The shape of the tree of life is the result of shifting diversification rates, and identifying the factors driving these shifts is one of the main aims in evolutionary biology. Various biotic and abiotic factors have been proposed to have an impact on mammal diversification, such as climatic and tectonic changes, the acquisition of new traits, and expansion into new ecosystems or landmasses.
We used phylogenetic comparative methods to quantify the influence of potential drivers on diversification patterns in extant squirrels (Sciuridae, Rodentia). We conducted a multilayer approach, comparing diversification rates among squirrel lineages depending on their degree of biome specialisation, biogeographic realm occupancy, locomotion adaptations, and presence in mountainous regions.
We generated the most complete phylogeny of squirrels to date, encompassing almost 80% of the extant species, and applied multiple and binary state¿dependent diversification models. All the traits examined showed an influence on diversification rates.
The biome specialist lineages showed the highest speciation rates, suggesting a major role of bioclimatic specialisation on macroevolutionary patterns. A single major event, the Miocene¿Pleistocene radiation of terrestrial¿adapted lineages in North America, left a signal that was recovered in two of our analyses. Both the Nearctic lineages and the terrestrial¿adapted lineages showed high speciation rates, highlighting the fact that that major evolutionary episodes may produce confounding effects in state¿dependent diversification models.
Ancestral reconstructions showed that cold and warm intervals in Earth¿s history had different effects on squirrels¿ diversification, depending on their climatic affinities. Tropical and arboreal squirrels evolved predominantly in the warm intervals, while terrestrial and cold¿adapted squirrels radiated in the cold intervals.
Our findings suggest that, while global climatic shifts are key for the speciation processes in mammalian lineages, lineage¿specific ecological adaptations are critical modulators of the responses of lineages to such environmental shifts, in an interplay that ultimately affects their diversification patterns.

The new fossil micromammal assemblage of Çapak represents a mixture of both Anatolian and European
faunal elements. The locality is very important for understanding faunal evolution in the less well-known time interval at the end of the early Miocene of western Anatolia. In Çapak, nine species of rodents and one species of ochotonid were encountered: the hamsters Democricetodon gracilis, Megacricetodon primitivus, Eumyarion aff. E. montanus, Cricetodon cf. C. aliveriensis, Cricetodon sp., and Karydomys cf. K. strati, the mole-rat Debruijnia sp., the squirrel Aliveria luteyni, the dormouse Myomimus tanjuae n. sp., and the pika Albertona balkanica. The assemblage is referable to Anatolian local zone E or MN unit 4. The relative abundance of the various genera is markedly different from that of the older early Miocene assemblages, suggesting that the environment in Anatolia became drier and had a more open landscape., This work was funded by National Geographic grant “Palaeogeography of Mammals Following the Collision of the African and Eurasian Plates” (GEFNE 140-15). This study was supported by Ege University (TTM/001/2016) and (TTM/002/2016) to SM
and MB. This study was also supported by grants UK/123/2019, UK/145/2020; Scientific Grant Agency of the Ministry of Education, Science, Research and Sport of the Slovak Republic VEGA 1/0164/19, 1/0533/21, and by the Slovak Research and
Development Agency under contract Nos. APVV-15-0575 and APVV-16-0121. This work was partially supported by project PGC2018-094122-B-I00 funded by FEDER/ Ministerio de Ciencia e Innovación-Agencia Estatal de Investigación., Peer reviewed

Mortality curves of extinct rodents can be reconstructed by using relative wear calculated from the amount of dentine exposed. MicroCT scanning of molars of the extinct Eomyid genus Ligerimys shows that this Wear Index increases gradually with wear; this means that wear classes can serve as a proxy for age and can therefore be used to reconstruct mortality curves. We reconstructed mortality curves of three species of Ligerimys from the late early Miocene of Spain. These curves show a similar trend, with decreasing numbers of molars as the amount of wear in each class increases. The curves show remarkable similarities to those of the wood mouse Apodemus. This suggests that Ligerimys was, like Apodemus, at the fast end of the fast-slow continuum, in other words r-selected, though not as extreme as some other rodents., This work was supported by the European Union [SYNTHESYS project, grant ES-TAF-1084]; Ministerio de Ciencia e Innovación-Agencia Estatal de Investigación; FEDER/ European Union [PGC2018-094122-B-I00]., Peer reviewed

The new micromammal site of Beydere 3 represents a typical Anatolian MN3 fauna in the high diversification and abundance of Eumyarion. Notably, two Eumyarion species are new; combined, they make up almost half of the assemblage. While Eumyarion beyderensis sp. nov. shows simple morphology in upper molars, Eumyarion aegeaniensis sp. nov. shows a more complex lophs pattern. In addition, Beydere 3 yielded nine species of rodents and an ochotonid: the hamsters Cricetodon kasapligili, Mirrabella crenulata, Megacricetodon hellenicus, Democricetodon doukasi, Vallaris zappai and Eumyarion sp., the dormice Glirulus ekremi and Glis sp., the beaver Steneofiber eseri, the squirrel Palaeosciurus fissurae and Ochotonidae indet. The fauna is significant in that it represents the first common occurrence of Megacricetodon in Anatolia. Eumyarion and Megacricetodon are both dominant groups which may indicate a signal of environmental change, but still closer to wet conditions. This, in combination with the new species of Eumyarion, suggests that Beydere 3 represents a time slice which was previously not recorded in Anatolia. Radiometric dating of ashes overlying the section indicates that the age of the locality is older than 18.21 (± 0.19) Ma., This work was supported by the Ege Üniversitesi (TTM/001/2016,TTM/002/2016);National Geographic Society (GEFNE 140-15); Spanish Ministry of Science and Innovation (PGC2018-094122-B-I00); Univerzita Komenského v Bratislave (UK/400/2021); Vedecká Grantová Agentúra MŠVVaŠ SR a SAV (1/0164/19, 1/0533/21) and Slovak Research and Development Agency under contracts (APVV-15-0575)., Peer reviewed

[Methods] The material and methods section of the article contains all details concerning the methods used in this study.
[Usage Notes] The readme file contains an explanation of each file uploaded in supplementary files., Establishing an evolutionary timescale is fundamental for tackling a great variety of topics in evolutionary biology, including the reconstruction of patterns of historical biogeography, coevolution and diversification. However, the tree of life is pruned by extinction and very generally molecular data cannot be gathered for extinct lineages. Methodological challenges have prevented until recently the application of tip-dating Bayesian approaches in morphology-based fossil-only datasets. Herein we present a morphological dataset for a group of cricetid rodents to which we apply a battery of methods fairly new in palaeontology that can be used by palaeontologists for the analysis of entirely extinct clades. We compare the tree topologies obtained by traditional parsimony, Bayesian dated and undated phylogenetic approaches and calculate stratigraphic congruence indices for each. Bayesian tip-dated clock methods seem to outperform parsimony in the case of our dataset, which includes highly homoplastic morphological characters. Regardless, all three topologies support the monophyly of Megacricetodontinae, Democricetodontinae and Cricetodontinae. Dispersal and speciation events inferred through Bayesian Binary Markov chain Monte Carlo and biodiversity analyses provide evidence for a correlation between biogeographic events, climatic changes and diversification in cricetids., Consejo Superior de Investigaciones Científicas, Award: PGC2018-094122-B-100 (MICU/AEI/FEDER,EU).
European Commission, Award: ES-TAF-2579., Peer reviewed

[EN] Badgers belong to the genus Meles Brisson, 1762, which comprise four extant species (M. anakuma
Temminck, 1844, M. leucurus (Hodgson, 1847), M. canescens Blanford, 1875, and M. meles (Linnaeus, 1758)). The genus is included in the subfamily Melinae Bonaparte, 1838, a polyphyletic group of Eurasian mustelids whose evolutionary relationships need further clarification. The evolutive relationships of the genus are complex and far from being resolved. This work aims to describe a nearly 1 Ma unpublished badger mandible from the Sierra de Quibas (Murcia) and to help clarify the evolutionary patterns of Euroasiatic badgers. To this end, we used 2D geometric morphometric techniques to measure 57 landmarks and semilandmarks in 79 first lower molars (m1) of Meles, ranging from Pleistocene to extant species. Our results show evidence for differentiating between primitive badgers and living species of Meles. The new m1 of Meles from the Quibas site is more gracile (relatively narrower and longer) than the other Eurasian extinct species, and shows that this
specimen can be placed in the subspecies M. meles meles (Linnaeus, 1758). Our results also show that
the denomination of M. meles atavus Kormos, 1914 as a related subspecies with a primitive morphology is morphologically supported. Therefore, we conclude that the living subspecies of badger M. meles meles was already differentiated in the south of the Iberian Peninsula at around 1 Ma, but some primitive remnant populations persisted in the north of the Iberian Peninsula, for which we recognize the subspecies M. meles atavus., [FR] Morphométrie géométrique 2D de la première molaire inférieure du genre Meles, incluant de nouvelles
preuves de la présence de blaireaux sur le site de Quibas du Pléistocène inférieur (Murcie, Espagne).
Les blaireaux sont un groupe polyphylétique de mustélidés au sein duquel on retrouve le genre Meles
Brisson, 1762. Il appartient à la sous-famille Melinae Bonaparte, 1838 de distribution eurasienne,
actuellement composée de quatre espèces vivantes (M. anakuma Temminck, 1844, M. leucurus
(Hodgson, 1847), M. canescens Blanford, 1875 et M. meles (Linnaeus, 1758)). Les relations évolutives
du genre sont complexes et loin d’être résolues. L’objectif de ce travail est de décrire une mandibule
de blaireau inédite de près de 1 Ma de la Sierra de Quibas (Murcie) et d’aider à clarifier les schémas
évolutifs des blaireaux euroasiatiques. À cette fin, nous avons utilisé des techniques morphométriques
géométriques 2D pour mesurer 57 repères et semi-repères dans 79 premières molaires inférieures
(m1) de Meles, allant du Pléistocène aux espèces existantes. Nos résultats montrent des preuves solides
de la différenciation entre les blaireaux primitifs et les espèces vivantes de Meles. Le nouveau m1 de
Meles du site de Quibas est plus gracile (relativement plus étroit et plus long) que les autres espèces
eurasiennes éteintes, et montre que ce spécimen peut être placé dans la sous-espèce M. meles meles
(Linnaeus, 1758). Nos résultats montrent également que la dénomination de M. meles atavus Kormos,
1914 en tant que sous-espèce apparentée avec une morphologie primitive est étayée morphologiquement.
Par conséquent, nous concluons que la sous-espèce vivante de blaireau M. meles meles
était déjà différenciée dans le sud de la péninsule Ibérique à environ 1 Ma, mais certaines populations
relictuelles persistent dans le nord de la péninsule Ibérique, pour lesquelles nous reconnaissons la
sous-espèce M. meles atavus., This work was supported by Ministerio de Ciencia e Innovación (MICINN/FEDER), Spain, Grant Number: CGL2016-75109-P; the Palarq Foundation, Spain; and the Comunidad Autónoma de la Región de Murcia,
Spain, Grant Number: ARQ115/2018 (Subvención para la Investigación e Intervención en el Patrimonio Arqueológico y Paleontológico de la Región de Murcia). The Comunidad Autónoma de la Región de Murcia allowed and supported the field work. AV is also funded by the Spanish Ministry of Economy and Competitiveness and FEDER/EU (Research Projects PGC2018-094122-B-100 and PID2020-
116220GB-100), the Government of Aragon (Group ref. E33_20R), and the Research Group UCM 910607. PP is beneficiary of a postdoctoral fellowship from the Argentinian Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET). The “Juan de la Cierva Formación” program (FJCI-2017-32157), from the Spanish Ministry of Science and Innovation, funds DGM., Peer reviewed

Part of the Vertebrate Paleobiology and Paleoanthropology book series (VERT), The aim of this review is, first of all, to summarize our knowledge on Miocene Moschidae and provide a helpful tool for the determination of moschids in Miocene faunas. Secondly, it gives an overview of our current knowledge on the spreading of Miocene Moschidae by characterizing the environmental conditions at key localities across Eurasia. Although many questions still remain unanswered, the spreading of moschids can clearly be correlated with the cooler, drier, more open, and especially more seasonal conditions that changed the terrestrial ecosystems across Eurasia during the Middle Miocene Climatic Transition. Moschids, far from being indicators for closed and humid environments as often assumed, were able to cope with drier and more seasonal environments, and could prove a helpful tool for paleoenvironmental reconstructions in the future; e.g., to also tackle environments of Miocene primates in more detail., This research received support from the SYNTHESYS Project https://www.synthesys.info/ (FR-TAF-1892, FR-TAF-6335, ES-TAF-6266) which is financed by European Community Research Infrastructure Action under the FP7 “Capacities” Program. IMS acknowledges support by the Spanish Government Agencia Estatal de Investigación—European Regional Development Fund of the European Union (projects MINECO/FEDER-UE; CGL2016-76431-P and CGL2017-82654-P, AEI/FEDER EU), the Spanish Ministry of Science and Innovation (MICINN) and FEDER funds (project ref. PID2020-116220GB-I00) and the Generalitat de Catalunya (CERCA Programme). This research was also supported by the Chinese Academy of Sciences (XDB26000000, XDA20070203, 2019QZKK0705, and XDB31030106). We also appreciate Luda Xing, IVPP, for data collection in respect to Chinese fossil Moschidae. JM received support from the Spanish project PGC2018-094122-B-100 (Ministerio de Investigación e Innovación). LC and BM acknowledge support from the Swiss National Science Foundation (SNF projects 200021_159854 and 200021_178853). BM wants to thank for access to collections J. Meng, S. Bell, J. Galkin, R. O’Leary, S. Goldberg, M. Norell, and A. Gishlick (AMNH New York, USA), as well as Daxner Höck and U. Göhlich (NHMW). BM’s gratitude goes to the Swiss National Science Foundation (SNF project P3003P2_161065 and P3P3P2_161066)., Peer reviewed

Identifying the drivers of adaptation is key to understanding the origin and evolution of diversity. Here we study the morphological evolution of tooth morphology, a classic example of a conserved structure, to gain insights into the conditions that can overcome resistance to evolutionary change. We use geometric morphometrics of the occlusal surface outline of the fourth lower premolar (p4) of squirrels, a paradigm of a stable tooth morphology, to explore morphological adaptations to diet. Although a versatile generalist dental morphology favors the retention of the ancestral shape, the acquisition of diets that require strong mechanical processing drives morphological change. In particular, species that eat both grass and dry fruits evolved disparate tooth shape morphologies, related to trade-offs between feeding performance that lead to a more or less pronounced change depending on the proportion of those items in their diet. Also, some folivores develop relatively large p4s, and most bark gleaners have relatively small p4s. Ultimately, despite the role of diet shaping these patterns, we showed that diet is not the only factor driving the evolution of tooth morphology., I.M. was funded by a predoctoral grant from the Complutense
University of Madrid (CT27/16-CT28/16) and two travel grants of the same institution to visit museum collections (EB40/17 and EB14/19). I.M. was awarded with the
collection study grant in 2018 granted by the American
Museum of Natural History (AMNH) in New York. This
work was partially supported by the project PGC2018-
094955-A-I00 and PGC2018-094122-B-I00 of the Spanish
Ministerio de Ciencia e Innovación (MCI/AEI/FEDER,
EU). This is a contribution of the Research Group Team
UCM-910607 on the Evolution of Cenozoic Mammals and
Continental Palaeoenvironments of UCM., Peer reviewed

The climatic evolution of the Neogene, with long-term cooling disrupted by the Middle Miocene Climatic Optimum (MMCO; ~17–14.75 Ma), arises as a suitable baseline to analyze the effects of these transcendent climatic changes on the mammalian community structures. The present investigation is an attempt to examine the palaeohabitat of a Neogene (Middle Miocene: ~15–11.5 Ma) geographically distant (i.e., from Spain, China, and India) extinct mammalian communities utilizing the cenogram approach (in both qualitative and quantitative framework). The detailed statistical analyses (presented herein) incorporating a total of eight mammalian communities allows us to infer predominance of Tropical Deciduous Forest environments between ~15 and ~11.5 Ma interval, with several pulses of distinctive aridity experienced by some communities thriving within the Iberian region. On the contrary, stable forested conditions were witnessed by the middle Miocene communities of Asia [i.e., the ~11.5 million-year-old mammalian community of Laogou (China), and the ~13.5 million-year-old mammalian community of Ramnagar (north India)]. Our present investigation also infers that additional mammalian remains (particularly of body mass of <35 kg) are warranted to decipher the habitat (based on cenogram approach) of the Middle Miocene (~13 Ma) mammalian community of Kalagarh (Himalayan Foreland Basin, north India) and the Middle Miocene (~14 Ma) mammalian community of Palasava (Kutch Basin, western India). Nonetheless, the Cenogram technique (being continuously developed over the past six decades) may become an important tool to decipher any habitat change(s) of western India’s mammalian communities considering renewed palaeontological efforts within the Neogene of the region.

The present study is an attempt to utilize cenogram methodology (both qualitative and quantitative) to consider mammalian communities from fve early to late Eocene localities across the globe (i.e., Polecat Bench, Bighorn Basin, North America; Abbey Wood, Blackheath Formation, UK; Cambay Shale, Cambay Basin, India; Wutu Formation, Wutu Basin, China; Pondaung Formation, Myanmar) so as to provide a comparative palaeohabitat framework. It is also a frst attempt to examine the palaeohabitat of an extinct mammalian community (i.e., from Cambay Shale) in India utilizing the cenogram approach. In addition, seven extinct middle-Miocene communities (Laogou, Linxia Basin, China; Estación Imperial, Spain; Paseo de las Acacias, Spain; Arroyo del Olivar-Puente de Vallecas, Spain; Somo-saguas, Spain; Paracuellos 5, Spain; Paracuellos 3, Spain) have also been considered, in order to provide a global perspective to the climatic inferences in a temporal context. The majority of statistical calculations for Paleogene communities expose forested and humid conditions, excluding the Cambay Shale mammalian community of India. A hidden diversity within the medium to large body-size category of mammals (disguising the mammal biodiversity expected in tropical forested habitats) from Cambay Shale (western India) is a plausible cause of digression in the results. This is refected in the histograms showing relationships between proportions of mammal species in various body-mass categories. Furthermore, the results show that Neogene mammalian communities were sustained in comparatively open habitats. Diferences between occidental European and Asian localities in the canopy and humidity of the Neogene environments are also refected in our analyses.

We describe two carnivoran coprolites found in the pseudokarst natural carnivore trap of Batallones-3, from the Late Miocene of Spain. The larger one, comprising multiple indistinguishable fragments of broken and corroded bones, indicates that the producer of the dropping might have been highly capable of crushing the softer parts of large bones. On the other hand, the smaller one shows several relatively larger and more complete bone fragments, thus exhibiting a greater capacity to break and swallow large portions of bone. The external morphology of the large coprolite is similar to that of extant bears, whereas the smaller one more closely resembles that of the living insectivorous hyaenid Proteles in morphology, on the one hand, and that of the viverrid Genetta in size, on the other hand. We hypothesize that the amphicyonid Magerycion anceps was the producer of the large coprolite and the jackal-sized basal hyaenid Protictitherium crassum excreted the smaller one. Thus, we present the first direct evidence of a bone durophagous diet in the carnivorans of Batallones.

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.

Els Casots is one of the richest fossil vertebrate sites of the Vallès-Penedès Basin (Catalonia, Spain). It was discovered in 1989 and excavated briefly during the 1990s, resulting in the recovery of thousands of remains and the erection of several new mammal species. Excavations were resumed in 2018 and continue to date. Here we provide updated results regarding the age, stratigraphy, biota and palaeoenvironment of the site. The age of the site is well constrained to ~15.9 Ma thanks to recent bio- and magnetostratigraphic data, thus coinciding with the Miocene Climatic Optimum (MCO). The stratigraphic succession at the site area indicates lacustrine to palustrine environments with cyclically oscillating water level. There are several fossiliferous layers that have yielded a vertebrate fauna comprising up to 75 different vertebrate species including amphibians, reptiles, birds and mostly mammals. The finding of several articulated partial skeletons indicate that the site records an autochthonous to parautochthonous assemblage. The abundance and completeness of the vertebrate remains together with a well-constrained age and detailed stratigraphic and palaeoenvironmental data, make els Casots a key site for understanding wetland ecosystems in southern Europe during the MCO.

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.

The present study deals with new material of carnivorans (Mustelidae, Mephitidae, Ailuridae, Potamotheriinae and Viverridae) from the basal Tortonian (Late Miocene, late Astaracian) hominid-bearing locality of Hammerschmiede (Bavaria, Germany). The small carnivoran fauna includes 20 species belonging to nine different subfamilies (Guloninae, Lutrinae, Mellivorinae, Potamotheriinae, Leptarctinae, Mephitinae, Simocyoninae, Genettinae and Viverrinae). The identified forms include: "Martes" sansaniensis, "Martes" cf. munki, "Martes" sp., Circamustela hartmanni n. sp., Laphyctis mustelinus, Guloninae indet., Eomellivora moralesi, Vishnuonyx neptuni, Paralutra jaegeri, Lartetictis cf. dubia, Trocharion albanense, Palaeomeles pachecoi, Proputorius sansaniensis, Proputorius pusillus, Alopecocyon goeriachensis, Simocyoninae indet., Potamotherium sp., Semigenetta sansaniensis, Semigenetta grandis and Viverrictis modica. The new species Circamustela hartmanni n. sp. is differentiated from the other members of the genus by its small size and the morphology of its dental cusps in the upper and lower carnassials. This is one of the highest reported taxonomic diversities for fossil small carnivorans in the Miocene of Europe, including also first and last occurrences for several genera and species. Additionally, the assemblage comprises some rare taxa such as Palaeomeles pachecoi and Eomellivora moralesi. An ecomorphological comparison of the discovered taxa reveals possible cases of competition and niche partitioning.