus, the lack of capitular tail [present in Epipliopithecus, dendropithecids, and cercopithecoids (84)] and the moderately globulous (although not posterolaterally expanded) capitulum with a well-developed zona conoidea [lacking in Epipliopithecus and dendropithecids (81–84)]; and, in the radius, the only slightly tilted and almost circular radial head with a small and flat area, a reduced lateral lip, and a beveled surface for the humeral zona conoidea. Pliobates also has a hominoid-like diarthrodial distal radioulnar joint (85–87), with a two-faceted expanded semilunar articulation in the ulnar head (Fig.6). In this regard, Pliobates departs from Epipliopithecus, dendropithecids, and cercopithecoids (17, 38, 82) and more closely resembles Proconsul (88), although the ulnar head is less extensive than in extant hominoids. In contrast with these derived features, the humeral shaft and humeroulnar joint are plesiomorphic: The former (Fig. 5) is anteriorly straight and somewhat proximally retroflexed; the latter (Fig. 6) lacks the stabilizing features of extant hominoids (83, 89), as shown by the narrow ulnar trochlear notch without a median keel (in agreement with the absence of spooling and the poorly defined trochlear lateral keel in the humerus of Pliobates). Humeral torsion in Pliobates is estimated at 101°, irrespective of the method employed (based on the posterior buttress for the humeral head or the bisector of the bicipital groove), with a confidence interval spanning 95.7° to 106.3° [based on the prediction error (5.23%) for the bicipital groove method (50)]. This degree of torsion is moderate, higher than that estimated for Proconsul heseloni (92°), but comparable to estimates for Dryopithecus fontani (102°) and Dendropithecus macinnesi (103.5°), and only slightly below the value estimated for Epipliopithecus vindobonensis (109°). The humeral torsion of Pliobates is thus most comparable to that of non-atelid platyrrhines and lower than that of Ateles and extant hominoids, especially African great apes and humans [although the high degree of humeral torsion of extant hominoids is related to increased mobility at the glenohumeral joint, the higher values of great apes and humans appear related to knucklewalking and enhanced manipulation, respectively, rather than suspensory behaviors (50)]. In contrast to the moderate humeral torsion, the forelimb of Pliobates appears somewhat elongated relative to its body size (fig. S5). Allometric computations of relative forelimb length in fossils (residuals are given in table S14) must be considered with caution, because they are dependent on the accuracy of body-size estimates. However, the forelimb of Pliobates (based on our BM estimate of 4.5 kg) appears more elongated than that of Epipliopithecus (based on our estimate of 11.5 kg). The latter taxon, contrary to previous assertions (81, 83), has the generalized proportions of quadrupedal monkeys. Pliobates, in contrast, has a forelimb elongation similar to that of female orangutans and Brachyteles, although it is less extreme than in Ateles and especially than in hylobatids. The same pattern holds when the humerus and radius are analyzed separately, although in Pliobates, relative length is somewhat higher for the radius than for the humerus. Pliobates further displays a high arm angle (8°), which is considerably greater than the average in most anthropoids, except Hylobates (9.8°), Pongo (6.3°), and Ateles (6.5°) (51). The ulnocarpal articulation of Pliobates is completely different from that of Epipliopithecus and dendropithecids (17, 38, 90), including a partially developed ulnar fovea (Fig. 6), which, in extant hominoids, is the attachment area of the triangular disc ligament and the intra-articular meniscus (85–87). The ulnar styloid process is relatively long and slender, with no discernible articular surfaces for the pisiform or triquetrum. This agrees with the lack of an articular facet for the styloid process on the pisiform, like extant hominoids but unlike monkeys and Proconsul (91). However, in contrast to Pierolapithecus (92), the triquetrum of Pliobates shows a proximal articular facet, which is more developed than that present in hylobatids and sometimes Pan (51, 87) but less developed than in monkeys. This suggests that ulnotriquetral contact might have been reduced by some kind of intra-articular tissue, similarly to some <

Hispanomeryx Morales, Moya-Sola and Soria, 1981, is a moschid pecoran from the middle-early late Miocene of Europe (MN 6 to MN 11; Morales et al., 1981; Azanza, 1986; Prothero, 2007; Vislobokova, 2...