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This work received financial support from the research projects CGL2015–68333-P (MINECO/FEDER, UE), Research Group BSCHUCM 910607 and Consolidated Research Group EO5 (Aragón Government). Referencias bibliográficas: • Adrover R (1986) Nuevas faunas de roedores en el Mio-Plioceno continental de la región de Teruel (España). Interés bioestratigráfico y paleoecológico. Instituto de Estudios Turolenses, Teruel • Adrover R, Feist M, Hugueney M, Mein P, Moissenet E (1982) L'áge et la mise en relief de la formation détritique culminante de la Sierra Pelarda (Province de Teruel, Espagne). C R Acad Sci Paris 295:231–236 • Aguirre E (1973) Conservación e historia de la Naturaleza. Boletín Estación Central. Ecología 2:89–97 • Aguirre E, Morales J, Soria D (1981) Accumulated bones in a Pliocene cave in Cerro Pelado, Spain. Natl Res Rep 13:69–81 • Agustí J, Santos-Cubero A, Furió M, De Marfa R, Blain H-A, Oms O, Sevilla P (2011) The late Neogene-early quaternary small vertebrate succession from the Almenara-Casablanca karst complex (Castellón, Eastern Spain): chronologic and paleoclimatic context. Quat Int 243:183–191 • Calvo JP, Daams R, Morales J, López-Martínez N, Agustí J, Anadón P, Armenteros I, Cabrera L, Civis J, Corrochano A, Díaz-Molina M, Elizaga E, Hoyos M, Martín-Suárez E, Martínez J, Moissenet E, Muñoz A, Pérez-García A, Pérez-González A, Portero JM, Robles F, Santisteban C, Torres T, van der Meulen J, Vera JA, Mein P (1993) Up-to-date Spanish continental Neogene synthesis and paleoclimatic interpretation. Rev Soc Geol Esp 6:29–40 • Daams R, Meulen A, Van Der Peláez-Campomanes P, Álvarez-Sierra MA (1999) Trends in rodent assemblages from the Aragonian (early-middle Miocene) of the Calatayud-Daroca Basin, Aragón, Spain. In: Agustí J, Rook L, Andrews P (eds) Hominoid evolution and climatic change in Europe (the evolution of Neogene terrestrial ecosystems in Europe), vol 1. Cambridge University Press, Cambridge, pp 390–412 • Hilgen FJ, Lourens LJ, Dam JA Van, with Contr, Corral de Lobato, a karstic site in the area of Molina de Aragón has been studied in a preliminary way. Even though there are not many Neogene karstic sites in the Iberian Chain, they occur in four clusters, with ages ranging from latest middle Miocene (MN7/8) to early Pleistocene (MN17). Correlations between these clusters and the reference stratigraphical units of the Tagus Basin, as well as with local and global events, are proposed. These karstic sites provide a complementary source of fossil vertebrate remains to that of the stratified sites formed lowland. The Heritage significance of such sites arises from the enhanced preservation of rare taxa or associations, and the operation of biotic concentrative processes., Gobierno de Aragón, Depto. de Didáctica de las Ciencias Experimentales , Sociales y Matemáticas, Fac. de Educación, TRUE, pub

The Pontils fossil site (middle Eocene, Ebro Basin, Spain) includes several vertebrate-bearing levels situated in a sequence recording a continental to marine transition. Although the locality has been known since the 1980s and scarce mammal remains have been already documented, an intensive sampling has not been developed until now. This work presents the first results of the recent field campaigns carried out at this site. Seven levels have yielded significant vertebrate remains, revealing a diverse assemblage which includes chondrichthyans, actinopterygians, amphibians, crocodilians, squamates, metatherians, eulipotyphlans, apatotherians, chiropterans, rodents, artiodactyls, perissodactyls, and primates, besides other non-vertebrate fossils. The Pontils assemblage indicates a mangrove swamp environment with warm and humid conditions and increasing marine influence towards the top of the sequence. Among mammals, primates are particularly diverse, including abundant remains of a minuscule, still undetermined omomyiform, and scarce teeth of Pseudoloris, Necrolemur, and an undetermined anchomomyin. The occurrence of larger benthic foraminifera allows the assignment of Pontils to Shallow Benthic Zone 17 (Bartonian), solving the debate about the age of the locality, previously assigned either to the Bartonian or the Lutetian. These data, together with previous magnetostratigraphic analyses, allow correlation to chrons C18r or C18n.1r, constraining the age of Pontils to between 39.58 and 41 Ma. Therefore, the Pontils site represents a new reference section for the correlation of marine and continental biostratigraphy during the middle Eocene.

The Pontils fossil site (middle Eocene, Ebro Basin, Spain) includes several vertebrate-bearing levels situated in a sequence recording a continental to marine transition. Although the locality has been known since the 1980s and scarce mammal remains have been already documented, an intensive sampling has not been developed until now. This work presents the first results of the recent field campaigns carried out at this site. Seven levels have yielded significant vertebrate remains, revealing a diverse assemblage which includes chondrichthyans, actinopterygians, amphibians, crocodilians, squamates, metatherians, eulipotyphlans, apatotherians, chiropterans, rodents, artiodactyls, perissodactyls, and primates, besides other non-vertebrate fossils. The Pontils assemblage indicates a mangrove swamp environment with warm and humid conditions and increasing marine influence towards the top of the sequence. Among mammals, primates are particularly diverse, including abundant remains of a minuscule, still undetermined omomyiform, and scarce teeth of Pseudoloris, Necrolemur, and an undetermined anchomomyin. The occurrence of larger benthic foraminifera allows the assignment of Pontils to Shallow Benthic Zone 17 (Bartonian), solving the debate about the age of the locality, previously assigned either to the Bartonian or the Lutetian. These data, together with previous magnetostratigraphic analyses, allow correlation to chrons C18r or C18n.1r, constraining the age of Pontils to between 39.58 and 41 Ma. Therefore, the Pontils site represents a new reference section for the correlation of marine and continental biostratigraphy during the middle Eocene.

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.

En los últimos años se han realizado estudios de análisis de morfometría geométrica de ocotónidos provenientes de diversos yacimientos europeos. Sin embargo, en la Península Ibérica, pese a ser muy abundantes, aún no se había realizado ningún estudio de este tipo. Se han escaneado a partir de microtomografía computerizada cinco restos craneales de Prolagus crusafonti (Ochotonidae, Lagomorpha) provenientes del Cerro de los Batallones (Mioceno superior, Madrid, España). Mediante el proceso de segmentación, a través del cual se consigue aislar y reconstruir de manera digital el elemento anatómico, se han obtenido cinco hemimandíbulas de diferentes individuos de P. crusafonti. Una vez aisladas comenzó el proceso de digitalización de landmarks de tres de estas hemimandíbulas junto con otras 77 de ocotónidos fósiles y actuales de los géneros Prolagus, Lagopsis y Ochotona. De esta manera se obtuvieron los datos geométricos de estas hemimandíbulas definidos en función de landmarks con los que se llevó a cabo un Análisis de Componentes Principales (PCA) para comparar y analizar la variabilidad morfológica de la muestra. Estos resultados muestran diferencias morfológicas entre los diferentes géneros analizados que aluden a estructuras fuertemente relacionadas con la biomecánica de la mandíbula. Estas diferencias podrían explicarse en términos de variación en la composición de la dieta, diferente desarrollo de las piezas dentales, diferente historia evolutiva y la insularidad y aislamiento de alguna de las especies. Las diferencias latitudinales y altitudinales en las distribuciones geográficas de las especies actuales y las inferidas para las especies extintas podrían influir en su dieta. De esta manera, la mandíbula reflejaría la dureza del alimento que consumían. En el caso de la erupción diferencial de las piezas dentales, se han observado diferencias morfológicas entre los ejemplares debido a la ausencia, en alguna de las especies, del tercer molar inferior. Por último, y

The modern era of analytical and quantitative palaeobiology has only just begun, integrating methods such as morphological and molecular phylogenetics and divergence time estimation, as well as phenotypic and molecular rates of evolution. Calibrating the tree of life to geological time is at the nexus of many disparate disciplines, from palaeontology to molecular systematics and from geochronology to comparative genomics. Creating an evolutionary time scale of the major events that shaped biodiversity is key to all of these fields and draws from each of them. Different methodological approaches and data employed in various disciplines have traditionally made collaborative research efforts difficult among these disciplines. However, the development of new methods is bridging the historical gap between fields, providing a holistic perspective on organismal evolutionary history, integrating all of the available evidence from living and fossil species. Because phylogenies with only extant taxa do not contain enough information to either calibrate the tree of life or fully infer macroevolutionary dynamics, phylogenies should preferably include both extant and extinct taxa, which can only be achieved through the inclusion of phenotypic data. This integrative phylogenetic approach provides ample and novel opportunities for evolutionary biologists to benefit from palaeontological data to help establish an evolutionary time scale and to test core macroevolutionary hypotheses about the drivers of biological diversification across various dimensions of organisms.

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.

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.

Establishing an evolutionary timeline 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 molecular data cannot be gathered for extinct lineages. Until recently methodological challenges have prevented the application of tip-dating Bayesian approaches in morphology-based fossil-only data sets. Herein, we present a morphological data set for a group of cricetid rodents to which we apply an array of methods fairly new in paleontology that can be used by paleontologists for the analysis of entirely extinct clades. We compare the tree topologies obtained by traditional parsimony, time-calibrated, and noncalibrated Bayesian inference phylogenetic approaches and calculate stratigraphic congruence indices for each. Bayesian tip-dated clock methods outperform parsimony in the case of our data set, 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.

The material of Cricetodon albanensis (Rodentia, Cricetodontinae) from La Grive-Saint Alban (carrière Milliat, fissure M) is described in detail for the first time. This species is a large sized Cricetodontinae that is characterised by having incomplete ectolophs and indistinct or very short mesolophs on the upper molars and by the presence of double metalophids and the absence of mesolophids on the lower molars. The acquisition of a dominant anterior metalophulid on the m1 has been developed independently in different lineages over time and could be related to a change in dietary habits towards more abrasive food, which may be linked to the cooling event Mi3 (circa 13.8 Ma). However, the development of this character does not seem to have been enough to prevent these Middle Miocene European cricetodontines to become extinct, while their more evolved European competitors (Hispanomys) flourished.

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.

El Cerro de los Batallones is one of the most important fossil sites of the Miocene. The quantity and preservation of its fossil remains have allowed us to describe many new mammal species. One of these is Hispanomys moralesi, a derived species of the Tribe Cricetodontini, a wide group of rodents with great importance in the faunal assemblage during the Miocene. Unlike most fossil micromammals in this fossil site, we have obtained skull remains that preserve never recorded anatomical structures in this time interval. Due to the fragility of the material, it could only be studied in detail using X-ray micro-computed tomography (µ-CT Scan). With this technique, we obtained three-dimensional models of 12 skulls of Hispanomys moralesi, in which different osteological parts have been described. In this way, we compared the morphological variation of the skull with other fossil cricetids and concluded that Batallones’ species is an opisthodont rodent, with a relatively elongated rostrum, a robust base of zygomatic arches and big-sized tooth rows. Opisthodont incisor characterises omnivore and herbivore diets in rodents. Therefore, these new findings of Hispanomys moralesi could allow us to describe the dietary affinities of this extinct species.

Extinct organisms provide crucial information about the origin and time of origination of extant groups. The importance of morphological phylogenetics for rigorously dating the tree of life is now widely recognized and has been revitalized by methodological developments such as the application of tip-dating Bayesian approaches. Traditionally, molecular clocks have been node calibrated. However, node calibrations are often unsatisfactory because they do not allow the fossil age to inform about phylogenetic hypothesis. The introduction of tip calibrations allow fossil species to be included alongside their living relatives, and the absence of molecular sequence data for these taxa to be remedied by supplementing the sequence alignments for living taxa with phenotype character matrices for both living and fossil taxa. Therefore, only phylogenetic analyses that take into account morphological characters can incorporate both fossil and extant species. Herein we present an unprecedented morphological dataset for a vast group of glirid rodents, to which different phylogenetic methodologies have been applied. We have compared the tree topologies resulting from traditional parsimony and Bayesian phylogenetic approaches and calculate stratigraphic congruence indices for each. Bayesian tip-dated clock methods seem to outperform parsimony with our dataset. The strict consensus tree recovered by tip dating invalidates the classic classification and allows dates to be proposed for the divergence and origin of the different clades.