palevol comptes rendus 2021  20  22 Directeurs de la publication / Publication directors : Bruno David, Président du Muséum national d’Histoire naturelle Étienne Ghys, Secrétaire perpétuel de l’Académie des sciences Rédacteurs en chef / Editors-in-chief : Michel Laurin (CNRS), Philippe Taquet (Académie des sciences) Assistante de rédaction / Assistant editor : Adenise Lopes (Académie des sciences ; cr-palevol@academie-sciences.fr) Mise en page / Page layout : Fariza Sissi (Muséum national d’Histoire naturelle ; fariza.sissi@mnhn.fr) Révisions linguistiques des textes anglais / English language revisions : Kevin Padian (University of California at Berkeley) Rédacteurs associés / Associate editors (*, took charge of the editorial process of the article/a pris en charge le suivi éditorial de l’article) : Micropaléontologie/Micropalaeontology Maria Rose Petrizzo (Università di Milano, Milano) Paléobotanique/Palaeobotany Cyrille Prestianni (Royal Belgian Institute of Natural Sciences, Brussels) Métazoaires/Metazoa Annalisa Ferretti (Università di Modena e Reggio Emilia, Modena) Paléoichthyologie/Palaeoichthyology Philippe Janvier (Muséum national d’Histoire naturelle, Académie des sciences, Paris) Amniotes du Mésozoïque/Mesozoic amniotes Hans-Dieter Sues (Smithsonian National Museum of Natural History, Washington) Tortues/Turtles Juliana Sterli (CONICET, Museo Paleontológico Egidio Feruglio, Trelew) Lépidosauromorphes/Lepidosauromorphs Hussam Zaher (Universidade de São Paulo) Oiseaux/Birds Eric Buffetaut (CNRS, École Normale Supérieure, Paris) Paléomammalogie (mammifères de moyenne et grande taille)/Palaeomammalogy (large and mid-sized mammals) Lorenzo Rook (Università degli Studi di Firenze, Firenze) Paléomammalogie (petits mammifères sauf Euarchontoglires)/Palaeomammalogy (small mammals except for Euarchontoglires) Robert Asher* (Cambridge University, Cambridge) Paléomammalogie (Euarchontoglires)/Palaeomammalogy (Euarchontoglires) K. Christopher Beard (University of Kansas, Lawrence) Paléoanthropologie/Palaeoanthropology Roberto Macchiarelli (Université de Poitiers, Poitiers) Archéologie préhistorique/Prehistoric archaeology Marcel Otte (Université de Liège, Liège) Référés / Reviewers : https://sciencepress.mnhn.fr/fr/periodiques/comptes-rendus-palevol/referes-du-journal Couverture / Cover : Scanning electronic microscope (SEM) micrographs of early Miocene Cricetidae from different sites of the Vallès-Penedès Basin. Democricetodon hispanicus Freudenthal, 1967 Comptes Rendus Palevol est indexé dans / Comptes Rendus Palevol is indexed by: – Cambridge Scientific Abstracts – Current Contents® Physical – Chemical, and Earth Sciences® – ISI Alerting Services® – Geoabstracts, Geobase, Georef, Inspec, Pascal – Science Citation Index®, Science Citation Index Expanded® – Scopus®. 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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 diff.pub@mnhn.fr / https://sciencepress.mnhn.fr Académie des sciences, Institut de France, 23 quai de Conti, 75006 Paris. © Publications scientifiques du Muséum national d’Histoire naturelle / © Académie des sciences, Paris, 2021 ISSN (imprimé / print) : 1631-0683/ ISSN (électronique / electronic) : 1777-571X Dispersal and early evolution of the first modern cricetid rodents in Western Europe: new data from the Vallès-Penedès Basin (Catalonia) Sílvia JOVELLS-VAQUÉ Isaac CASANOVAS-VILAR Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, C/de les Columnes, s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona (Spain) silvia.jovells@icp.cat isaac.casanovas@icp.cat (corresponding author) Submitted on 3 December 2019 | Accepted on 27 April 2020 | Published on 7 June 2021 urn:lsid:zoobank.org:pub:AD9A368B-11D2-49D9-A1C0-3451D239361D Jovells-Vaqué S. & Casanovas-Vilar I. 2021. — Dispersal and early evolution of the first modern cricetid rodents in Western Europe: new data from the Vallès-Penedès Basin (Catalonia). Comptes Rendus Palevol 20 (22): 401-439. https://doi.org/10.5852/cr-palevol2021v20a22 ABSTRACT Modern cricetids originated in Asia and dispersed into Western Europe by the end of the early Miocene, where they quickly became major components of the rodent faunas. Here we review the early Miocene rodent record of the genera Democricetodon Fahlbusch, 1964 and Megacricetodon Fahlbusch, 1964 in the Vallès-Penedès Basin (Catalonia, Spain). Democricetodon is represented by four species in the studied sites (D. hispanicus Freudenthal, 1967, D. cf. decipiens (Freudenthal & Daams, 1988), D. gracilis Fahlbusch, 1964 and a large-sized undetermined species) and Megacrice- todon by one (M. primitivus (Freudenthal, 1963)). The cricetid succession bears several similarities KEY WORDS with that of the nearby Calatayud-Montalbán Basin (East-Central Spain) to the point that the same Democricetodon, Megacricetodon, detailed local biostratigraphy could be extended to the Catalan basin. The rare presence of certain early Miocene, Democricetodon species (D. gracilis) and other small mammal taxa also reveal affinities with regions Iberian Peninsula, biostratigraphy, beyond the Iberian Peninsula and indicate that the Vallès-Penedès Basin was more humid and forested western Europe. than inland Iberian basins during the early Miocene. RÉSUMÉ Dispersion et évolution précoce des premiers rongeurs cricétidés modernes en Europe occidentale : nouvelles données du bassin de Vallès-Penedès (Catalogne). Les cricétidés modernes sont originaires d’Asie et se sont dispersés en Europe occidentale à la fin du Miocène inférieur. Nous passons ici en revue les signalements du Miocène inférieur des genres Democricetodon Fahlbusch, 1964 et Megacricetodon Fahlbusch, 1964 du bassin du Vallès-Penedès (Catalogne, Espagne). Democricetodon est représenté par quatre espèces (D. hispanicus Freudenthal, 1967, D. cf. decipiens (Freudenthal & Daams, 1988), D. gracilis Fahlbusch, 1964 et une espèce indéterminée de grande taille) et Megacricetodon par une (M. primitivus (Freudenthal, 1963)). La succession de cricétidés présente plusieurs similarités avec celle du bassin voisin de Calatayud-Montalbán (Centre- MOTS CLÉS Est de l’Espagne), à tel point que la même biostratigraphie locale détaillée pourrait être étendue au Democricetodon, Megacricetodon, bassin catalan. La rare présence de certaines espèces de Democricetodon (D. gracilis) ainsi que celle Miocène inférieur, d’autres taxons de petits mammifères révèle également des affinités avec des régions au-delà de la péninsule Ibérique, biostratigraphie, Péninsule Ibérique et indique que le bassin du Vallès-Penedès était plus humide et boisé que les bassins Europe occidentale. ibériques intérieurs durant le Miocène inférieur. COMPTES RENDUS PALEVOL • 2021 • 20 (22) © Publications scientifiques du Muséum et/and Académie des sciences, Paris. www.cr-palevol.fr 401 Jovells-Vaqué S. & Casanovas-Vilar I. INTRODUCTION France and Spain (for an updated review see Oliver & Peláez Campomanes 2016). From then on, both genera diversify in The cricetids Democricetodon Fahlbusch, 1964 and Megacrice- the different geographic regions and become characteristic of todon Fahlbusch, 1964 are major components of the early and the middle Miocene faunas. middle Miocene rodent faunas of Eurasia, to the point that Early modern cricetid faunas have been particularly well one or two species of these cricetids may account for more studied in the Calatayud-Montalbán Basin of Spain, where than 90% of the recovered rodent remains in some sites (e.g. they are very well represented (Daams & Freudenthal 1988; Daams & Freudenthal 1988). Both are incredibly diverse, Freudenthal & Daams 1988; Van der Meulen et al. 2003; each one comprising more than 30 recognized species with a Oliver Pérez 2015; Oliver & Peláez-Campomanes 2014, relatively limited stratigraphic and geographic range. In turn, 2016). These genera are also common in other Iberian basins, these species define multiple independent lineages character- including the Foia de Bunyol (Daams & Freudenthal 1974; ized by marked temporal changes in size and morphology. Adrover et al. 1987) or the Riu Magre Basin (Ruiz-Sánchez Therefore, it is not surprising that these cricetids have been et al. 2003), both in Valencia, and the Ebro Basin (Ruiz-Sánchez widely used in biostratigraphy and biochronology. Indeed, et al. 2003; Suárez-Hernando 2017). However, these include the first appearance and evolution of different Democriceto- relatively few fossil sites and the recovered material of the don and Megacricetodon species in combination with other earliest modern cricetids is generally scarce. The coastal Vallès- rodent taxa is used to define most of the local biozones for Penedès Basin, located in Catalonia, northeast Spain comprises the early and middle Miocene in several European regions a rich record covering most of the Miocene (Casanovas-Vilar (Daams et al. 1999; Abdul Aziz et al. 2008, 2010; Kälin & et al. 2016b). Until recently, the early Miocene successions of Kempf 2009; Van der Meulen et al. 2012; Casanovas-Vilar this area had been little studied in comparison to middle and et al. 2016a; Prieto & Rummel 2016). They also play an late Miocene ones, yet these included a handful of remarkable important role in continental biochronology, the first appear- sites. However, recent survey and sampling campaigns have ance of these genera (as well as that of certain widely dispersed focused on the early Miocene part of the record, resulting species) marking the boundaries between higher-ranking in the recovery of rich fossil microvertebrate samples from biochronological units, such as mammal ages in China (Qiu “classical” and newly-discovered localities. About 30 sites are et al. 2013) and MN (Mammal Neogene) zones in Europe known to date, and these have delivered more than 2.000 iden- (Mein 1999; Agustí et al. 2001; Hilgen et al. 2012). tifiable rodent cheek teeth. Only a small part of this material Democricetodon and Megacricetodon are two of the first genera has been described and biostratigraphical studies are now in commonly referred to as “modern cricetids” (see comments course. Cricetids are common in almost all sites and deserve on classification in section Classification of the cricetids) to special attention because of their biostratigraphical implica- distinguish them from older muroid taxa that had characterized tions. Detailed descriptions of the cricetid fauna of Can Martí the Oligocene faunas of Eurasia (e.g. the genera Eucricetodon Vell (Agustí 1983), Les Cases de la Valenciana (Jovells-Vaqué Thaler, 1966, Pseudocricetodon Thaler, 1969 and Melissiodon et al. 2018) and els Casots (Jovells-Vaqué et al. 2017) sites have Schaub, 1920). Modern cricetids, namely the genera Spanocri- been published, as well as a review of the genus Melissiodon cetodon Li, 1977 and Primus de Bruijn et al., 1981, first (Jovells-Vaqué & Casanovas-Vilar 2018). However, most of appear simultaneously in Anatolia and Pakistan, respectively, the cricetid material had not been studied yet. Here we identify by the latest Oligocene (Theocharopoulos 2000). The old- and describe all the early Miocene material belonging to the est Democricetodon are recorded slightly later, by the earliest genera Democricetodon and Megacricetodon. This allows for a Miocene (c. 22 Ma) from Anatolia, Pakistan and China (Meng refined biostratigraphical correlation of the studied localities et al. 2003; Maridet et al. 2011; Flynn & Wessels 2013; Flynn based on high-resolution local zonations defined for other et al. 2013). On the other hand, Megacricetodon is also first Iberian basins. Finally, the occurrence of certain species with recorded from the early Miocene (c. 19-17 Ma) of Anatolia Central European affinities and their paleobiogeographical (Wessels et al. 2001). In western Europe, part of the early implications are also discussed. Miocene (MN3, from about 19.5 to 17 Ma) is devoid of cricetids because of the extinction of all the Oligocene taxa, except for the rare occurrence of Melissiodon. This interval, GEOLOGICAL SETTING AND AGE called the “cricetid vacuum”, comes to an abrupt end with de dispersal of the modern cricetids Democricetodon and The record of the Vallès-Penedès Basin (Barcelona, Catalo- Megacricetodon (Daams & Freudenthal 1989). Democriceto- nia) covers most of the Miocene, ranging from the Ramblian don is already recorded during the late MN3 (17 Ma) in the (c. 20 Ma) to the middle Turolian (c. 7 Ma) European land Calatayud-Montalbán Basin of east-central Spain, where it mammal ages (Casanovas-Vilar et al. 2016b). This basin is occurs in low numbers (Van der Meulen et al. 2003, 2012). an elongated half-graben parallel to the coastline bounded The first common occurrence of this genus is used to define the by the Catalan Littoral Ranges (Fig. 1) that was formed in lower boundary of biozone MN4 (Agustí et al. 2001; Hilgen the context of the opening of the northwestern Mediter- et al. 2012). In its turn, Megacricetodon is recorded slightly ranean during the late Oligocene (Cabrera & Calvet 1996; later in western Europe, its first occurrences corresponding Roca et al. 1999; Cabrera et al. 2004). Major features of the to MN4 sites from Greece, the Czech Republic, Switzerland, stratigraphic record of the Vallès-Penedès were controlled 402 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe IBERIAN PENINSULA N Igualada SPAIN Terrassa Sabadell Martorell Sant Cugat del Vallès Sant Sadurní d’Anoia MAMMAL SITES Ramblian (MN3) Barcelona Vilafranca early Aragonian (MN4) del Penedès middle Aragonian (MN5) Fault Inferred fault Normal fault CLA Thrustfault 10 km Vilanova i la Geltrú NEOGENE-QUATERNARY Quaternary sediments: alluvial fan and fluvial facies Neogene volcanic rocks (basalts) EBRO FORELAND BASIN INFILL Pliocene marine units: transitional tomarine breccias, Paleogene (conglomerates, sandstones, lutites) sandstones, marls PALEOZOIC BASEMENT AND MESOZOIC COVER Upper Continental Unit (Langhian-Tortonian): alluvial fan facies Mesozoic (carbonates, sandstones, lutites) Marine and Transitional Units 2 (late Burdigalian-Langhian): coralgal carbonates facies Hercynian intrusive rocks (mostly granitoids) Marine and Transitional Units 1 (late Burdigalian-Serravallian): malts, silts, bioclastic sands and sandstones Paleozoic sedimentary and metamorphic rocks Lower Continental units ( early and late Burdigalian): alluvial fan and shallow lacustrine facies Fig. 1. — Geographical location and simplified geological map of the Vallès-Penedès Basin indicating the main mammal localities ranging from MN3 to MN5 (modified from Casanovas-Vilar et al. 2016a, b). For locality acronyms see section Abbreviations of the main text. by tectonic activity of the main limiting faults and sea level alluvial fan facies sourced from the surrounding reliefs as well changes in the western Mediterranean. The sediment infill as occasional shallow lake deposits consisting of carbonates, of the basin is mostly continental, except for brief episodes evaporites, carbons and mostly greyish to greenish lutites of marine transgression that occurred during the latest early (Cabrera 1979, 1981; Cabrera et al. 1991; Agustí et al. 1985; Miocene and the beginning of the middle Miocene. The Casanovas-Vilar et al. 2011, 2016b; de Gibert & Casanovas- studied material was recovered from early and early middle Vilar 2011). Lake deposits include the richest early Miocene Miocene localities that have been dated by means of litho- and fossil sites in the basin (Fig. 2). The Subirats lacustrine unit, biostratigraphy (Agustí 1982; Agustí et al. 1985; Casanovas- extending a few kilometers between the towns of Subirats Vilar et al. 2011, 2016b). and Gelida (Cabrera 1979, 1981; Agustí & Cabrera 1980; The early Miocene record of the Vallès-Penedès starts at Cabrera et al. 1991; Casanovas-Vilar et al. 2011; Jovells-Vaqué the Ramblian, biozone MN3 (c. 19.5-17.2 Ma). Ramblian et al. 2017, 2018) includes the localities of Les Cases de la sites correspond to the so-called “cricetid vacuum” (Daams & Valenciana, Can Martí Vell (levels 1 to 3) and Els Casots Freudenthal 1989) and do not include cricetids other than (levels 72, 73 and 74). Each one of these levels has delivered Melissiodon (Casanovas-Vilar et al. 2011, 2016b; Jovells- more than 150 identifiable rodent teeth, about half of them Vaqué & Casanovas-Vilar 2018). The oldest sites recording cricetids. At Els Casots area the Subirats lacustrine unit lies the modern cricetids Democricetodon and Megacricetodon are directly over of the pre-Miocene basement of the basin, there correlated to the early Aragonian, biozone MN4 (Casanovas- consisting of Cretaceous carbonates (Fig. 1). The nearby site Vilar et al. 2011, 2016b; Jovells-Vaqué et al. 2017, 2018). of Cal Sutxet corresponds to a yellowish lutite layer which These belong to the Lower Continental Units, a lithostrati- occasionally includes sharp boulders eroded from the basement. graphical formation that crops out near the southern mar- Other sites belonging to the Lower Continental Complexes gin of the basin (Fig. 1). These units consist of intensely red include, el Canyet, Sant Mamet and Les Escletxes del Papiol, COMPTES RENDUS PALEVOL • 2021 • 20 (22) 403 Jovells-Vaqué S. & Casanovas-Vilar I. which are all located in the distal, mudstone-dominated facies between 2011 and 2017 (Les Cases de la Valenciana, Can Martí of alluvial fan systems sourced from the southern reliefs (Fig. 2). Vell, Cal Sutxet, Sant Mamet, Can Cabanes W, Riera del Morral Between 17 and 15 Ma several sea level changes took place in 1). Agustí (1983) collected a remarkably rich sample from the context of the Mid-Miocene Climatic Optimum (Zachos Can Martí Vell, but this material could not be found within et al. 2001) and, since the southwestern basin end was open the MGB collections. The rich sample recovered during the to the sea, the Vallès-Penedès was flooded repeatedly. At 2015 field campaign at this site is described instead. Similarly, least three episodes of marine transgression and regression Agustí (1981) described a few Megacricetodon primitivus are recorded: late Burdigalian, Langhian and early Serraval- (Freudenthal, 1963) specimens from Can Julià 6, a nearby lian (Cabrera et al. 1991; Cabrera & Calvet 1996; de Gib- classical site already reported in Crusafont et al. (1955), which ert & Casanovas-Vilar 2011; Casanovas-Vilar et al. 2016b). we could not find within the ICP collections. The material The sequences deposited at this time define the Marine and from old collections, including the specimens from El Canyet Transitional Units (Fig. 2). Sediments corresponding to the as well as some from Sant Mamet, probably derives from oldest late Burdigalian (latest early Miocene) transgression surface surveys. Dental terminology follows Oliver & Peláez- only crop out in the Penedès sector of the basin, where they Campomanes (2013), whereas measurement method is after define the Vilobí evaporitic unit (Fig. 2). This consists of Daams & Freudenthal (1988: 42, fig. 1). Morphotype coding gypsum which was deposited in coastal playa lakes (sabkha) follows Van der Meulen et al. (2003) and Maridet (2003) for as well as littoral sandstones and lutites (Ortí & Pueyo 1976; Democricetodon and Oliver & Peláez-Campomanes (2013) for Magné 1978; Agusti et al. 1990). The Vilobí del Penedès Megacricetodon with slight modifications. All measurements are fossil site is particularly rich, having delivered more than given in millimeters and were taken with an optical micrometer 300 identifiable rodent specimens, and is located in a ter- to the nearest 0.01 mm. Estimated measurements (because of rigenous coquina full of ostreids which presumably indicate minor damage or distortion) are between brackets, whereas “>” bay deposits (Casanovas-Vilar et al. 2019). The Langhian indicates that the measurement cannot be confidently taken (beginning of the middle Miocene) transgression is the best but certainly exceeded the reported value. Summary statistics represented marine episode. At that time, marine environ- and scatterplots were performed using the R software (R Core ments extended up to the Vallès sector, with the development Team 2017). MN zone definitions for western Europe follow of shallow marine and transitional deposits (Figs 1; 2). On Agustí et al. (2001), whereas age boundaries are as defined in the other hand, carbonate coralgal shelf deposits (including Hilgen et al. (2012). minor fringing reefs), marine bay and transitional fan-delta siliciclastic systems persistently occupied the Penedès sector, Classification of the cricetids directly connected to the sea (Cabrera et al. 1991; Cabrera & The contents and definition of the cricetid family are a matter Calvet 1996; de Gibert & Casanovas-Vilar 2011; Casanovas- of debate. McKenna & Bell (1997) ranked cricetids as a mere Vilar et al. 2016b). In the Vallès sector, the sites of Riera del subfamily within the huge Muridae family, but most classifications Morral and Can Cabanes W are both located in transitional upgrade the Cricetidae to the family rank (e.g. Hartenberger 1998; Langhian deposits associated with marine invertebrates and Steppan et al. 2004; Wilson & Reeder 2005; Fabre et al. 2012, ichnofauna (Fig. 2; de Gibert & Robles 2005; Casanovas- 2015; Steppan & Schenk 2017; D’Elía et al. 2019). Recent Vilar & Jovells-Vaqué 2017). Finally, Les Escletxes del Papiol molecular phylogenetic analyses recognize the Cricetidae as a and Sant Mamet sites, also in the Vallès sector, are located just distinct family from the Muridae, both being part of the larger a few meters below Langhian marine deposits, but still placed Muroidea superfamily (Steppan et al. 2004; Fabre et al. 2012, in distal alluvial fan facies belonging to the Lower Continental 2015; Steppan & Schenk 2017). The Cricetidae include the extant Units (Fig. 2; Agustí et al. 1985; Casanovas-Vilar et al. 2011, hamsters (Cricetinae), voles and lemmings (Arvicolinae) as well as 2016b). Detailed lithostratigraphical data, as well as associated the three subfamilies of New World rats and mice (Neotominae, faunal data, allow for a confident stratigraphical succession of Sigmodontinae and Tylominae; Steppan et al. 2004; Fabre the studied localities which is further reinforced by biostrati- et al. 2012, 2015; Steppan & Schenk 2017). Other muroid graphical results (Fig. 2; see also section Biostratigraphy and groups (e.g. Nesomyidae, Spalacidae, Platacanthomyidae) that correlations). However, the age of these sites will be surely had been included within the cricetids are now considered refined after ongoing magnetostratigraphical studies. separate families. The phylogenetic relationships of most extinct genera are uncertain, and even their ascription to the Cricetidae is questionable, specially for older forms which might be as closely MATERIAL AND METHODS related to extant cricetids as they are to other muroid families. As far as the genera Democricetodon and Megacricetodon are The described material is housed at the Institut Català de concerned, these are generally considered to be closer to extant Paleontologia Miquel Crusafont in Sabadell (Barcelona, Spain) cricetids (Flynn 1985; Kälin 1999; Lindsay 2008). Democricetodon and the Museu de Ciències Naturals de Barcelona (MGB). in particular has been related to extant cricetines and is generally Collection numbers are given in the main text as well as for placed in a subfamily within the cricetids (the Democricetinae) figured specimens. Almost all the material was recovered after together with the closely-related North American genus (or systematic screen-washing campaigns during the early 1990s subgenus) Copemys Wood, 1936 (Lindsay 2008). On the other (Els Casots, Vilobí del Penedès, Les Escletxes del Papiol) and hand, Megacricetodon is generally considered to be closer to 404 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Baix Penedès Alt Penedès Baix Llobregat Valès Occ. Ser. MN5 Langh. CLA MN4 Burfigalian MN3 Aquit. Proximal alluvial Proximal alluvial Terrigenous bau fan facies fan facies (nothern-sourced) (southern-sourced) Carbonate platform Distal alluvial Lacustrine carbonates fan facies Litoral sabkha and evaporites MARINE AND CONTINENTAL FACIES TRANSITIONAL FACIES Fig. 2. — Generalized stratigraphic framework of the early to early middle Miocene sedimentary record of the Vallès-Penedès Basin showing the main lithostrati- graphic units and their relationships (modified after Cabrera et al. 1991 and De Gibert & Casanovas Vilar 2011). Note the different early Miocene lacustrine systems (including most localities) as well as the marine deposits associated to the Langhian transgression. For locality acronyms see section Abbreviations of the main text. CMV1 Can Martí Vell – level 1; the Muridae (including murines, gerbillines and other minor CMV2 Can Martí Vell – level 2; extant subfamilies) since its dental morphology resembles that CMV3 Can Martí Vell – level 3; of early murids such as Potwarmus Lindsay, 1988 and Antemus CS72 els Casots – level 72; Jacobs, 1977 (Jacobs 1977; Flynn 1985; Lindsay 1988; Jacobs & CS73 els Casots – level 73; Down 1994). Megacricetodon is sometimes included within a CS74 els Casots – level 74; separate subfamily (Megacricetodontinae; e.g. Lindsay 1988, CSU Cal Sutxet; 2008; Wessels et al. 2001; Wessels 2009) that may form a EC El Canyet; monophyletic clade with the Muridae (Flynn 1985). In any LCV Les Cases de la Valenciana – level 1 (classical site); case, most authors agree in including both generea within the MC Molí de Can Calopa; MOR1 Riera del Morral – level 1; cricetids in a broad sense. This is the classification scheme PA Les Escletxes del Papiol; followed here, although we acknowledge that Megacricetodon SAB1 Sant Andreu de la Barca – level 1; may be closer to murids. SM Sant Mamet – level 1; TFR1 Turó de les Forques – level 1; Abbreviations VI Vilobí del Penedès. Institutions IPS Institut de Paleontologia de Sabadell (now ICP, Institut Other abbreviations Català de Paleontologia Miquel Crusafont), Sabadell; MN European Mammal Neogene zones. MGB Museu de Geologia de Barcelona (now Museu de Ciències Naturals de Barcelona), Barcelona; “V” preceding collection number indicates that the speci- men belongs to the Villalta Collection, donated to the SYSTEMATIC PALEONTOLOGY museum in the 1980s. Order RODENTIA Bowdich, 1821 Measurements Family Cricetidae Fischer von Waldheim, 1817 L anteroposterior length; Genus Democricetodon Fahlbusch, 1964 W labiolingual width. Democricetodon hispanicus Freudenthal, 1967 Localities (Figs 3, 4; Table 1; Appendix 2) CB Costablanca; CCW Can Cabanes W; A catalog of the studied material and measurements is CJ Can Julià; given in Appendix 1. See Table 1 for summary statistics COMPTES RENDUS PALEVOL • 2021 • 20 (22) 405 Jovells-Vaqué S. & Casanovas-Vilar I. and comparisons. See also Figure 3 for measurements and is no trace of the protosinus and the lingual anteroloph is comparisons. Morphotype frequency tables are given in reduced to just a tiny lingual cingulum. The protolophule is Appendix 2. simple and connects with the anterior part of the paracone. There is no mesoloph and the metalophule is simple and Description short. The posteroloph is well developed and closes the very M1 (Fig. 4A, B) narrow posterosinus. The metacone is reduced and can be All studied specimens have a simple anterocone, except one slightly displaced to the lingual side (e.g. IPS45008, Fig. 4A). from CS74 in which this cusp is slightly subdivided (IPS45001 The mesosinus is closed by a thin low ridge continuous with from CS74). The arms of the anteroloph are well developed the posteroloph. The sinus is highly reduced, transverse and and close the anterior valleys. The anterolophule is simple and narrow, being closed by a low cingulum descending from placed somewhat lingually. However, IPS45002 from CS74 the hypocone. shows a forked anterolophule with both arms reaching the anterocone. The protosinus is relatively reduced. The protolo- m1 (Fig. 4C) phule normally consists of a single posterior arm that joins the The anteroconid is simple and rounded. The anterior valleys entoloph behind the protocone, but it may also be double with are closed in some specimens from CS72 and CS74 (e.g. a better-developed anterior arm (Appendix 2C). In IPS45008 IPS19481, Fig. 4C) by the arms of the anterolophid, which from CS74 (Fig. 4A), the anterior arm is interrupted before are quite low. The mesolophid is variable in length, ranging reaching the paracone. In some specimens, there is a short from short to long, although a medium-length mesolophid is ectoloph on the paracone. The mesoloph varies from short to most common. The sinusid is wide and generally points for- long in all the studied specimens (Appendix 2E), however this wards, although it may be transverse in a few specimens. The ridge is most commonly long (LCV1, CMV3) or of medium sinusid is closed by a low cingulum. The mesosinusid is more length (CS74). The metalophule consists of a posterior arm often open except in one specimen from CMV1 (IPS96841) only that connects the metacone with the posteroloph just in which it is closed by the metaconid ridge (Appendix 2M). behind the hypocone. In one specimen, V7846 from el Canyet, The hypolophulid is short and merges with the entolophid the metalophule is double with an anterior arm merging the anteriorly to the hypoconid. The metalophulid, is absent in entoloph anteriorly to the hypocone. In IPS19491 from IPS19481 from CS74 (Fig. 4C). In the other specimens, the CS74 the metalophule is completely absent. The posterosinus metalophulid is very short and may be either transverse or is highly reduced and closed by the posteroloph. The sinus slightly anterior to the protoconid. The posterolophid closes is closed by a well-developed cingulum originating from the the posterosinusid, reaching the posterior wall of the entoconid. hypocone in all studied specimens. m2 (Fig. 4D) M2 (Fig. 4A, B) The lingual anterolophid and the anterosinusid are reduced The anteroloph shows a long and high labial arm that closes so that they completely disappear with moderate dental wear, the narrow anterosinus. The protosinus is vestigial and is as in IPS86281 from CSU. The protosinusid is closed by also closed by a much lower lingual arm of the anteroloph. the labial arm of the anterolophid. The mesolophid is highly The protolophule generally consists of an anterior arm and a variable, ranging from short to long but also being frequently posterior one (Appendix 2G). The latter merges the entoloph absent (Appendix 2Q). In all the remaining morphological posteriorly to the protocone in most cases, however, in a few features the m2 resemble the m1. specimens the posterior arm of the protolophule is incomplete (for example in IPS45008 and IPS19491 from CS74, Fig. 4A, m3 (Fig. 4E) B), being interrupted before merging with the paracone.The The anterior valleys are very reduced to the point that in some ectoloph on the paracone is variable being present or absent specimens the lingual arm of the anterolophid is vestigial, the (Appendix 2H). When present, the ectoloph does not merge anterosinusid thus being absent. The metaconid connects with with the mesoloph. The length of the mesoloph ranges from the anterolophid by means of an extremely short metalophulid. long to short but is more commonly long (Appendix 2I). The mesolophid is absent in all the specimens. The entoconid The sinus is transverse and closed by a cingulum originating is highly reduced being integrated into the hypolophulid. The from the hypocone. The metalophule is short and simple, sinusid is posteriorly directed. This valley is generally open joining the entoloph anteriorly to the hypocone. However, in the CS74 specimens as well as in IPS85740 from CCW in V7846 from EC the metalophule is simple and transverse while in those from LCV and CMV3 it is closed by a low to the hypocone, and in IPS19475 from CS74 it is double cingulid. The posterosinusid is reduced and completely closed with similarly well-developed anterior and posterior arms. by the posterolophid which is very high. The mesosinusid is The posteroloph closes the posterosinus, while the mesosinus also closed by a high ridge departing from the posterior wall is closed by a low cingulum. of the metaconid. M3 (Fig. 4A) Comparisons and remarks The anterosinus is almost closed by the well developed labial The Vallès-Penedès specimens fit within the size and morpho- arm of the anteroloph in all the studied specimens. There logical range of Democricetodon hispanicus Freudenthal, 1967 406 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe D. mutilus M1 m1 D. moralesi 1.6 1.2 D. mutilus D. moralesi D. hispanicus 1.4 D. hispanicus 1.0 W 1.2 D. decipiens W D. decipiens 0.8 D. franconicus 1.0 D. franconicus D. gracilis D. gracilis 1.0 0.6 1.2 1.4 1.6 1.8 2.0 2.1 1.2 1.4 1.6 1.8 L L m2 D. mutilus M2 1.4 D. decipiens 1.4 D. mutilus D. hispanicus 1.2 D. hispanicus D. moralesi 1.2 D. moralesi W 1.0 D. decipiens W 1.0 D. franconicus D. franconicus D. gracilis 0.8 0.8 D. gracilis 0.6 0.6 1.0 1.2 1.4 1.6 1.8 1.0 1.2 1.4 1.6 1.8 L L M3 m3 D. mutilus D. mutilus 1.2 1.2 D. decipiens D. hispanicus D. moralesi CMV1 CMV2 1.0 1.0 CMV3 CS72 D. moralesi CS73 W W D. decipiens CS74 CSU 0.8 0.8 EC D. hispanicus Democricetodon sp. LCV1 D. gracilis D. cf. decipiens D. gracilis MOR1 D. gracilis PA D. hispanicus SM 0.6 0.6 VI 0.6 0.8 1.0 1.2 0.8 1.0 1.2 1.4 L L Fig. 3. — Length/width scatter plot for Democricetodon hispanicus Freudenthal, 1967, D. gracilis Fahlbusch, 1964, D. cf. decipiens (Freudenthal & Daams, 1988) and D. sp. 4 from different Vallès-Penedès sites. Locality acronyms and measurement abbreviations are explained in section Abbreviations of the main text. For details on the measurement methods see section Material and methods, and Daams & Freudenthal (1988: 42, Fig. 1). The ellipses show the 95% confience interval for the following species: Democricetodon hispanicus from Villafeliche 2A, D. moralesi Van der Meulen et al., 2003 from La Col D and D. franconicus Fahlbusch, 1966 from La Col D (data provided by P. Peláez-Campomanes, Museo Nacional de Ciencias Naturales, Madrid), D. gracilis and D. mutilus Fahlbusch, 1964 from Sandelzhausen (Wessels & Reumer 2009) and D. decipiens from Bunyol (Daams & Freudenthal 1974). COMPTES RENDUS PALEVOL • 2021 • 20 (22) 407 Jovells-Vaqué S. & Casanovas-Vilar I. (see Van Der Meulen et al. 2003) being remarkably similar to of Villafeliche 2A (Calatayud-Montalbán Basin), the type the material from the type locality, Villafeliche 2A (Calatayud- locality of this species (Freudenthal 1967). Villafeliche 2A is Montalbán Basin, Aragon, Spain; Freudenthal 1967). This correlated to local subzone B (estimated age 16.63 Ma; Van species is distinguished from the chronologically and geographi- der Meulen et al. 2012), thus being somewhat older than cally close species Democricetodon decipiens (Freudenthal & the Vallès-Penedès sites. Considering the evolutionary trends Daams, 1988), Democricetodon franconicus Fahlbusch, 1966 towards size increase and reduction of mesolophs/ids within and Democricetodon koenigswaldi (Freudenthal, 1963) by its the D. hispanicus-D. lacombai lineage, these differences could smaller size (Van der Meulen et al. 2003). The Vallès-Penedès well be explained by the younger age of the Vallès-Penedès samples (particularly the youngest ones such as CS73 and specimens. Indeed, when the whole intraspecific variation is CS74) overlap with upper size range of D. hispanicus, with a taken into account (and not that existing in just one site), few specimens, generally M2 and m2 being larger, well within the described material perfectly agrees with size variation in the size range of D. decipiens (Fig. 3; Appendix 1). However, D. hispanicus, although being close to the upper boundary. these specimens can be distinguished from D. decipiens by As far as the development of the mesolophid is concerned, the development of mesolophs and mesolophids, which are medium length and short mesolophids generally predominate always longer in D. hispanicus (Van der Meulen et al. 2003). in most D. hispanicus from the Calatayud-Montalbán sites Besides its smaller size, D. hispanicus further differs from (see Van der Meulen et al. 2003). D. koenigswaldi by its less developed metalophule on the M2 (Van der Meulen et al. 2003). In D. koenigswaldi the metalo- phule is frenquently double or merges the entoloph posterior Democricetodon gracilis Fahlbusch, 1964 to the hypocone, whereas in the studied specimens this ridge (Figs 3, 4; Table 1; Appendix 2) is generally simple and merges the entoloph anterior to the hypocone. Similarly, the protolophule is predominantly double A catalog of the studied material and measurements is given in D. koenigswaldi, and although double protolophules occur in in Appendix 1. See Table 1 for summary statistics and com- D. hispanicus, simple ones are almost equally common (Van der parisons. See also Figure 3 for measurements and compari- Meulen et al. 2003; see also Appendix 2C, G). The described sons. Morphotype frequency tables are given in Appendix 2. specimens show all these diagnostic features that unambigu- ously indicate an ascription to D. hispanicus, which would be Description the first Democricetodon species to be recorded in the basin. M1 (Fig. 4F) Several of the morphological trends that have been described The anterocone is simple in the two specimens from SM for the Calatayud-Montalbán material (Van der Meulen (IPS10342 and IPS103751). The protolophule is always double. et al. 2003) can also be recognized in the Vallès-Penedès. In the Both arms of the protolophule reach the paracone, but the upper molars simple protolophules become more common in anterior arm is equally developed or is thinner than the poste- younger samples, such as CS74 (see Appendix 2C). Similarly, rior and weakly reaches the paracone (Fig. 4F; Appendix 2C). the spur on the paracone is more frequent in these sites. The The anterosinusid is closed by the labial anteroloph. On the mesoloph is predominantly long in the older sites, whereas contrary, the low lingual anteroloph does not reach the pro- in younger ones we may also find medium-sized and short tocone, resulting in an open protosinus. The anterolophule is mesolophs (Appendix 2E, I). Compared to the upper molars simple. The paracone shows a very short posterior ectoloph in the mesolophids are always more reduced than the mesolophs, the two specimens. The mesoloph is long, but it does not reach particularly for the m2. In the younger sites (CS74, CCW, SM) the labial margin of the teeth (Appendix 2E). The mesosinus the mesolophid is predominantly short or absent on the m2 and is closed by a low cingulum. The metalophule is simple and short to medium-length on the m1. The only remarkable differences merges the entoloph posterior to the hypocone. The sinus is with the Calatayud-Montalbán samples include the slightly larger transverse and closed by a low and arched cingulum originat- size of some Vallès-Penedès specimens and the higher occurrence ing from the hypocone. of double protolophules on the M2. In the Calatayud-Montalbán basin, double protolophules occur in a maximum of 25% of the M2 (Fig. 4G) specimens (Villafeliche 2A; see Freudenthal & Daams 1988; Two specimens have been recovered from SM (IPS103752 and Van der Meulen et al. 2003), whereas in the Vallès-Penedès IPS103753). The labial anteroloph is well developed whereas material they predominate (Appendix 2C, G). Another mor- the lingual one is vestigial. The protosinus is also vestigial. phological feature that only occurs in the Vallès-Penedès is the In one specimen the protolophule is double (IPS103753, presence of forked anterolophules, which occur in one specimen Fig. 4G), whereas in the other one it is simple and transverse from LV1 and another one from CS74. Agustí (1981, 1983) (IPS103752, Appendix 2G). Both molars show a very short already noted some of these features regardigng the CMV ectoloph on the paracone. The mesoloph is long but not and EC specimens, particularly stressing the larger size and reaching the labial margin of the teeth in any of the studied shorter mesolophids, and he ascribed the material to D. aff. specimens. The metalophule is simple and anterior to the hispanicus. Similarly, Agustí & Llenas (1993) also ascribed hypocone in the specimen IPS103752, and it is not observable the material of CS73 and CS74 to D. aff. hispanicus. These in IPS103753, Fig. 4G), likely because it was very low and authors only compared the described specimens to those this area is covered with encrusted sediment. 408 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe A B 1 mm C D E F G H I J K L M N O P Q R S T U V W X Y Z A’ Fig. 4. — Scanning electronic microscope (SEM) micrographs of early Miocene Cricetidae from different sites (indicated in parentheses) of the Vallès-Penedès Basin. Democricetodon hispanicus Freudenthal, 1967: A, IPS45008, right M1-M3 (reversed) (CS74); B, IPS19491, right M1-M2 (reversed) (CS74); C, IPS19481, left m1 (CS74); D, IPS88884 left m2 (LCV1); E, IPS86457, right m3 (reversed) (LCV1). Democricetodon gracilis Fahlbusch, 1964; F, IPS103751 left M1 (SM); G, IPS103753 left M2 (SM); H, IPS103754 left m1 (SM); I, IPS103755, right m2 (reversed) (SM); J, IPS105145 left m3 (SM). Democriceton cf. decipiens (Freudenthal & Daams, 1988); K, IPS19548, left M1(VI); L, IPS87106 left M2 (VI); M, IPS86941 left M3 (VI); N, IPS87109 left m1 (VI); O, IPS86387 left m2 (PA). Democriceton sp. 4; P, IPS88868 left M2 (MOR1); Q, IPS 45052 left m1 (CS73); R, IPS105188 left m2 (MOR1). Megacricetodon primitivus (Freudenthal, 1963); S, IPS86997 left M1 (VI); T, IPS86432 right M1 (reversed) (LCV1); U, IPS86435 left M2 (LCV1); V, IPS87022 left M2 (VI); W, IPS19479 left M3 (CS74); X, IPS44950 right m1- m2 (reversed) (CS74); Y, IPS86440 left m1(LCV1); Z, IPS86444 left m2 (LCV1); A’, IPS86451 right m3 (reversed) (LCV1). For the locality acronyms see section Abbreviations of the main text. Scale bar: 1 mm. COMPTES RENDUS PALEVOL • 2021 • 20 (22) 409 Jovells-Vaqué S. & Casanovas-Vilar I. m1 (Fig. 4H) length ones on the M2; a predominantly double protolophule The anteroconid and the anterior region of the molar is on M2; a posterior metalophule on M1 and an anterior or reduced. The anteroconid is simple and round. The arms transverse one on most M2; and a short anterior region of the of the anterolophid are thin and long, closing the anterior m1 (Maridet 2003; Wessels & Reumer 2009). Neverthe- valleys. The metalophulid is simple and very short, merg- less, there are a few differences. In the material of the type ing with the anterolophulid anterior to the protoconid. The locality (Sandelzhausen, Germany, MN5) the mesolophids anterolophulid is generally low where it joins the metalo- are predominantly of medium length or short on m1 and m2 phulid and the protoconid. The mesolophid is very long and and the protolophule is generally posterior on the M1, not reaches the lingual margin of the tooth, sometimes merging partially double as in the specimens from SM (IPS103753). with a very low cingulid (Appendix 2N). The sinusid points Agustí (1981) already ascribed a few specimens from SM to slightly forward and is closed by a low cingulid descend- D. cf. gracilis and a few more to D. aff. hispanicus, further ing from the protoconid. The mesosinusid is enclosed by a remarking that they certainly belonged to a smaller and more cingulid in the SM specimen IPS103754 (Fig. 4H) whereas primitive species than the one present at EC and CMV (that for CS73 and VI it is open. The hypolophulid is simple and he identified as D. aff. hispanicus, see above). Here we confirm merges with the entolophid just anterior to the hypoconid. the ascription of the small-sized specimens to D. gracilis further The posterolophid is thin and long, it reaches the base of including those previously identified as D. aff. hispanicus and the entoconid and closes the posterosinusid. ascribe to this species material from the chronologically close sites (CS73, VI, CCW). m2 (Fig. 4I) Democricetodon gracilis is known from central Europe (Ger- The anterior valleys of the teeth are very narrow and closed many, Switzerland, Austria, Czech Republic; Maridet 2003; by the arms of the anterolophid (Appendix 2O). The sinusid, Wessels & Reumer 2009), France (Maridet 2003) and Turkey which points slightly forward, and the mesosinusid are closed (Theocharopoulos 2000), but it is very rare in Spain. Indeed, by low cingulids. In the CCW specimen (IPS85738) the this species had only been previously reported from two localities mesolophid is of medium length and merges with the entoco- from the Bardenas Reales de Navarra (Ebro Basin): Loma Negra nid. In the other specimen this ridge is also medium length, 64 and Pico del Fraile 2 (Suárez-Hernando 2017). These two but it does not merge with the entoconid (Appendix 2Q). sites belong to the middle Aragonian (zone D) while magneto- All the remaining morphological features are analogous to stratigraphical data indicate a correlation to chron C5Br, yielding those described for the m1. an estimated age of 15.7-15.9 Ma (Suárez-Hernando 2017). The Vallès-Penedès sites that have delivered D. gracilis are m3 (Fig. 4J) slightly older, belonging to the latest early Aragonian (sub- The mesolophid is absent. The sinusid is transverse and closed zone Cb; see below). Democricetodon gracilis is distinguished by a low cingulid. The posterosinus is closed by a very high from the similarly-aged Democricetodon franconicus by its posterolophid that merges with the reduced entoconid. The smaller size and more reduced m1, amongst other morpho- mesosinusid is closed by a low cingulid (e.g. IPS105145 from logical features (Maridet 2003). In the Calatayud-Montalbán SM, Fig. 4J). All other morphological features are analogous Basin D. franconicus is first recorded in sites from subzone Cb to those described for the m1 and m2. (late early Aragonian, MN4), which are about the same age as CS73, VI and SM (Van der Meulen et al. 2003, 2012). Comparisons and remarks In that area, D. franconicus would give rise to an anagenetic A few specimens of small size evidence the presence of a second lineage including (in chronological order) D. koenigswaldi, Democricetodon species in SM and VI, where it coexists with D. larteti (Schaub, 1925) and D. crusafonti (Agustí, 1978) D. cf. decipiens (see below and Fig. 3). Similarly, a single m1 (Van der Meulen et al. 2003). Democricetodon larteti and from CS73 stands out for its small size, clearly smaller than D. crusafonti are common components of the late Aragonian D. hispanicus and much smaller than Democricetodon sp. from (MN6, MN7+8) Vallès-Penedès faunas being index fossils for the same site (see Fig. 3). Finally, the Democricetodon speci- local biostratigraphy (Casanovas-Vilar et al. 2016a, b). While mens recovered at CCW, geographically and chronologically D. koenigswaldi might be present in some Vallès-Penedès sites very close to SM, are of small-size, particularly the m2 and the (see section Biostratigraphy and correlations), D. franconicus, m3. The recovered upper molars are further characterized by the oldest member of this lineage, appears to be absent. long mesolophs, the predominantly double protolophule on M1 and M2, and posterior metalophule on M1. The mesolophid is long in lower molars except m3, in which it is absent. The Democricetodon cf. decipiens anteroconid region of the m1 is very short, particularly in the (Freudenthal & Daams, 1988) specimen from SM (IPS103754). The described specimens are (Figs 3, 4; Table 1; Appendix 2) smaller than all other Democricetodon species present at West- ern Europe at the time but D. gracilis Fahlbusch, 1964 (see A catalog of the studied material and measurements is given Fahlbusch 1964; Maridet 2003; Wessels & Reumer 2009). in Appendix 1. See Table 1 for summary statistics and com- Furthermore, their morphology fits within the variation range of parisons. See also Figure 3 for measurements and compari- this species, which shows long mesolophs on M1 and medium sons. Morphotype frequency tables are given in Appendix 2. 410 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Table 1. — Summary measurements (in mm) of Democricetodon hispanicus Freudenthal, 1967, D. gracilis Fahlbusch, 1964, D. cf. decipiens (Freudenthal & Daams, 1988) and Democricetodon sp. 4 from different early Miocene sites of the Vallès-Penedès Basin. Only complete specimens were measured. Locality acronyms and measurement abbreviations are explained in section Abbreviations of the main text. For details on the measurement methods see section Material and methods, and Daams & Freudenthal (1988: 42, Fig. 1). L W Locality Species min. mean max. s.d. N min. mean max. s.d. N M1 SM D. cf. decipiens – 1.87 – – 1 – 1.20 – – 1 VI D. cf. decipiens – 1.91 – – 1 – 1.25 – – 1 SM D. gracilis 1.54 – 1.58 – 2 0.99 – 1.03 – 2 CS74 D. hispanicus 1.58 1.69 1.77 0.07 8 1.05 1.12 1.19 0.05 10 CMV3 D. hispanicus 1.47 1.52 1.63 – 3 1.04 1.05 1.08 – 3 LCV1 D. hispanicus 1.61 1.64 1.66 – 3 0.93 1.03 1.09 – 3 EC D. hispanicus – 1.63 – – 1 – 1.17 – – 1 M2 MOR1 D. sp. 4 – 1.6 – – 1 – 1.2 – – 1 SM D. cf. decipiens – 1.34 – – 1 – 1.17 – – 1 PA D. cf. decipiens – 1.39 – – 1 – – – – - VI D. cf. decipiens 1.29 – 1.45 – 2 1.18 – 1.23 – 2 SM D. gracilis 1.18 – 1.2 – 2 1.01 - 1.07 – 2 CS74 D. hispanicus 0.93 1.25 1.49 0.16 13 0.84 1.07 1.16 0.09 14 CMV3 D. hispanicus 1.06 1.22 1.35 0.10 8 0.97 1.09 1.16 0.07 8 LCV1 D. hispanicus 1.06 1.25 1.33 0.09 8 0.93 1.06 1.14 0.06 8 EC D. hispanicus 1.29 – 1.30 – 2 1.06 – 1.13 – 2 M3 VI D. cf. decipiens – 0.93 – – 1 – 1.01 – – 1 CS74 D. hispanicus 0.73 0.84 0.91 – 3 0.77 0.88 0.96 – 3 CS73 D. hispanicus – 0.93 – – 1 – 0.98 – – 1 CS72 D. hispanicus 0.7 – 0.94 – 2 0.75 – 0.97 – 2 CMV3 D. hispanicus 0.91 – 0.96 – 2 0.95 – 0.96 – 2 LCV1 D. hispanicus 0.81 0.88 0.93 – 3 0.88 0.91 0.95 – 3 m1 CS73 D. sp. 4 – 1.67 – – 1 – 1.08 – – 1 VI D. cf. decipiens 1.48 – 1.56 – – 0.80 0.98 1.09 – 3 VI D. gracilis 1.21 – 1.35 – 2 0.86 – 0.89 – 2 SM D. gracilis – 1.28 – – 1 – 0.91 – – 1 CS73 D. gracilis – 1.28 – – 1 – 0.92 – – 1 CCW D. gracilis – 1.37 – – 1 – 0.90 – – 1 CS74 D. hispanicus 1.4 1.44 1.49 0.03 12 0.93 0.97 1.01 0.02 12 CS73 D. hispanicus 1.42 1.48 1.57 0.05 8 0.94 0.98 1.11 0.05 8 CSU D. hispanicus – 1.31 – – 1 – 0.86 – – 1 CMV3 D. hispanicus 1.32 1.39 1.48 – 5 0.96 1.01 1.06 – 5 LCV1 D. hispanicus 1.38 1.45 1.50 - 3 0.96 0.98 1.00 - 3 m2 MOR1 D. sp. 4 – 1.51 – – 1 – 1.31 – – 1 PA D. cf. decipiens – 1.31 – – 1 – 1.21 – – 1 VI D. cf. decipiens 1.26 – 1.33 – 2 1.01 1.12 1.17 – 3 SM D. gracilis – 1.13 – – 1 – 0.96 – – 1 CCW D. gracilis – 1.17 – – 1 – 0.99 – – 1 CS74 D. hispanicus 1.19 1.26 1.39 0.06 15 0.93 1.07 1.20 0.07 15 CSU D. hispanicus 1.15 1.22 1.29 – 3 1.03 1.05 1.07 – 3 CMV3 D. hispanicus – 1.12 – – 1 – 0.91 – – 1 LCV1 D. hispanicus 1.06 1.21 1.31 0.1 10 0.88 0.97 1.05 0.07 10 m3 PA D. cf. decipiens – 1.21 – – 1 – 1.04 – – 1 VI D. gracilis – 1.11 – – 1 – 0.81 – – 1 SM D. gracilis 0.97 – 1.02 – 2 – 0.85 – – 2 CCW D. gracilis – 0.99 – – 1 – 0.77 – – 1 CS74 D. hispanicus 1.09 1.17 1.23 0.05 7 0.88 0.92 0.95 0.02 6 CS73 D. hispanicus 1.09 1.11 1.14 0.02 5 0.9 0.94 1 0.03 5 CMV3 D. hispanicus – 1.24 – – 1 – 0.89 – – 1 CMV2 D. hispanicus – 1.12 – – 1 – 0.94 – – 1 LCV1 D. hispanicus 1.11 1.14 1.19 – 3 0.899 0.92 0.94 – 3 COMPTES RENDUS PALEVOL • 2021 • 20 (22) 411 Jovells-Vaqué S. & Casanovas-Vilar I. Description mesolophid is either short (IPS86387 from PA, Fig. 4O) M1 (Fig. 4K) or absent (IPS87116 from VI). The metalophulid is simple The anterocone is simple in all studied specimens. The and anterior as well as the hypolophulid. The remaining anterolophule is simple IPS19548 from VI (Fig. 4K) and morphological features resemble those described for the forked in IPS19368 from SM, with both arms reaching the m1 (see Appendix 2K-N). anterocone separately (Appendix 2B). The protolophule is simple and merges with the entoloph posterior to the pro- m3 tocone. The ectoloph on the paracone is present in the SM Just IPS86388 from PA has been recovered. The anterosi- specimen and absent in the VI one. The mesoloph is short, nusid is absent and the anterolophid presents only a labial almost vestigial, and the metalophule is simple and poste- arm, which is well developed and closes the protosinusid. rior to the hypocone. All valleys are closed by low cingula. The mesolophid is completely absent and the mesosinusid is closed by a ridge. The entoconid is barely distinguishable. M2 (Fig. 4L) The sinusid is transverse and closed by a low cingulid that The anterior valleys are narrow and closed by the well- originates from the protoconid and reaches the hypoconid. developed arms of the anteroloph. The protolophule is double (Appendix 2G). Nevertheless, in the VI specimens Comparisons and remarks (Fig. 4L) the posterior arm is incomplete, being interrupted In the sites VI, PA and SM a Democricetodon species larger before reaching the paracone. In the same specimens the than D. hispanicus is present. However, the material in all these anterior arm is transverse and constricted at the point it sites is very scarce, so species attribution is tentative. These merges with the paracone. The ectoloph on the paracone specimens are distinguished not only by their larger dimen- is absent. The mesoloph ranges from short to long, but it sions but also by the more reduced mesoloph/ids as compared is more commonly of medium length. This ridge may be to D. hispanicus (Appendix 2). In the studied specimens the arched posteriorly as in IPS87106 from VI (Fig. 4L). The mesoloph/ids are predominantly of medium length or short, metalophule is posterior to the hypocone in all specimens whereas in D. hispanicus these ridges are more frequently long but in IPS86386 from PA, which shows a transverse met- (Appendix 2N, Q). Van der Meulen et al. (2003) reviewed alophule. The sinus is closed by a low cingulum. the genus Democricetodon in the Calatayud-Montalbán Basin and recognized two different anagenetic lineages (but see M3 (Fig. 4M) Freudenthal 2006 for a completely opposed view). Democri- Only two specimens have been recovered: IPS86941 and cetodon hispanicus and D. decipiens are recognized as succes- IPS87108 from VI. Both molars show well-developed sive members of the same lineage. Later species in the same anterolophs closing the anterior valleys. The protolophule lineage are (in chronological order): Democricetodon moralesi is double with a better developed posterior arm. The meso- (Freudenthal & Daams, 1988) , Democricetodon jordensi loph is either absent or fused with the posterior arm of the Van der Meulen et al., 2003 and Democricetodon lacombai protolophule. The sinus is vestigial and closed by a low cin- (Freudenthal & Daams, 1988). This lineage shows a set of gulum. Similarly, the posterosinus is also reduced to a tiny anagenetic trends including: size increase, reduction of meso- fossete and closed by the thick posteroloph. The metacone loph/ids, loss of the anterior protolophule on upper molars, is undistinguishable and the mesosinus is closed by a high and presence of a forked anterolophule on the M1 (Van der and thick ridge. Meulen et al. 2003). Many of these morphological features are observed in the studied specimens, particularly the M1. m1 (Fig. 4N) As far as size is concerned, the specimens fit within the size Only two complete specimens have been recovered: IPS87109 range of D. decipiens, being larger than D. hispanicus and at and IPS87110, both from VI. The anteroconid is simple the same time smaller than D. moralesi or the similarly-aged and bean shaped. The labial anterolophid is well developed, Democricetodon koenigswaldi, which belongs to the second thus closing the protosinusid (Appendix 2K), while the lineage (Van Der Meulen et al. 2003) (Fig. 3). However, anterosinusid is open because the lingual anterolophid does the material is too scarce for a confident attribution to this not reach the base of the metaconid. The mesolophid is of species, so it is assigned to D. cf. decipiens until further medium length in all studied specimens. The metalophulid specimens are recovered. and the hypolophulid are anterior to the protoconid and hypoconid, respectively. A cingulid descending from the metaconid partially closes the mesosinusid, but this valley Democricetodon sp. 4 remains open. The sinusid is proverse and closed by a low (Figs 3, 4; Table 1; Appendix 2) cingulid. A catalog of the studied material and measurements is m2 (Fig. 4O) given in Appendix 1. See Table 1 for summary statistics The molars are squared, being relatively wide. The lingual and comparisons. See also Figure 3 for measurements and anterolophid is highly reduced in most of the studied comparisons. Morphotype frequency tables are given in specimens, thus resulting in very narrow anterosinusid. The Appendix 2. 412 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Description (Southern Germany; Wessels & Reumer 2009). Interestingly, M2 (Fig. 4P) the CS73 m1 had been previously ascribed to the genus Fahl- Only a single M2 has been recovered, IPS88868 from buschia Mein & Freudenthal, 1971 (Agustí & Llenas (1993) MOR1 which is highly worn and slightly damaged on its which was later synomymized with Democricetodon (Van Der labial margin. Both arms of the anteroloph are high and long Meulen et al. 2003; but see Freudenthal 2006). Jovells-Vaqué and close the anterior valleys. The protolophule is double et al. (2017:146) ascribed this specimen to a second, larger- with both arms similarly well developed. The paracone is sized Democricetodon species that would have coexisted with damaged, but apparently it did present a short ectoloph. D. hispanicus. These authors already noted some affinities with The mesoloph is of medium length and the mesosinus is D. moralesi and D. koenigswaldi. Concerning the specimens from wide and closed by a cingulum. The sinus is transverse and MOR1, the single M2 and m2 are the only identifiable rodent closed by a merging the protocone with the hypocone. The material found at the site. Clearly the material is too scarce to metalophule is simple and transverse. The posteroloph is confidently ascribe it to the species level. Furthermore, it may long and reaches the metacone, thus resulting in a closed well be possible that the specimen from CS73 and those from posterosinus. MOR1 belong to different Democricetodon species of similar size. In the Calatayud-Montalbán Basin D. moralesi, ranges from m1 (Fig. 4Q) local subzone Cb (late Early Aragonian, MN4) to Db (middle Only one m1 has been recovered: IPS45052 from CS73. The Aragonian, MN5; Van Der Meulen et al. 2003, 2012). On the anteroconid is asymmetric. The anterior valleys of the teeth are contrary, D. koenigswaldi is not recorded until much later, its narrow, especially the anterosinusid, and closed by the arms first occurrence defining the lower boundary of subzone Dd (late of the anterolophid. The metalophulid is short and merges middle Aragonian, MN5; Van Der Meulen et al. 2003, 2012). with the anterolophulid just anterior to the protoconid. The Since CS73 is correlated to zone C (Jovells-Vaqué et al. 2017; mesolophid is long, almost reaching the lingual margin of see section Biostratigraphy and correlations) D. moralesi would the molar and pointing anteriorly. The mesosinusid is closed be a more likely candidate than D. koenigswaldi. MOR1 is by a low cingulid which presents a thickening resembling younger, coinciding with the Langhian transgression at the a mesostylid. The sinusid points markedly forward and is Vallès-Penedès Basin, but probably not as late as subzone Dd, closed by a low cingulid merging the protoconid with the so an attribution to D. moralesi is also more probable in this hypoconid. The hypolophulid is simple and merges with the case. It is worth remarking that the morphology and size of entolophid anterior to the hypoconid. Finally, the poster- the few recovered specimens also fits with D. mutilus, a species olophid is thick and reaches the base of the entoconid, thus known from Serbia, Hungary, Austria, Germany, Switzerland closing the posterosinusid. and France, but not recorded in the Iberian Peninsula. This species co-occurs with D. gracilis in certain Swiss (Kälin & m2 (Fig. 4R) Kempf 2009) and German sites (Wessels & Reumer 2009), IPS105188 from MOR1 is the only m2 recovered. The molar is which is also present at CS73. While D. mutilus is also a likely relatively wide. It shows a highly reduced lingual anterolophid candidate for the ascription of the limited Vallès-Penedès and a vestigial anterosinusid. On the contrary, the labial arm of sample, only the recovery of further material will allow clari- the anterolophid is well developed and closes the protosinusid. fying the identity of this larger-sized Democricetodon species The mesolophid is vestigial. The sinusid is narrow and transverse. in CS73 and MOR1. The remaining morphological features resemble those described for the m1. Genus Megacricetodon Fahlbusch, 1964 Comparisons and remarks These three specimens stand out among all other early Mio- Megacricetodon primitivus Freudenthal, 1963 cene Democricetodon specimens found in the Vallès-Penedès (Figs 4, 5; Table 2; Appendix 4) Basin because of their larger size (Fig. 3). They are larger than D. hispanicus and D. decipiens, the few recovered specimens A catalog of the studied material and measurements is given fitting within the range size of D. moralesi, D. koenigswaldi in Appendix 3. See Table 2 for summary statistics and com- and Democricetodon mutilus (Fig. 3). Some characters of the parisons. See also Figure 5 for measurements and compari- M2 of MOR1 fit better with D. moralesi (Van der Meulen sons. Morphotype frequency tables are given in Appendix 4. et al. 2003) or D. mutilus (Wessels & Reumer 2009) such as the symmetrically double protolophule and simple transverse Description metalophule. While these characters may also occur in D. M1 (Fig. 4S, T) koenigswaldi, double and posterior metalophules predominate The anterocone is deeply split with the labial cusp larger on the M2 instead of transverse ones. Furthermore, the antero- than the lingual one in most molars, but in the material conid of the m1 recovered at CS73 is characteristically asym- from younger sites (PA, VI) both cusps are more frequently metric as in D. moralesi. However, this specimen presents a long of similar size (Appendix 4A). Most of the specimens, such mesolophid, which is a rare feature in both D. moralesi and D. as IPS44939 from CS74 or IPS86432 from LCV (Fig. 4T), koenigswaldi but very common in D. mutilus from Sandelzhausen show a well-developed cingulum or anterior platform at the COMPTES RENDUS PALEVOL • 2021 • 20 (22) 413 Jovells-Vaqué S. & Casanovas-Vilar I. Table 2. — Summary measurements (in mm) of Megacricetodon primitivus (Freudenthal, 1963) from different early Miocene sites of the Vallès-Penedès Basin. Only complete specimens were measured. Locality acronyms and measurement abbreviations are explained in section Abbreviations of the main text. For details on the measurement methods see section Material & methods, and Daams & Freudenthal (1988: 42, Fig. 1). L W Locality min. mean max. s.d. N min. mean max. s.d. N M1 PA 1.41 1.50 1.64 0.07 12 0.94 0.99 1.07 0.03 11 VI 1.37 1.45 1.58 0.06 23 0.89 0.95 1.07 0.05 24 CSU – 1.51 – – 1 1.00 – 1.02 – 2 CS73 1.32 1.45 1.64 0.12 7 0.88 0.97 1.16 0.10 7 CS74 1.40 1.46 1.53 0.06 5 0.87 0.93 1.05 0.07 5 CMV3 1.31 1.44 1.55 0.07 10 0.77 0.91 0.97 0.04 18 LCV1 1.34 1.43 1.91 0.17 11 0.83 0.90 1.03 0.06 12 M2 PA 0.99 1.09 1.17 0.04 14 0.86 0.94 1.02 0.04 14 VI 1.01 1.08 1.22 0.05 40 0.81 0.90 0.97 0.04 41 CSU – – – – – – 0.98 – – 1 CS72 0.98 – 1.11 – 2 0.95 – 0.98 – 2 CS74 1.10 – 1.14 – 2 0.83 – 0.92 – 2 CMV3 0.97 1.05 1.12 0.04 18 0.80 0.88 0.94 0.03 18 CMV1 – 1.05 – – 1 – 0.96 – – 1 LCV1 0.96 1.05 1.11 0.03 15 0.77 0.87 0.96 0.04 16 M3 PA 0.71 0.77 0.86 – 4 0.75 0.79 0.86 – 4 VI 0.77 0.92 0.92 – 2 0.76 0.93 0.96 – 2 CS72 0.64 – 0.65 – 2 0.74 – 0.75 – 2 CS73 – 0.84 – – 1 – 0.72 – – 1 CS74 – 0.75 – – 1 – 0.71 – – 1 CMV3 0.64 0.73 0.82 0.04 13 0.67 0.74 0.83 0.05 13 CMV1 0.66 0.70 0.73 – 3 0.68 0.75 0.81 – 3 LCV1 0.68 0.76 0.78 0.10 3 0.66 0.70 0.79 0.10 3 m1 PA 1.48 1.38 1.48 0.07 7 0.78 0.84 0.94 0.05 7 VI 1.21 1.33 1.47 0.06 19 0.75 0.81 0.89 0.03 22 SM – [1.37] – – 1 – [0.92] – – 1 CSU – 1.34 – – 1 – 0.82 – – 1 CS72 1.31 1.36 1.39 – 3 0.85 1.36 1.39 – 3 CS73 1.10 1.27 1.40 0.11 5 0.64 0.77 0.84 0.09 6 CS74 1.08 1.35 1.45 0.09 17 0.76 0.85 0.93 0.04 18 CMV3 1.18 1.25 1.33 0.05 18 0.69 0.81 0.90 0.04 18 CMV1 1.14 1.18 1.27 – 3 0.73 – 0.83 – 2 LCV1 1.13 1.23 1.35 0.05 16 0.71 0.79 0.88 0.04 17 m2 PA 0.95 1.12 1.20 0.06 18 0.70 0.91 0.97 0.05 19 VI 0.99 1.09 1.15 0.04 30 0.78 0.90 0.99 0.04 32 CSU – 1.10 – – 1 – 0.91 – – 1 CS72 1.01 1.06 1.13 – 3 0.81 0.84 0.87 – 3 CS74 1.03 1.09 1.09 0.05 12 0.82 0.89 0.98 0.05 12 CMV3 0.98 1.05 1.10 0.03 17 0.80 0.85 0.91 0.02 17 CMV2 – 1.10 – – 1 – 0.89 – – 1 LCV1 0.85 1.02 0.85 0.06 15 0.72 0.84 0.95 0.04 15 m3 PA 0.87 0.93 1.00 – 4 0.70 0.76 0.84 – 4 VI 0.87 0.92 1.00 0.04 13 0.66 0.73 0.92 0.07 13 CS73 – 1.00 – – 1 – 0.81 – – 1 CS74 0.84 0.92 1.08 0.09 5 0.72 0.74 0.77 0.02 5 CMV3 0.70 0.84 0.92 0.05 17 0.57 0.68 0.74 0.04 18 LCV1 0.79 0.83 0.89 – 4 0.68 0.69 0.71 – 4 base of the anterocone. The presence of a labial spur on the with the anterolophule posteriorly to the protocone. The anterolophule is variable, but it is more frequently absent. mesoloph is always present, most frequently being long, When present, it is very short, almost vestigial, see IPS86997 short in just a few specimens. The ectoloph on the paracone from VI (Fig. 4S) and IPS86432 from LCV1 (Fig. 4T; is also variable. In the specimens from LV1 and PA it is Appendix 4C). The protolophule is always simple and merges more frequently absent, whereas in the other sites a short 414 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe m1 M1 1.2 1.2 M. primitivus 1.0 1.0 W W M. collongensis 0.6 0.8 M. collongensis M. primitivus 0.6 M. minor M. minor 0.6 1.2 1.4 1.6 1.8 1.0 1.2 1.4 1.6 L L m2 M2 1.2 1.1 M. collongensis M. collongensis 1.0 1.0 M. primitivus W 0.9 W 0.8 0.8 M. primitivus M. minor 0.7 M. minor 0.6 0.8 0.9 1.0 1.1 1.2 1.3 0.8 1.0 1.2 1.4 L L m2 M3 1.0 Locality 1.0 M. collongensis CCW CMV1 0.9 CMV2 0.9 CMV3 CS72 0.8 0.8 CS73 CS74 W W CSU 0.7 0.7 M. collongensis EC LCV1 0.6 M. primitivus MOR1 0.6 PA M. primitivus M. minor SM M. minor 0.5 0.5 VI 0.5 0.6 0.7 0.8 0.9 1.0 0.6 0.7 0.8 0.9 1.1 1.1 L L Fig. 5. — Length/width scatter plot for Megacricetodon primitivus (Freudenthal, 1963) from different Vallès-Penedès sites. Locality acronyms and measurement abbreviations are explained in section Abbreviations of the main text. For details on the measurement methods see section Material and methods, and Daams & Freudenthal (1988: 42, fig. 1). The ellipses show the 95% confience interval for Megacricetodon primitivus (Oliver & Peláez-Campomanes 2014), M. minor (Lartet, 1851) (Wessels & Reumer 2009) and M. collongensis (Mein, 1958). COMPTES RENDUS PALEVOL • 2021 • 20 (22) 415 Jovells-Vaqué S. & Casanovas-Vilar I. ectoloph is frequently present but it never merges with the short or of medium length, and only rarely long or absent. mesoloph. In all the specimens from the VI the ectoloph is The mesosinusid is open (e.g. IPS86440 from LCV1, Fig. 4Y). relatively longer, almost touching the long mesoloph, such as The metalophulid and the hypolophulid are both simple, in IPS86997 (Fig. 4S). The metalophule consists of a single anterior and quite short in all specimens. The sinusid is gen- arm posterior to the hypocone that may be placed quite erally wide and transverse, although in some specimens this lingually, particularly in younger samples such as VI and PA, valley points forward (e.g. IPS44950 from CS74, Fig. 4X). thus resulting in a highly reduced posterosinus. The sinus is The posterolophid is well developed and reaches the entoconid transverse in all studied specimens. thus closing the posterosinusid. M2 (Fig. 4U, V) m2 (Fig. 4X, Z) The anterior valleys of the teeth are closed by the well-devel- The anterior valleys of the teeth are closed by the arms of the oped arms of the anterolophule. The labial arm is notably anterolophid. The lingual arm of the anterolophid is highly higher than the lingual one. The protosinus is reduced. The reduced (e.g IPS86444 from LCV) and sometimes absent (e.g. protolophule is simple and slightly anterior to the protocone IPS44950 from CS74, Fig. 4X). As in the case of the m1, the in most cases (Appendix 4L), except in a few specimens mesolophid is variable, but it is more frequently short or of from VI (e.g. IPS87022, Fig. 4V) and PA (e.g. IPS86335) medium length, only very rarely long or absent (e.g. IPS86444 that present a double protolophule with a stronger anterior from LCV1, Fig. 4Z). The remaining morphological features arm. In a few other specimens, such as IPS86435 from LCV resemble those of the m1. (Fig. 4U), the protolophule shows a vestigial posterior arm. The ectoloph is variable and in some specimens such as IPS86328 m3 (Fig. 4A’) from PA, IPS87022 from VI (Fig. 4V) and IPS 86888 from All the valleys of the m3 are reduced, including the sinusid VI the ectoloph is well developed and weakly connects the and the mesosinusid, as compared to those of the m1 and m2. paracone and the mesoloph. The mesoloph is present in all The protosinusid is particularly narrow and it is closed by the specimens but IPS86444 from LV1. This ridge is generally of labial anterolophid, which is quite low. The anterosinusid and medium length or long. The metalophule is simple in all the lingual anterolophid are vestigial or have completely disap- studied specimens and generally transverse or slightly ante- peared in some molars (e.g. IPS86451 from LCV1, Fig. 4A’). rior (Appendix 4O). The sinus normally is transverse in the The sinusid and mesosinusid are closed by low cingulids. younger localities such as VI and PA but in most specimens The mesolophid is absent in all the studied specimens. The from LCV and CMV3 it points forwards (Appendix 4N). entoconid is highly reduced, being barely recognizable in most teeth. The posterolophid is well developed and closes M3 (Fig. 4W) the posterosinusid. The teeth are extremely reduced and button shaped. The labial anteroloph is well developed and delimits the narrow anter- Comparisons and remarks osinus (IPS19479 from CS74, Fig. 4W). The lingual anter- The described specimens are similar in size to those of small oloph and protosinus are generally vestigial as in IPS86439 Megacricetodon species (Fig. 5) such as Megacricetodon primitivus from LCV1. The protolophule is simple and merges with (Freudenthal, 1963), Megacricetodon collongensis (Mein, a very short anterolophule anterior to the protocone. The 1958) and Megacricetodon minor (Lartet, 1851), which are mesoloph is highly variable, in some cases such as IPS19479 abundant in the Iberian Peninsula during the Aragonian from CS74 (Fig. 4W) the mesoloph is long and has a small (Daams & Freudenthal 1988; Oliver Pérez 2015; Oliver & mesostyle at the end. The metacone is recuded, being barely Peláez-Campomanes 2016). However, several morphological distinguishable in many specimens such in IPS45094 from features allow for the ascription of the material to M. primitivus. CS72. The metalophule is anterior to the hypocone. The The size of the specimens overlaps with that of M. primitivus posteroloph is well developed and closes the posterior valley and M. minor, but in the studied material the anterocone of the tooth. The transverse and narrow sinus is closed by a in the M1 is deeply split as in M. primitivus and not simple low cingulum. Some specimens present a reduced sinus and to slightly split as in M. minor. Megacricetodon collongensis the protocone and hypocone show a weak direct connection is of similar size, although slightly larger than M.primitivus (IPS86439 from LCV1). (Oliver & Peláez-Campomanes 2014; Fig. 5). This species further differs from M. primitivus by the much more frequent m1 (Fig. 4X, Y) occurrence of subdivided anteroconids on the m1 and the The anteroconid is simple and round in all the studied speci- higher proportion of double protolophules and metalophules mens but in a few teeth from PA and CS74 which show a sligh on M1 and M2 (Oliver & Peláez-Campomanes 2014). Except subdivision of this cusp (Appendix 4P). In these cases, the for a few specimens from PA and CS74, in the studied material two cusps of the anteroconid are of similar size. A few speci- the anterocone is predominately simple and rounded, while mens from CS74 (6 out of 17), such as IPS44950 (Fig. 4X), and the protolophule and metalophule are predominantly show a vestigial and low labial spur on the anterolophulid simple in the M1. However, in a few M2 from LV1, PA and (Appendix 4Q). Such spur is absent in all the remaining VI, the protolophule shows an incomplete posterior arm and material. The mesolophid is variable, being more frequently in a few others from PA and VI, the protolophule is double. 416 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Calatayud-Montalbán Basin Vallès-Penedès Basin Northern Alpine Foreland Basin Democricetodon franconicus Magnetic Local Local Magnetic Democicetodon gracilis polarity MN zones Eumyarion weinfurteri Ma zones MN polarity Ma Eumyarion weinfurteri D. mutilus C5ACn E Osch- C5ACn 14 14 Democricetodon decipiens Megacricetodon primitivus graben C5ACr C5ACr MN6 Eumyarion weinfurteri Democricetodon moralesi Rümikon Megacricetodon primitivus C5ADn C5ADn C5ADr Uzwil- C5ADr Netzen- MN5 Democricetodon gracilis D Democricetodon sp.4 buech Melissiodon dominans Apistobel 15 C5Bn 15 Megacricetodon bavaricus Democricetodon decipens C5Bn Ligerimys ellipticus 520m Ligerimys ellipticus Tobel Hombre chtikon Democricetodon hispanicus Democricetodon hispanicus PA C5Br Democicetodon franconicus C5Br Ligerimys florancei Ligerimys florancei Melissiodon dominans MN5 4 Vermes 1 4 MOR Ligerimys florancei SM, CCW Melissiodon dominans 16 16 Ligerimys antiquus VI C Ligerimys antiquus CS, CSU, CJ, C5Cn 3 Oberkulm- MN4 C5Cn Pseudotheridomys sp. CMV, LCV1 EC Sälem Pseudotheridomys fejfari 1 3 1 4 B 2 3 Tägernau- 17 2 strasse 17 C5Cr MN4 C5Cr C5Dn MC 2 Glovelier C5Dn 1 C5Dr TFR1, CB C5Dr SAB 18 18 Trub-Sältenbach MN3 A MN3 C5En C5En C5Er C5Er 19 19 C6n C6n Fig. 6. — Biostratigraphic distribution of the early Miocene cricetid species in the Vallès-Penedès Basin. Selected eomyid taxa are also included because of their use in biostratigraphical correlation. Black dots represent the occurrence of a taxon in a given site while grey dots indicate uncertain attribution to that taxon. The Calatayud-Montalbán basin record and local biostratigraphy is after Van der Meulen et al. (2012) while those for the Swiss Molasse Basin is after Kälin & Kempf (2009). Four main biochronlogic events are distinguished: 1) First occurrence (FO) of the eomyid Ligerimys florancei; 2) FO of the cricetid Democricetodon; 3) FO of the cricetid Megacricetodon; 4) Last occurrence (LO) of the eomyid genus Ligerimys. Note the different order and timing of bioevents in the different regions. For locality acronyms see section Abbreviations of the main text. Black indicate normal magnetic polarity chrons, white indicate inversed magnetic polarity chrons. The metalophule is simple in all M2. The mesolophs/ids Megacricetodon primitivus is the first Megacricetodon spe- are generally long or of medium length in all molars but in cies to be recorded in the Iberian Peninsula, its oldest record the m3, which has no mesolophid. This contrasts with the being at the site of Artesilla in the Calatayud-Montalbán type material from Valtorres (Calatayud-Montalbán Basin, Basin (Oliver Pérez et al. 2008, 2015; Oliver & Peláez-Cam- Spain), in which this ridge is less developed, more frequently pomanes 2014). This site is correlated to local subzone Ca and being short to medium length in all molars but the m3, its age is estimated as 16.49 Ma (Van der Meulen et al. 2012). where it is generally absent (Freudenthal 1963; Daams & This species persists in the Calatayud-Montalbán Basin for Freudenthal 1988; Oliver & Peláez-Campomanes 2014). Other almost one million years, its last record being at Valdemoros differences with the type material include the larger size of 8A correlated to local subzone Db, with an estimated age of some of the specimens from the younger sites (VI, PA), which 15.68 Ma (Oliver & Peláez-Campomanes 2014). Its disappear- is however within the size range for the species (see Daams & ance coincides with the first record of additional, larger-sized Freudenthal 1988; Oliver & Peláez-Campomanes 2014). Megacricetodon species in the region, such as Megacricetodon Agustí (1981, 1982) recognized a second larger-sized species vandermeuleni (Oliver & Peláez-Campomanes 2013, 2016). of Megacricetodon in CMV represented by a single m1. This Megacricetodon primitivus does not show directional changes specimen measures 1.38 by 0.78 (length by width) and therefore in size or morphology throughout its range, despite showing fits perfectly within the range of M. primitivus (see Oliver & a high inter- and intra-populational variability (Oliver & Peláez-Campomanes 2014), not being even especially large. Peláez-Campomanes 2014). This contrasts with the situation Therefore, we ascribe it to the latter species, which would be in most Megacricetodon lineages and other coeval cricetids, the only Megacricetodon species present at the Vallès-Penedès such as Democricetodon, which do show clear anagenetic trends during the early Miocene. during the same time span. The Vallès-Penedès material also COMPTES RENDUS PALEVOL • 2021 • 20 (22) 417 Jovells-Vaqué S. & Casanovas-Vilar I. reflects this situation. Size of the molars remains stable, with Basin the Ramblian sites corresponding to the “cricetid only a minor increase in the length of m1 (see Table 2). As vacuum” are correlated to local zone A (c. 19.5-16.77 Ma; far as morphology is concerned, the only temporal trends are Daams & Freudenthal 1988; Daams et al. 1999; Van der the rare occurrence of split anteroconids on the m1 in the Meulen et al. 2012). In the Vallès-Penedès Basin Ramblian specimens from younger sites as well as an apparent trend sites include Sant Andreu de la Barca, Turó de les Forques, towards the reduction of the posterosinus on M1. In the La Costablanca, and Molí de Can Calopa (Casanovas-Vilar younger localities, such as VI and PA, the metalophule is et al. 2011, 2016b). These have delivered a glirid-dominated connected to the posteroloph more obliquely, thus delimiting fauna that also includes the cricetid Melissiodon (Jovells-Vaqué & a more reduced posterosinus. While this is also observed in Casanovas-Vilar 2018) but no modern cricetids. Besides the a significant number of specimens from older sites, such as absence of cricetids other than Melissiodon, these sites are also in those from CMV3, it is by far more common in younger characterized by a high abundance of glirids and eomyids, ones. Such pattern is not observed in the Calatayud-Montalbán represented by the genera Ligerimys Stehlin & Schaub, 1951 material, the posterosinus of the M1 being generally more and (more infrequently) Pseudotheridomys Schlosser, 1926. reduced, comparable to the specimens from VI and PA, even Local zone B of the Calatayud-Montalbán Basin (16.77- in the oldest sites (see Oliver & Peláez-Campomanes 2014: 16.49 Ma) marks the beginning of the Aragonian and is table 10). characterized by Democricetodon hispanicus as the only mod- ern cricetid present (Daams & Freudenthal 1988; Daams et al. 1999; Van der Meulen et al. 2012). This interval, cor- DISCUSSION responding to the earliest part of the MN4, would be entirely missing in the Vallès-Penedès. Casanovas-Vilar et al. (2016b: Biostratigraphy and correlations 802) mentioned scarce remains of Democricetodon from Turó Overall the early Miocene cricetid succession in the Vallès- de les Forques (indeed just a single m1). However, this was Penedès Basin is similar to that of other regions, such as the later found to be an unfortunate contamination from LCV, Calatayud-Montalbán Basin (Van der Meulen et al. 2012) or which was being sorted at the same time. The specimen was the Swiss Molasse Basin (Kälin & Kempf 2009) (Fig. 6). The distinguished because of its different color and aspect of the order of events in all these regions is as follows: 1) after the sediment still trapped in the main valleys, both matching LCV. last occurrence of Eucricetodon, there is a “cricetid vacuum” The presence and abundance of the eomyid Ligerimys ellipticus (Daams & Freudenthal 1989) with only the genus Melis- Daams, 1976 characterizes zone C (16.49-15.93 Ma) together siodon present; 2) the first modern cricetid to be recorded is with the first appearance of Megacricetodon (M. primitivus) and Democricetodon; 3) Megacricetodon, a second genus of modern Eumyarion (Daams & Freudenthal 1988; Daams et al. 1999; cricetid, appears slightly later, together with the paracriceto- Van der Meulen et al. 2012). The sites of the Subirats lacus- dontine Eumyarion Thaler, 1966; and 4) by the end of the trine unit (LCV, CMV and CS) are the oldest MN4 sites in early Miocene additional Democricetodon species are recorded the basin and they already record both Megacricetodon and in all the regions. Even though Democricetodon and Megacri- Eumyarion (Agustí 1981, 1983; Jovells-Vaqué et al. 2017, cetodon are represented by distinct species in Central Europe 2018). Ligerimys ellipticus is present in most of the studied sites and Iberia well since their first occurrence, and there is a (see Fig. 6). Its absence in EC, CSU and CCW can certainly clear diachrony in the timing of the events (Van der Meulen be attributed to the scarce sample recovered at these sites. et al. 2012), their order is the same in both regions (Fig. 6). However, these localities can be correlated to zone C because The first modern cricetid recorded in the Swiss Molasse is of the presence of M. primitivus in CSU and Eumyarion in D. franconicus, which first occurs at 17.6 Ma. On the other EC. Regarding CCW, it is lithostratigraphically correlated to hand, in Calatayud-Montalbán D. hispanicus is present in SM, which, in turn, unambiguously belongs to zone C. Van low numbers at 17 Ma (Van der Meulen et al. 2003, 2012). der Meulen et al. (2012) subdivided zone C into subzones The genus Megacricetodon, represented by M. collongensis, first Ca (16.49-16.30 Ma) and Cb (16.3-15.93 Ma) according occurs at 17.2 Ma in Switzerland, whereas in Spain it first to the Democricetodon species present and the replacement appears at 16.6 Ma and is represented by M. primitivus. Finally, of Ligerimys florancei Stehlin & Schaub, 1951 by Ligerimys Eumyarion, an archaic “cricetid” genus related to Oligocene ellipticus. The replacement occurred within subzone Ca, but forms such as Eucricetodon (Freudenthal et al. 1992), first in a couple of Vallès-Penedès sites, LCV and CMV, both appears simultaneously with Megacricetodon in Switzerland Ligerimys species co-occur, even though L. ellipticus is by far (Kälin & Kempf 2009) and perhaps slightly later in Spain more abundant. In all the remaining sites only L. ellipticus is (Van der Meulen et al. 2012). present and it is rare. While this would apparently support Previous works have remarked that the early Miocene rodent a distinction of subzones Ca and Cb in the Vallès-Penedès succession of the Vallès-Penedès shows several coincidences with Basin as well, it is contradicted by the different cricetid suc- that of Calatayud-Montalbán, thus indicating that the same cession. In LCV and CMV, D. hispanicus is present, while in detailed local zonation could be used in both areas (Casanovas- the Calatayud-Montalbán Basin, its descendant D. decipiens Vilar et al. 2011, 2016b; Jovells-Vaqué et al. 2017, 2018). is the only Democricetodon that occurs in subzone Ca (Van Our results support these earlier conclusions albeit they also der Meulen et al. 2003, 2012). In CS, which would be cor- show some significant differences. In the Calatayud-Montalbán related to subzone Cb, two additional Democricetodon species 418 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe are recorded (D. gracilis and the larger-sized D. sp. 4), but the Vallès-Penedès Basin. Ongoing magnetostratigraphical D. hispanicus is most abundant. Democricetodon cf. decipiens is studies in the early and middle Miocene successions of this recorded in VI, SM and PA, coexisting with D. gracilis in VI. basin will surely refine these correlations. In MOR1 D. cf. moralesi is represented by scarce remains. In Calatayud-Montalbán, the replacement of D. decipiens by its Paleobiogeographic descendant D. moralesi defines the lower boundary of subzone and paleoenvironmental implications Cb (Van der Meulen et al. 2012). A second Democricetodon Several studies have stressed the differences in faunal com- lineage first occurs in that basin during subzone Cb (D. fran- position between the Vallès-Penedès Basin and other, more conicus, Van der Meulen et al. 2003, 2012), which is not pre- inland, areas of the Iberian Peninsula during the Miocene sent in the Vallès-Penedès. On the contrary, D. gracilis is not (Agustí 1990; Casanovas-Vilar et al. 2005, 2008; Casanovas- recorded in Calatayud-Montalbán, but is first recorded in the Vilar & Agusti 2007; Maridet et al. 2007, 2013). As far as Swiss Molasse during the late MN4 (Kälin & Kempf 2009), rodents are concerned, differences with the central Iberian more or less at the same time it is first recorded in the Vallès- basins are particularly clear during the latest Aragonian (mid- Penedès Basin. The Ebro Basin sites that have delivered this dle Miocene, MN7 + 8) and the Vallesian (late Miocene, species are slightly younger, being correlated to the earliest MN9-MN10), with more diverse faunas including a greater MN5 (Suárez-Hernando 2017). Clearly, the Vallès-Penedès number of purported forest-dwelling taxa in the Vallès- sites can be correlated to zone C of the Calatayud-Montalbán Penedès (Casanovas-Vilar & Agustí 2007; Casanovas-Vilar Basin, but the distinction of subzones in the Catalan basin is et al. 2008). This pattern is also recognized for the larger questionable. Yet, it seems clear that LCV and CMV, which mammals, for example hominoid primates, chalicotheres still record the eomyid L. florancei together with L. ellipticus, and tapirs are present at the Vallès-Penedès and other Catalan are older than the remaining sites. At CS, VI and SM addi- Basins (Cerdanya and Seu d’Urgell basins, in the Pyrennees) tional Democricetodon species are recorded, which would argue but unknown from elsewhere in Spain at that time. Catalo- for a younger age of these sites, even if the Democricetodon nia is consistently recognized as a transitional area between species differ. the humid and forested environments of west and central For PA and MOR1, two of the younger sites (see Fig. 2), Europe and the drier landscapes of inner Iberia (Casanovas- the correlation is not clear. The rodent fauna of PA includes Vilar et al. 2005, 2008). three cricetids (M. primitivus, D. cf. decipiens and Eumyarion This situation probably already existed during the early cf. weinfunteri (Schaub & Zapfe, 1953)) but no eomyids. Miocene, although available data are still scarce. Nevertheless, Megacricetodon primitivus is by far the most abundant cricetid Costeur & Legendre (2008a, b) showed that the early Miocene in the sample, D. cf. decipiens being represented by just four (MN3-MN4) large mammal faunas from the Vallès-Penedès molars (one broken) and Eumyarion by one. Total sample are distinct from those of other Iberian basins, and include a size includes more than 100 identifiable rodent specimens so slightly higher diversity of herbivores, particularly suiforms. that the absence of Ligerimys is intriguing. If confirmed, the In this regard, they are closer to contemporary faunas from presence of D. decipiens would suggest a correlation to zone southern France and Germany. The insectivores (Eulipotyphla) C. The PA site is located just tens of meters below Langhian show a clear latitudinal diversity gradient throughout the marine limestones (Casanovas-Vilar et al. 2011, 2016b), so Miocene in western Europe, with decreasing generic diversity it may be middle Miocene rather early Miocene. The absence at lower latitudes, mostly because of the absence of many of Ligerimys, which disappears at the beginning of the mid- genera and even entire families in the inner basins of the dle Aragonian (MN5), zone D (Van der Meulen et al. 2012), Iberian Peninsula at particular time intervals (Furió et al. 2011; would support this tentative correlation. Regarding MOR1, Madern & Van den Hoek Ostende 2015). During the early this locality is located in transitional facies associated to the Miocene central Europe presented a remarkable diversity of Langhian marine transgression and it has only delivered two talpids and soricids, but only a few species are recorded in inner molars which are attributed to an undetermined, large-sized Spain. Indeed, the early Miocene (MN3-MN4) insectivore Democricetodon species. Considering its stratigraphical posi- faunas of the Calatayud-Montalbán Basin generally include tion we correlate it to the beginning of the middle Miocene. the erinaceid Galerix Pomel, 1848 and the talpid Desmanodon Recently, a series of karstic fissure fillings have been located Engesser, 1980 besides a few more genera (such as Heterosorex within Langhian limestones in Clariana (Castellet i La Gornal, Gaillard, 1915 and Miosorex Kretzoi, 1959) (Van der Meulen Penedès sector). Several fissures have delivered a rich sample et al. 2012; Furió et al. 2017). The early Miocene insectivore comprising thousands of small mammals including insec- faunas of the Vallès-Penedès have been recently described tivorans, chiropterans, lagomorphs and rodents. The rodent (Van den Hoek Ostende et al. 2020). Overall, the insectivores fauna has yet to be prepared and studied, but it includes confirm the intermediate position of the Vallès-Penedès, but Cricetodon and a medium-sized Megacricetodon species, prob- show that differences with the inland were less pronounced than ably M. collongensis. Cricetodon first occurs during the late during the middle and late Miocene (Van den Hoek Ostende MN5 (zone E) in Calatayud-Montalbán, where it coexists et al. 2020). The presence of dimylids (Plesiodimylus Gaillard, with M. collongensis. Clariana sites may well be of similar age 1897, Chainodus Ziegler, 1990), particularly in Ramblian and, together with PA and MOR1, fill the gap between the sites would suggest somewhat more humid conditions in early (MN4) and late Aragonian (MN6-MN7 + 8) sites in Catalonia. Dimylids were specialized insectivores, presumably COMPTES RENDUS PALEVOL • 2021 • 20 (22) 419 Jovells-Vaqué S. & Casanovas-Vilar I. malacophagous, which would have favored moist forested Ruiz-Sánchez et al. 2003). On the other hand, Melissiodon is environments (Ziegler 1999). Finally, this latitudinal pattern even rarer and generally disappears much earlier than in the from more humid and forested environments in central Europe Vallès-Penedès, where it last occurs at SM, just a few meters to drier landscapes in the Iberian Peninsula is also confirmed below the marine sediments of the Langhian transgression by paleobotanical data from the earliest middle Miocene (Jovells-Vaqué & Casanovas-Vilar 2018). (Bruch et al. 2004; Fauquette et al. 2007; Jiménez-Moreno & Numerous works have provided ecological assigments of Suc 2007; Jiménez-Moreno et al. 2010). fossil rodent taxa which allow for paleoenvironmental infer- As far as rodents are concerned, Agustí (1990) and Maridet ences based on dental morphology or consider the ecological et al. (2007, 2013) recognized the unique character of the preferences and distribution of extant relatives (e.g. Daams early Miocene Vallès-Penedès faunas, particularly during the et al. 1988; Van der Meulen & Daams 1992; Van Dam & early Aragonian (MN4). In the Calatayud-Montalbán Basin, Weltje 1999; Van Dam 2006; Casanovas-Vilar & Agusti 2007). the rodent faunas at the time include a diverse assemblage The genera Democricetodon and Megacricetodon are medium to of ground-dwelling glirids, the cricetids Democricetodon and small-sized cricetids with brachyodont and bunodont molars Megacricetodon as well as the eomyid Ligerimys and occasional often assumed to have been generalists (e.g. Van Dam & terrestrial sciurids (see Van der Meulen et al. 2012; García- Weltje 1999; Casanovas-Vilar & Agustí 2007). However, Paredes et al. 2016). On the other hand, rodent faunas from Daams et al. (1988: 291) assigned different ecological prefer- France, Switzerland and Germany are enriched with a higher ences to the Iberian species of those genera, with small-sized diversity of both species and families, including arboreal forms with long mesolphs/ids (e.g. Democricetodon gaillardi sciurids (Dehmisciurus Marković et al., 2016, Miopetaurista (Schaub, 1925), Megacricetodon minor) presumably favor- Kretzoi, 1962), semiaquatic castorids (Steneofiber Geoffroy- ing more humid environments. Heshkovitz (1967) noted Saint-Hilaire, 1833), and several other “cricetids” including than in extant sigmodontine cricetids, the forms with a Anomalomys Gaillard, 1900, Eumyarion and Melissiodon (Mari- well-developed mesoloph/id (pentalophodont) are usually det et al. 2007, 2011). Some of these taxa are shared with the forest-dwellers while those in which these ridges are reduced Vallès-Penedès, including the arboreal squirrel Dehmisciurus or absent (tetralophodont) tend to prefer open country. All (present at EC, CS and SM; Aldana Carrasco 1991, 1992) the Democricetodon and Megacricetodon species present at the or the cricetids Eumyarion (Agustí 1981, 1983; Jovells-Vaqué Vallès-Penedès during the early Miocene present mesolophs/ et al. 2018) and Melissiodon (Jovells-Vaqué & Casanovas- ids of variable length, but long ridges are only particularly Vilar 2018). In addition, presumably arboreal glirids (character- common in Megacricetodon primitivus and Democricetodon ized by molars with many transverse ridges, e.g. Muscardinus gracilis. Other cricetid genera, such as Eumyarion, show Kaup, 1829, Glis Brisson, 1762, Glirudinus De Bruijn, 1966; additional transverse ridges, such as an ectomesolophid in see Van der Meulen & de Bruijn 1982) are more common in the lower molars or a long labial spur on the anterolophule Central Europe and the Vallès-Penedès than in the Iberian of the M1. Eumyarion is generally considered to have pre- inland. Yet, in the Catalan basin, taxa of Iberian affinities gener- ferred humid and forested environments (Daams et al. 1988; ally predominate over those originating from Central Europe. Van Dam 2006; Casanovas-Vilar & Agustí 2007). As already Iberian taxa include certain ground-dwelling glirids, such as said, Eumyarion weinfurteri, a Central European form, is the hypsodont Armantomys De Bruijn, 1966, only recorded recorded at the Vallès-Penedès during the early Aragonian at Turó de les Forques – level 1 and Costablanca (Ramblian, (LCV, CMV, SM), although it is not common. In contrast, MN3), and Simplomys simplicidens (de Bruijn, 1966), which is this genus is completely absent from the Iberian inland at that more widespread and abundant. These are characterized by a time, which is taken as an evidence for the exitence of more simpler molar pattern, with fewer transverse ridges as in extant arid environments. Regarding Melissiodon, the fourth cricetid ground-dwelling mouse dormice (Myomimus Ognev, 1924; genus recorded at the early Miocene Vallès-Penedès site, its see Van der Meulen & de Bruijn 1982). unique molar pattern and aberrant mandible shape, resem- The early Miocene cricetids of the Vallès-Penedès show an bling that of shrew rats, have been related to an insectivorous interesting mixture of elements belonging to the two differ- diet (Hordijk et al. 2015). Melissiodon would have preferred ent biogeographical provinces. Democricetodon hispanicus humid forested environments where its food resources would and D. decipiens are restricted to Iberia while M. primitivus be more abundant. Indeed, both Eumyarion and Melissiodon also occurs in southern France (Oliver & Peláez-Campo- are generally more common in faunas which include a high manes 2016). On the other hand, Democricetodon gracilis is number of forest-dwelling elements. Melissiodon is far more a common component of central European faunas (Daxner- common and persists for a longer time in the Vallès-Penedès Höck et al. 1998; Maridet 2003; Abdul Aziz et al. 2008, Basin as compared to the inner Iberian basins (Jovells-Vaqué & 2010; Kälin & Kempf 2009; Wessels & Reumer 2009). Casanovas-Vilar 2018). The presence of these two cricetid Other “cricetids” shared with central Europe include Eum- genera during the early Miocene in the Vallès-Penedès indi- yarion weinfurteri and Melissiodon dominans Dehm, 1950. cates affinities with Central Europe and would be indicative The former is very rare in the inner Spanish basins (Van der of forested environments. Democricetodon gracilis which is also Meulen et al. 2012), and when present it is represented by the a Central European element, may have also favored such kind species Eumyarion valencianum Daams & Freudenthal, 1974, of environments since it is abundant in faunas including a which also occurs in Valencia (Daams & Freudenthal 1974; large proportion of forest-dwellers, such as Oberdof (MN4, 420 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Austria; Daxner-Höck et al. 1998) or Sandelzhausen (MN5, D. gracilis, a species mostly recorded from central Europe, is Germany; Wessels & Reumer 2009). Therefore, the cricetid present in some sites (CS73, VI, SM, CCW), thus indicating faunas support the general impression that during the early also affinities with northern faunas (Fig. 6). The fourth, for the Miocene the Vallès-Penedès already represented a transition moment undetermined, Democricetodon species is represented zone between the forested regions of west and central Europe by just a few molars that fit within the morphological and size and the more arid inner Iberian Peninsula. However, this is a range of D. moralesi and D. mutilus. While an attribution to preliminary conclusion that should be further explored and the former species would indicate affinities with Calatayud- tested using more refined, quantitative, paleoecological analyses. Montalbán, the latter would show greater influence of central European faunas and, if confirmed, would be the first record of this species in the Iberian Peninsula. Eumyarion weinfurteri, CONCLUSIONS another central European form, is sporadically recorded. The occurrence of these cricetids may indicate the occurrence of The early Miocene rodent record of the Vallès-Penedès Basin more humid and forested environments as compared to more ranges from the Ramblian (MN3) to the end of the early inland Iberian basins. Evidence provided by large mammals, Aragonian (MN4) and is richer and more continuous than insectivores, other rodents and paleobotanical data supports previously thought. Modern cricetids are a common compo- this interpretation. nent of MN4 faunas and include four species of the genus Democricetodon (D. hispanicus, D. cf. decipiens, D. gracilis, D. sp. 4) and one of the genus Megacricetodon (M. primitivus). In FUNDING turn, these modern (crown) forms co-occur with two archaic “cricetids”: Melissiodon dominans and Eumyarion weinfurteri. This research has been funded by the Agencia Estatal de Inves- The rich and detailed rodent succession of the Calatayud-Mon- tigación from the Spanish Ministerio de Economía, Industria talbán Basin (Spain) and the Swiss Molasse Basin has shown y Competitividad and the Agencia Estatal de Investigación that Democricetodon is the first genus of modern cricetids to (AEI) from Spain/European Regional Development Fund disperse in western Europe, Megacricetodon occurring slightly of the European Union [projects CGL2017-82654-P and later (Kälin & Kempf 2009; Van der Meulen et al. 2012). CGL2016-76431-P and RYC-2013-12470 research contract However, in the Vallès-Penedès both genera appear simulta- to I.C.V.], by the Agència de Gestió d’Ajuts Universitaris i neously at LCV, the oldest MN4 site, where they are repre- de Recerca of the Generalitat de Catalunya [2017 SGR 116 sented by D. hispanicus and M. primitivus. This may evidence research group, 2018FI B1 00201 predoctoral grant to S.J.V.], a brief hiatus in the record, corresponding to the earliest MN4 and by the CERCA Programme of the Generalitat de Catalu- (equivalent to local zone B of Calatayud-Montalbán). In the nya. Fieldwork was supported by the Departament de Cultura older sites (LCV, CMV) only D. hispanicus and M. primitivus of the Generalitat de Catalunya [project 2014/100584] and are present, but later localities (CS) record additional Democri- the National Geographic Society [grant number 9640-15]. cetodon species (D. gracilis, D. sp. 4). In even younger sites, close to middle Miocene marine deposits, D. hispanicus is replaced by its descendent D. cf. decipiens (VI, SM, PA). The Acknowledgements cricetid and eomyid succession indicates close affinities with We thank V. Vicedo and L. Troya (Museu de Ciències Natu- the nearby Calatayud-Montalbán Basin, to the point that the rals de Barcelona) for their assistance when consulting the same local zonation could be used in the Vallès-Penedès (see specimens of the Villalta Collection. We are indebted to Pablo Daams et al. 1999; Van der Meulen et al. 2012). The oldest Peláez-Campomanes (Museo Nacional de Ciencias Naturales, Vallès-Penedès sites, which are devoid of cricetids other than Madrid) who provided valuable data for graphs and compari- Melissiodon, would correlate to local zone A (Ramblian). All sons. We thank the constructive comments by the editor and other sites would correlate to local zone C, zone B being reviewers of this manuscript which improved the final result. entirely missing in the Vallès-Penedès. Van der Meulen et al. (2012) subdivided zone C into subzones Ca and Cb owing to the species of Ligerimys and Democricetodon present. 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Appendix 1. — Catalogue number and length and width measurements (in mm) of Democricetodon hispanicus Freudenthal, 1967, D. gracilis Fahlbusch, 1964, D. cf. decipiens (Freudenthal & Daams, 1988) and Democricetodon sp. 4 from different early Miocene sites of the Vallès-Penedès Basin. Locality acronyms and measurement abbreviations are explained in section Abbreviations of the main text. For details on the measurement methods see section Material and methods, and Daams & Freudenthal (1988: 42, fig. 1): https://doi.org/10.5852/cr-palevol2021v20a22_s1 Appendix 2. — Morphotype frequency tables for the Democricetodon Fahlbusch, 1964 species. For locality acronyms see section Abbreviations of the main text. For details on morphotype coding see section Material and methods, and references therein. Appendix 2A. — Anterocone M1. Anterocone M1 Species Site N D. decipiens SM 1 – 1 VI 1 – 1 D. gracilis SM 2 – 2 D. hispanicus CS74 8 1 9 CMV3 5 – 5 LCV1 1 – 1 EC 1 – 1 Appendix 2B. — Anterolophule M1. Anterolophule M1 Species Site N SM – 1 – 1 D. decipiens VI 1 – – 1 D. gracilis SM 2 – – 2 CS74 8 1 – 9 CMV3 5 – – 5 D. hispanicus LCV1 – – 1 1 EC 1 – – 1 Appendix 2C. — Protolophule M1. Protolophule M1 Species Site N SM 1 – – 1 D. decipiens VI 1 – – 1 D. gracilis SM – 2 – 2 CS74 7 2 – 9 D. hispanicus CMV3 2 3 – 5 LCV1 1 – – 1 EC – 1 – 1 426 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Appendix 2D. — Ectoloph on the Paracone M1. Ectoloph on the Paracone M1 Species Site N D. decipiens SM 1 1 VI 1 1 D. gracilis SM 2 2 D. hispanicus CS74 6 3 9 CMV3 4 4 LCV1 2 2 EC 1 1 Appendix 2E. — Mesoloph M1. Mesoloph M1 Species Site N D. decipiens SM 1 – – – 1 VI 1 – – – 1 D. gracilis SM – – 2 – 2 D. hispanicus CS74 4 5 – – 9 CMV3 – – 4 – 4 LCV1 – – 2 – 2 EC – 1 – – 1 Appendix 2F. — Metalophule M1. Metalophule M1 Species Site N D. decipiens SM 1 – – – 1 VI 1 – – – 1 D. gracilis SM 2 – – – 2 D. hispanicus CS74 9 – – – 9 CMV3 4 – – – 4 LCV1 2 – – – 2 EC – – 1 – 1 COMPTES RENDUS PALEVOL • 2021 • 20 (22) 427 Jovells-Vaqué S. & Casanovas-Vilar I. Appendix 2G. — Protolophule M2. Protolophule M2 Species Site N D. sp. 4 MOR1 – – 1 – – 1 D. decipiens SM – – 1 – – 1 PA – – 1 – – 1 VI – 2 – – – 2 D. gracilis SM 1 – 1 – – 2 D. hispanicus CS74 6 – 6 – – 12 CMV3 – – 7 – – 7 LCV1 2 1 6 – – 9 EC 1 2 – – – 3 Appendix 2H. — Ectoloph on the Paracone M2. Ectoloph on the Paracone M2 Species Site N D. sp. 4 MOR1 1 – 1 D. decipiens SM – 1 1 PA – – 1 VI – 2 2 D. gracilis SM 2 – 2 D. hispanicus CS74 8 4 12 CMV3 – 7 7 LCV1 2 – 2 EC 2 1 3 Appendix 2I. — Mesoloph M2. Mesoloph M2 Species Site N D. sp. 4 MOR1 – 1 – – 1 D. decipiens SM 1 – – – 1 PA – – 1 – 1 VI – 2 – – 2 D. gracilis SM 2 – – – 2 D. hispanicus CS74 6 5 1 – 12 CMV3 6 1 – – 7 LCV1 2 – – – 2 EC 1 – 2 – 3 428 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Appendix 2J. — Metalophule M2. Metalophule M2 Species Site N D. sp. 4 MOR1 1 1 D. decipiens SM 1 1 PA 1 1 VI 2 2 D. gracilis SM 1 1 D. hispanicus CS74 11 1 12 CMV3 7 7 LCV1 2 2 EC 2 1 3 Appendix 2K. — Lingual Anterolophid m1. Lingual Anterolophid m1 Species Site N D. sp. 4 CS73 – 1 1 D.decipiens VI – 2 2 D. gracilis SM – 1 1 VI – 2 2 CS73 – 1 1 CCW – 1 1 D. hispanicus CS74 8 4 – CS72 2 6 8 CSU 1 – 1 CMV3 4 – 4 CMV1 1 – 1 LCV1 6 – 6 Appendix 2L. — Metalophulid m1. Metalophulid m1 Species Site N D. sp. 4 CS73 1 – – 1 D. decipiens VI 2 – – 2 D. gracilis SM 1 – – 1 VI 2 – – 2 CS73 1 – – 1 CCW 1 – – 1 D. hispanicus CS74 5 6 1 12 CS72 7 1 – 8 CSU – 1 – 1 CMV3 4 – – 4 CMV1 1 – – 1 LCV1 5 1 – 6 COMPTES RENDUS PALEVOL • 2021 • 20 (22) 429 Jovells-Vaqué S. & Casanovas-Vilar I. Appendix 2M. — Metaconid ridge m1. Metaconid ridge m1 Species Site N D. sp. 4 CS73 1 – 1 D. decipiens VI – 2 2 D. gracilis SM 1 – 1 VI – 2 2 CS73 – 1 1 CCW – 1 1 CS74 – 12 12 D. hispanicus CS72 – 8 8 CSU – 1 1 CMV3 – 4 4 CMV1 1 – 1 LCV1 – 6 6 Appendix 2N. — Mesolophid m1 Mesolophid m1 Species Site N D. sp. 4 CS73 1 – – – 1 D. decipiens VI – 2 – – 2 D. gracilis SM 1 – – – 1 VI 2 – – – 2 CS73 1 – – – 1 CCW 1 – – – 1 D. hispanicus CS74 3 6 3 – 12 CS72 – 5 3 – 8 CSU – 1 – – 1 CMV3 1 2 1 – 4 CMV1 – 1 – – 1 LCV1 – 3 3 – 6 430 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Appendix 2O. — Anterosinusid m2 Anterosinusid m2 Species Site N D. sp. 4 MOR1 1 – 1 D. decipiens PA 1 – 1 VI 2 1 3 D. gracilis SM 1 – 1 CCW 1 – 1 D. hispanicus SM – 2 2 CS74 5 11 16 CSU 3 1 4 CMV3 2 2 4 CMV2 – 1 1 LCV1 5 6 11 Appendix 2P. — Metaconid ridge m2 Metaconid ridge m2 Species Site N D. sp. 4 MOR1 – 1 1 D. decipiens PA – 1 1 VI – 3 3 D. gracilis SM – 1 1 CCW 1 1 D. hispanicus SM – 2 2 CS74 3 13 16 CSU – 4 4 CMV3 2 2 4 CMV2 1 – 1 LCV1 6 4 10 Appendix 2Q. — Mesolophid m2 Mesolophid m2 Species Site N D. sp. 4 MOR1 1 – – – 1 D. decipiens PA 1 – – – 1 VI 1 – – 2 3 D. gracilis SM – 1 – – 1 CCW – 1 – – 1 D. hispanicus SM 2 – – – 2 CS74 8 4 – 4 16 CSU 3 – – 1 4 CMV3 – 2 1 1 4 CMV2 – – 1 – 1 LCV1 4 2 6 – 12 COMPTES RENDUS PALEVOL • 2021 • 20 (22) 431 Jovells-Vaqué S. & Casanovas-Vilar I. Appendix 3 — Catalogue number and length and width measurements (in mm) of Megacricetodon primitivus (Freudenthal, 1963) from different early Miocene sites of the Vallès-Penedès Basin. Locality acronyms and measurement abbreviations are explained in section Abbreviations of the main text. For details on the measurement methods see section Material and methods, and Daams & Freudenthal (1988: 42, fig. 1): https://doi.org/10.5852/cr-palevol2021v20a22_s2 Appendix 4. — Morphotype frequency tables for Megacricetodon primitivus (Freudenthal, 1963). For locality acronyms see section Abbreviations of the main text. For details on morphotype coding see section Material and methods, and references therein. Appendix 4A. — Anterocone M1. Anterocone M1 Site N PA – 6 15 – 21 VI – 4 12 9 25 CS74 – 4 1 – 5 CS73 – 5 – – 5 CSU – 1 – – 1 CMV3 – 6 6 1 13 CMV2 – 2 – – 2 CMV1 – – 1 – 1 LCV1 – 9 4 – 13 Appendix 4B. — Anterior platform of the M1 anterocone. Anterior platform of the M1 anterocone M1 Site N PA 8 6 14 VI 14 9 23 CS74 4 1 5 CS73 4 4 8 CSU – 1 1 CMV3 7 6 13 CMV2 2 – 2 CMV1 1 – 1 LCV1 6 4 10 432 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Appendix 4C. — Labial spur of the M1 anterolophule. Labial spur of the M1 anterolophule Site N PA 1 3 16 20 VI 7 7 16 30 CS74 1 1 4 6 CS73 – 3 3 6 CSU – 1 – 1 CMV3 1 4 5 10 CMV2 – – 2 2 CMV1 – – 1 1 LCV1 2 4 9 15 Appendix 4D. — Protolophule M1. Protolophule M1 Site N PA 20 – – – VI 29 – 1 – CS74 5 – – 5 CS73 6 – – 6 CSU 2 – – 2 CMV3 12 – – – CMV2 2 – – 2 CMV1 1 – – 1 LCV1 14 – – – Appendix 4E. — Ectoloph M1. Ectoloph M1 Site N PA 3 5 13 21 VI 10 6 12 28 CS74 – 2 1 3 CS73 1 1 4 6 CSU – – 2 2 CMV3 5 3 5 13 CMV2 1 1 – 2 CMV1 – – 1 1 LCV1 3 8 3 14 COMPTES RENDUS PALEVOL • 2021 • 20 (22) 433 Jovells-Vaqué S. & Casanovas-Vilar I. Appendix 4F. — Ectoloph-Mesoloph connection M1. Ectoloph – Mesoloph connection M1 Site N PA – 1 19 20 VI – 22 7 22 CS74 – – 5 5 CS73 – – 6 6 CSU – 2 – 2 CMV3 – 12 – 12 CMV2 – – 2 1 CMV1 – – 1 1 LCV1 – – 14 14 Appendix 4G. — Mesoloph M1. Mesoloph M1 Site N PA 11 6 3 – 20 VI 18 7 5 – 30 CS74 1 3 2 – 5 CS73 2 3 1 – 6 CSU – 2 – – 2 CMV3 8 8 1 – 17 CMV2 1 1 – – 2 CMV1 – 1 – – 1 LCV1 9 4 1 – 14 Appendix 4H. — Metalophule M1. Metalophule M1 Site N PA 1 4 12 17 VI 3 6 19 28 CS74 1 3 1 5 CS73 – 2 4 6 CSU – 2 – 2 CMV3 – 6 8 14 CMV2 – 1 1 2 CMV1 – 1 – 1 LCV1 – 14 – 14 434 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Appendix 4I. — Posterosinus M1 Posterosinus M1 Site N PA 1 19 – 20 VI 12 16 – 28 CS74 2 3 – 5 CS73 – 6 – 6 CSU – 2 – 2 CMV3 5 6 3 14 CMV2 – 2 – 2 CMV1 – 1 – 1 LCV1 7 5 1 13 Appendix 4J. — Protolophule M2. Protolophule M2 Site N PA 13 – – 3 3 16 VI 22 13 3 2 2 38 CS74 1 1 – – – 2 CS72 1 1 – – – 2 CSU 1 – – – – 1 CMV3 8 4 1 – – 13 CMV1 – 1 – – – 1 LCV1 10 6 – – 1 17 Appendix 4K. — Ectoloph M2. Ectoloph M2 Site N PA 2 7 7 16 VI 14 11 10 35 CS74 – 2 – 2 CS72 – 1 1 2 CSU 1 – – 1 CMV3 4 8 1 13 CMV1 – – 1 1 LCV1 4 9 3 16 COMPTES RENDUS PALEVOL • 2021 • 20 (22) 435 Jovells-Vaqué S. & Casanovas-Vilar I. Appendix 4L. — Ectoloph-Mesoloph connection M2. Ectoloph-Mesoloph connection M2 Site N PA 5 11 11 VIO 8 31 39 CS74 2 2 CS72 2 2 CSU 1 1 CMV3 13 13 CMV1 1 1 LCV1 17 17 Appendix 4M. — Mesoloph M2. Mesoloph M2 Site N PA 7 9 – – 16 VI 19 20 6 – 35 CS74 – 2 – – 2 CS72 – 2 – – 2 CSU – – 1 – 1 CMV3 3 8 3 – 14 CMV1 – 1 – – 1 LCV1 7 5 4 1 17 Appendix 4N. — Sinus M2. Sinus M2 Site N PA 12 4 16 VI 30 9 39 CS74 1 1 2 CS72 2 – 2 CSU 1 – 1 CMV3 – 13 13 CMV1 1 – 1 LCV1 – 16 16 436 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Appendix 4O. — Metalophule M2. Metalophule M2 Site N PA 6 6 4 – 16 VI 7 21 7 – 35 CS74 1 1 – – 2 CS72 2 – – – 2 CSU 1 – – – 1 CMV3 12 2 – – 14 CMV1 – 1 – – 1 LCV1 17 – – – 17 Appendix 4P. — Anteroconid m1. Anteroconid m1 Site N SM 1 – – 1 PA 4 3 – 7 VI 25 – – 25 CS74 13 2 – 15 CS73 3 – – 3 CS72 5 – – 5 CSU 1 – – 1 CMV3 17 – – 17 CMV1 5 – – 5 LCV1 11 – – 11 Appendix 4Q. — Labial spur on m1 anterolophulid. Labial spur on m1 anterolophulid Site N SM 1 – – 1 PA 10 – – 10 VI 29 – – 29 CS74 11 6 – 17 CS73 3 – – 3 CS72 6 – – 6 CSU 1 – – 1 CMV3 21 – – 21 CMV1 5 – – 5 LCV1 17 – – 17 COMPTES RENDUS PALEVOL • 2021 • 20 (22) 437 Jovells-Vaqué S. & Casanovas-Vilar I. Appendix 4R. — Metalophulid m1 Metalophulid m1 Site N SM 1 – – 1 PA 10 – – 10 VI 32 – – 32 CS74 17 – – 17 CS73 3 – – 3 CS72 6 – – 6 CSU 1 – – 1 CMV3 20 – – 20 CMV1 5 – – 5 LCV1 17 – – 17 Appendix 4S. — Mesolophid m1. Mesolophid m1 Site N SM – – 2 – 2 PA 6 3 1 – 10 VI 1 17 7 4 29 CS74 1 2 13 1 17 CS73 – 2 1 – 3 CS72 – 1 4 1 6 CSU – – 1 – 1 CMV3 – 11 7 – 18 CMV1 – 4 1 – 5 LCV1 1 9 7 – 17 438 COMPTES RENDUS PALEVOL • 2021 • 20 (22) Dispersal and early evolution of the first modern cricetid rodents in Western Europe Appendix 4T. — Lingual anterolophid lingual m2. Lingual anterolophid m2 Site N PA 5 10 2 17 VI 8 21 2 31 CS74 2 7 2 11 CS72 – 3 – 3 CSU – 1 – 1 CMV3 4 5 2 11 CMV2 1 – – 1 CMV1 1 – – 1 LCV1 3 6 8 17 Appendix 4U — Mesolophid m2. Mesolophid m2 Site N PA 4 12 4 – 20 VI – 24 6 4 34 CS74 – 2 7 2 11 CS72 – 2 1 – 3 CSU – 1 – – 1 CMV3 – 4 5 2 11 CMV2 3 – – – 3 CMV1 – 1 – – 1 LCV1 3 8 5 1 17 COMPTES RENDUS PALEVOL • 2021 • 20 (22) 439