comptes rendus palevol 2021 z 20 z 28 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) COUVERTURE / COVER : Mandible of Prolagus sp. from the early Pliocene of Hungary. Author of photos: Stanislav Čermák. 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®. Les articles ainsi que les nouveautés nomenclaturales publiés dans Comptes Rendus Palevol sont référencés par / Articles and nomenclatural novelties published in Comptes Rendus Palevol are registered on: – ZooBank® (http://zoobank.org) Comptes Rendus Palevol est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris et l’Académie des sciences, Paris Comptes Rendus Palevol is a fast track journal published by the Museum Science Press, Paris and the Académie des sciences, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Geodiversitas, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie. L’Académie des sciences publie aussi / The Académie des sciences also publishes: Comptes Rendus Mathématique, Comptes Rendus Physique, Comptes Rendus Mécanique, Comptes Rendus Chimie, Comptes Rendus Géoscience, Comptes Rendus Biologies. 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 Prolagus Pomel, 1853 (Lagomorpha, Mammalia) in the framework of the Pliocene faunal rearrangements in central Europe Stanislav ČERMÁK Institute of Geology of the Czech Academy of Sciences, Rozvojová 269, 165 00 Prague 6 (Czech Republic) cermaks@gli.cas.cz (corresponding author) Chiara ANGELONE Department of Sciences, University Roma Tre, Largo S. L. Murialdo 1, 00146 Roma (Italy) and Institut Català de Paleontologia Miquel Crusafont, Edifici Z ICTA–ICP, Carrer de les Columnes s/n, Campus de la Universitat Autònoma de Barcelona, 08193 Cerdanyola del Vallès, Barcelona (Spain) Blanca MONCUNILL-SOLÉ Centro de Investigacións Científicas Avanzadas (CICA), As Caballeiras s/n, Campus de Elviña, Universidade da Coruña, 15071 A Coruña (Spain) and Department of Sciences, University Roma Tre, Largo S. L. Murialdo 1, 00146 Roma (Italy) Submitted on 21 February 2020 | Accepted on 25 May 2020 | Published on 28 June 2021 urn:lsid:zoobank.org:pub:4FD1B8AF-7957-469E-A627-8126691743C3 Čermák S., Angelone C. & Moncunill-Solé B. 2021. — Prolagus Pomel, 1853 (Lagomorpha, Mammalia) in the framework of the Pliocene faunal rearrangements in central Europe. Comptes Rendus Palevol 20 (28): 597-617. https://doi.org/10.5852/cr-palevol2021v20a28 ABSTRACT Pliocene occurrences of Prolagus Pomel, 1853 in central Europe represent anomalies out of the peri-Mediterranean area, at that time the core distribution of the genus. Though known for several decades, those materials never underwent a general revision. The detailed analysis and comparison of all available materials performed here revealed two phenotypic entities: 1) Prolagus bilobus Heller, 1936 (Gundersheim localities, Raciszyn 1), for which were defined additional diagnostic characters and ontogenetic patterns of variation (d3/p3 and mandible); and 2) Prolagus sp. (Beremend 26/39), probably a new species. All the available occurrences are dated to MN15b. The morphological trends towards the reduction of p3 entoconid and of enamel folding evidenced in Pliocene Prolagus of western Europe cannot be recognized in coeval central European forms. Evidently, P. bilobus and Prolagus sp. do not pertain to the western European clade, whose separation is known since the early late Miocene. KEY WORDS We hypothesize that at least P. bilobus originated from a dispersal of Prolagus from south-eastern re- Prolagidae, Prolagus bilobus, gions of Europe rather than from an autochthonous speciation of isolated populations left as a relict Gundersheim, after the southward displacement of Prolagus distribution area. The dispersal is likely to be related to Raciszyn 1, Beremend, the Pliocene global environmental changes during which extensive faunal rearrangements took place palaeobiogeography. in Europe, in particular to those near the early/late Pliocene boundary. COMPTES RENDUS PALEVOL • 2021 • 20 (28) © Publications scientifiques du Muséum et/and Académie des sciences, Paris. www.cr-palevol.fr 597 Čermák S. et al. RÉSUMÉ Prolagus Pomel, 1853 (Lagomorpha, Mammalia) dans le cadre des réarrangements fauniques du Pliocène de l’Europe centrale. Le peu de données dont nous disposons sur le Prolagus Pomel, 1853 du Pliocène d’Europe centrale représentent des anomalies dans la distribution géographique du genre, qui, à cette époque, est limité à la zone péri-Méditerranéenne. Bien que la présence de Prolagus dans le Pliocène d’Europe centrale est connue depuis plusieurs décennies, de tels matériels n’ont jamais fait l’objet d’une révision géné- rale. Une analyse détaillée et une comparaison de tous les matériels disponibles étudiés ici ont permis d’identifier deux éntités phénotypiques : 1) Prolagus bilobus Heller, 1936 à Gundersheim (plusieurs localités) et Raciszyn 1, pour lequel des caractères diagnostiques additionnels et des modèles onto- génétiques de variation (d3/p3 et mandibule) ont été définis ; et 2) Prolagus sp. à Beremend 26/39, qui représente probablement une nouvelle espèce. Toutes les occurrences disponibles sont datées du MN15b. Les tendances morphologiques vers une réduction de l’entoconide de p3 et une diminution du repli de l’émail, mises en évidence dans les espèces pliocènes de Prolagus d’Europe occidentale, n’ont pas été reconnues dans des formes contemporaines d’Europe centrale. Prolagus bilobus et Prolagus sp. n’appartiennent manifestement pas au clade ouest-européen, dont la séparation est connue depuis le début du Miocène supérieur. Notre hypothèse est qu’au moins P. bilobus trouve son origine dans une dispersion des populations à partir de l’Europe sud-orientale plutôt que, comme on le croyait MOTS CLÉS autrefois, dans une spéciation autochtone de populations isolées, laissées telles des vestiges à la suite Prolagidae, Prolagus bilobus, du déplacement vers le sud de l’aire de distribution géographique de Prolagus. Cette dispersion est Gundersheim, probablement liée aux changements environnementaux à l’échelle globale survenus au Pliocène, et Raciszyn 1, Beremend, responsables de changements environnementaux globaux pendant lesquels des réarrangements fau- paléobiogéographie. niques majeurs ont pris place en Europe, et particulièrement à la transition Pliocène inférieur/supérieur. INTRODUCTION Shushpanov 1988; Averianov & Tesakov 1998; Tesakov & Averianov 2002). Prolagus Pomel, 1853 represents a successful, long-lasting, We decided to focus our attention to Pliocene central Euro- species-rich lagomorph genus that during most of the Mio- pean Prolagus, in order to summarize the available materials, cene was widely distributed throughout Europe and formed to clarify their taxonomic status, and to try to unravel the a significant part of the small mammal assemblages (López reasons that led to the palaeobiogeographical distribution Martínez 1974, 1989, 2001). Probably, since the latest Mio- observed after the Miocene. cene, its geographical range started to dwindle. The Pliocene is a critical moment in the evolution of Prolagus. The changes in its palaeobiogeographical distribution from pan-European THE LOCALITIES: HISTORICAL BACKGROUND to patchy and mainly peri-Mediterranean distribution become AND GEOLOGICAL SETTING evident in this period. The Pliocene record of Prolagus in the peri-Mediterranean area is quite common and well doc- The Pliocene record of Prolagus in Central Europe is limited umented, in contrast to central Europe, where the genus is to the following localities; their geographical locations are very rare and occurs sporadically in a few localities. shown in Figure 1. Isolated Pliocene populations of Prolagus in central Europe have been known since the early 20th century (Heller 1936). GUNDERSHEIM The proven occurences of the genus have been limited until Most of the fossil faunas at Gundersheim (Alzey-Worms dis- now to Prolagus bilobus Heller, 1936 documented from Gunder- trict, Rheinland-Pfalz, Germany) come from three quarries, sheim sites (MN15b) in Germany and Raciszyn 1 (MN15b) in located SW from the homonymous village. A study of the Poland (Heller 1936; Fejfar & Storch 1990; Fejfar et al. 2006; fossil record from Gundersheim was first performed by Hel- Fostowicz-Frelik 2010; Čermák & Angelone 2013). Apart from ler (1936). However, although the material originated from punctual taxonomic disputes (cfr. Fostowicz-Frelik 2010 vs several fissures of quarries, F. Heller published the different Čermák & Angelone 2013), these remains have never been faunas considering them as one assemblage, with only sporadic the subject of a detailed, dedicated study. notes concerning their original provenance. Thus, the exact Some studies exist about the taxonomy and phylogeny of location, age, and name of each site/assemblage have since then Pliocene Prolagus from western Europe (López Martínez & remained unclear. Until the 1960s, all the samples were gener- Thaler 1975; López Martínez 1989; Angelone 2008a), and ally considered as pertaining to one faunal unit, and referred to as well, some enigmatic isolated remains have been reported simply as “Gundersheim”. Eventually, Kretzoi (1962) distin- from eastern Europe (Agadjanian & Erbajeva 1983; Erbajeva & guished two faunal units in Heller’s original material: an older 598 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe N 2 4 1 5 3 200 km FIG. 1. — Geographical localization of Pliocene Prolagus-bearing localities in central Europe: 1, Gundersheim 1 (sensu Tobien 1980; age uncertain, most likely MN15b), Gundersheim-Findling (MN15b), Gundersheim-fissure 4 (MN15b); 2, Raciszyn 1 (MN15b); 3, Beremend 26 and 39 (MN15b); 4, Wölfersheim (MN15b); 5, Kisláng (Prolagus age uncertain, most likely MN15b). Black circles, available and studied material; white circles, unavailable material mentioned in literature (not figured, nor described). one, correlated with the Csarnotanian (sensu Kretzoi 1962; Čermák & Wagner (2013) and Hypolagus petenyii Čermák & c. MN15b), called Gundersheim 1 by Tobien (1980), and a Fladerer in Čermák, 2009 (p3 dex [morphotype I/a/A], 2 lower younger one, correlated with the late Villányian (sensu Kret- molariforms); Gundersheim-fissure 4 – Leporidae gen. et sp. zoi 1962; c. MN17), called Gundersheim 2 by Tobien (1980). indet. (fragmentary lower molariform); and “Hasenfunds- Tobien (1980) also numbered particular Gundersheim telle I” (after label by Heller) – Hypolagus published by Hel- fissures. Since the 1950s, he discovered and studied 22 fis- ler (1936: 137-139; figs 1-2, pl. XI: 13-14) as H. brachygnathus sures of which five yielded identifiable faunal remains. The and reassigned here as H. cf. brachygnathus (Kormos, 1930) most interesting was fissure no. 4, in which were recovered (2 mandibles – sin with p3-p4 [morphotype II/c/D] and dex Trilophomys schaubi, Bjoernkurtenia canterranensis, Baranomys with p3 [(I/II)/c/C]). longidens, Germanomys weileri, and Mimomys gracilis, which indicate a late Ruscinian age (MN15b). This site, published RACISZYN 1 in detail by Fejfar et al. (2006), is known as Gundersheim- The locality Raciszyn 1 corresponds to a small abandoned fissure 4 (Gundersheim 3 sensu Koenigswald & Tobien 1990). quarry located c. 500 m W from the village of Raciszyn (district Tobien (1980) assumed that the most of Heller’s older faunal Gmina Działoszyn, province Łódź, central Poland). The fossil unit (Gundersheim 1) most likely originated from this fissure. material comes from the terra rossa filling of a collapsed cave The last fauna at Gundersheim was discovered in 1976 in formed in Oxfordian limestones. Preliminary information about an isolated block of calcified red loam found in the open field the locality and its fauna was provided by Kowalski (1990), near the Rosengartenbruch quarry. The locality, known as Nadachowski (1990), Sulimski & Szynkiewicz (1994), Szyn- Gundersheim-Findling (Gundersheim 4 sensu Koenigswald & kiewicz & Jagiełło (2015), and Nadachowski et al. (2015). Tobien 1990), yielded a faunal assemblage analogous to the The locality yielded a fossil assemblage consisting of about 40 one found in Gundersheim-fissure 4 described by Fejfar mammalian taxa (Sulimski in Szynkiewicz & Jagiełło 2015). et al. (2006), and indicating a late Ruscinian (MN15b) age The assemblage includes biostratigraphically valuable taxa such as well. Rodents from this locality were described in detail as a small Mimomys of the gracilis-stehlini lineage (with prev- by Fejfar & Storch (1990), insectivores by Dahlmann & alence of primitive forms), Mimomys hassiacus, Baranomys or Storch (1996), and prolagid lagomorphs partly by Čermák & Trilophomys (Nadachowski 1990; Szynkiewicz & Jagiełło 2015; Angelone (2013). Nadachowski et al. 2015), indicating a late Ruscinian (MN15b) Apart from Prolagus, herein studied, Gundersheim locali- age (Kowalski 1990; Nadachowski et al. 2015). The remains ties yielded other lagomorph taxa: Gundersheim-Findling – of Prolagus addressed in this paper were first studied by Fos- Pliopentalagus dietrichi (Fejfar, 1961) published in detail by towicz-Frelik (2010) and described as a new species, Prolagus COMPTES RENDUS PALEVOL • 2021 • 20 (28) 599 Čermák S. et al. osmolskae Fostowicz-Frelik, 2010. Eventually, Čermák & and described by Kretzoi (1954) is currently curated in the Angelone (2013) revised and synonymized the species with collection of the MFGI, but the last surveys (SČ in 2005) Prolagus bilobus. The only other lagomorph species found in were not successful to track down the specimen. At any rate, Raciszyn 1 is Hypolagus petenyii (Nadachowski et al. 2015). the Kisláng mammalian assemblage is a mixture of material of different geological ages with a minimum estimated range BEREMEND between c. 1.7-3.5 Ma (Mayhew 2012). Thus, the age of the The limestone quarry at Beremend is situated in the western- p3 reported by Kretzoi (1954) is unclear. Nevertheless, con- most part of the Villány Hills (Baranya region, Hungary), an sidering the presence of Mimomys cf. hassiacus in the locality area palaeontologically investigated since the half of the 19th (Mayhew 2012), the late Ruscinian (MN15b) age cannot be century. The first fossiliferous fissures in the quarry were dis- excluded (see for details Čermák & Angelone 2013). covered by S. J. Petényi in 1847 (Petényi 1864; Kordos 1991). During the 20th century, the quarry yielded several dozens of vertebrate-bearing sites, whose age span from the early Plio- MATERIAL AND METHODS cene to the early Pleistocene (see Kretzoi 1956, 1959, 1962; Jánossy 1986; Kordos 1991; Pongrácz 1999; Császár & Kor- All the material from the Gundersheim localities is stored in dos 2004 for details). The material, comprising remains of the the collections of the SMF and the material from Raciszyn 1 genus Prolagus studied herein, was collected by L. Pongrácz is curated in the collections of the ZPAL. The material from from fissure fillings Beremend 26 and 39. The site no. 26 Beremend 26 and 39 comes from the private collection of yielded a very extensive and diverse fossil assemblage of mam- L. Pongrácz (Győr, Hungary) and is temporarily housed in the mals including the biostratigraphical markers Mimomys gracilis, collection of the GLI. Following Čermák & Angelone (2013), Dolomys nehringi, and Propliomys hungaricus (cf. Császár & we used Gundersheim 1 and Gundersheim 2 for Heller’s original Kordos 2004; Čermák 2007; Čermák unpubl. data). Such taxa localities/assemblages (sensu Tobien 1980), and Gundersheim- indicate a late Ruscinian (MN15b) age for Beremend 26, i.e., fissure 4 and Gundersheim-Findling for localities published slightly younger than Csarnóta 2 (L. Kordos pers. comm.). by Fejfar et al. (2006) and Fejfar & Storch (1990). Apart from Prolagus, the locality yielded also other lagomorphs, For teeth nomenclature and metrics we follow López namely Ochotonoma csarnotana Kretzoi, 1959 (partly studied Martínez (1989) and Angelone & Sesé (2009), for jaws we by SČ; see Čermák 2007) and Hypolagus petenyii. follow Wible (2007) and Čermák (2009). Drawings and mea- In comparison with Beremend 26, the fossil record from sures were taken with the aid of a Dino-Lite digital microscope Beremend 39 is much scantier and less diverse, consisting of and of a binocular microscope with ocular micrometer. Dental “only” about 15 mammalian taxa (Pongrácz in litt.; Čermák & measurements were taken as maximum antero-posterior or Wagner 2013). Nevertheless, the presence of Beremendia fis- bucco-lingual dimensions of the respective two-dimensional sidens, Eliomys intermedius, M. cf. gracilis, Propliomys hunga- dental structures orthogonal to prismatic shaft. For consis- ricus, H. petenyii, and Trischizolagus dumitrescuae, together tency, only adult specimens (recognized by their prismatic with the evolutionary degree of the arvicolids, indicate also tooth shape) were used for interspecific size comparisons. All a late Ruscinian (MN15b) age, analogous to Beremend 26 measured data are given in millimeters [mm]. (Čermák & Wagner 2013). The biostratigraphic terminology used in this paper follows Fejfar & Heinrich (1983) and Fejfar et al. (1998). The term OTHER LOCALITIES “Central Europe” is understood here as a geographic region in In addition to the above mentioned localities, there are other the center of Europe including Germany, Switzerland, Liech- two relevant sites from which the presence of Prolagus was tenstein, Austria, Poland, the Czech Republic, Slovakia, and reported. Unfortunately, however, we were not successful to Hungary. Country abbreviations follow ISO 3166-1 alpha-2 track down the lagomorph material from those localities. The codes. Following the recommendations of the International taxonomic position of those samples remained thus unclear. Commission of Stratigraphy, the first letter of informal sub- – Wölfersheim (Wetterau district, Germany; MN15b) – the epochs is not capitalized. occurrence of P. bilobus in this locality was reported in a fau- Interspecific comparisons were made with the early Miocene nal list by Tobien (1977). Dahlmann (2001) studied in detail to Pleistocene European species of Prolagus using original or small mammals from this locality, but not lagomorphs, which additional materials (indicated by “*”), or on a bibliographical were reported only in a faunal list (ibid.: 95) with reference to basis (unless otherwise stated, data were taken from the original Tobien’s list (1977). The collection of mammals from Wölfer- descriptions of species): *P. oeningensis (König, 1825); *P. sar- sheim is deposited in the SMF, but lagomorphs are missing. dus (Wagner, 1829); P. vasconiensis Viret in Roman & Viret, – Kisláng (Polgárdi district, Hungary; MN15 or MN17 1930; *P. savagei Berzi, 1967; *P. crusafonti López Martínez in according to Mayhew 2012) – Kretzoi (1954) reports from López Martínez & Thaler, 1975; P. depereti López Martínez in this locality a sole left p3 of Prolagus. Based on the presence López Martínez & Thaler, 1975; *P. figaro López Martínez in of the crochet (“Der Sporn am Mittelgraben der Kauflache”), López Martínez & Thaler, 1975; *P. ibericus López Martínez in Kretzoi (1954: 247) hypothesized a taxonomic proximity of López Martínez & Thaler, 1975; *P. michauxi López Martínez the Kisláng specimen to Prolagus from Gundersheim, and in López Martínez & Thaler, 1975; *P. tobieni López in López referred to it as P. cf. bilobus. The material of Kisláng collected Martínez et al., 1977; *P. sorbinii Masini, 1989; P. fortis López 600 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe Martínez & Sesé in Álvarez-Sierra et al., 1990; P. caucasicus Martínez & Thaler (1975), revised by Čermák & Angelone (2013). Averianov & Tesakov, 1998; *P. italicus Angelone, 2008a; Gundersheim (probably Gundersheim 1 sensu Tobien 1980; age uncertain, most likely MN15b), Rheinland-Pfalz, Germany (see *P. latiuncinatus Angelone & Čermák, 2015; and *P. pannoni- Čermák & Angelone 2013 for details). cus Angelone & Čermák, 2015. Other metric data were taken from Álvarez-Sierra et al. (1990), Tesakov & Averianov (2002), EMENDED DIAGNOSIS. — See Čermák & Angelone (2013: 49). and Angelone & Veitschegger (2015). MATERIAL. — Gundersheim (probably Gundersheim 1, original material of F. Heller, partly published in Heller [1936]): 1 p3 sin, ABBREVIATIONS SMF 1996/136; 1 fragment of mandibular body dex with p4-m2, Institutions SMF 1994/895; 1 talonid of p4 dex, SMF 1994/896; 1 talonid of GLI Institute of Geology of the Czech Academy of Sci- p4 or m1 dex, SMF 1994/897; 1 trigonid of p4 or m1 dex, SMF 1996/137; 1 mandibular body dex with p4-m2, SMF 1996/156; ences, Prague; 1 mandibular ramus dex with articular head, SMF 1996/157; 1 MFGI Magyar Földtani és Geofizikai Intézet [Hungarian mandibular ramus dex with p4-m1, SMF 1996/158. Gundersheim- Institute of Geology and Geophysics], Budapest; fissure 4 (material published by Fejfar et al. 2006): 2 P2 sin, SMF SMF Senckenberg Museum, Frankfurt; 2005/1, 2; 1 P3 sin, SMF 2005/3; 1 d3 dex, SMF 2005/4; 1 p3 sin, ZPAL Institute of Palaeobiology, Polish Academy of Sci- SMF 2005/5; 1 fragment of mandibular body dex, SMF 2005/6. Gundersheim-Findling: 11 I1 sin, SMF 1985/171–181; 6 I1 dex, ences, Warszawa. SMF 1985/182-187; 2 P2 sin, SMF 1985/70, 71; 5 P2 dex, SMF 1985/72-76; 5 P3 sin, SMF 1985/77, 80, 82, 84, 85; 7 P3 dex, Teeth SMF 1985/78, 79, 81, 83, 86–88; 8 P4 sin, SMF 1985/89-96; 12 D/d Upper/lower deciduous teeth; P4 dex, SMF 1985/97-108; 1 fragment of P4 sin, SMF 1985/242; I/i Upper and lower incisors; 4 fragments of P4 dex, SMF 1985/240, 241, 243, 244; 9 M1 sin, SMF 1985/109-112, 115, 124, 126, 247, 248; 17 M1 dex, SMF M/m Upper/lower molar; 1985/113, 114; 116-123, 125, 141, 142, 245, 246, 249, 255; 4 M2 P/p Upper/lower premolar. sin, SMF 1985/129, 133, 135, 140; 12 M2 dex, SMF 1985/127, 128, 130-132, 134, 136-139, 178, 179; 3 enamel fragments of up- Measurements per molariforms, SMF 1985/252-254; 6 fragments of mandibular AA Partial width; bodies dex with various teeth, SMF 1985/165-170; 1 i1 sin, SMF 1985/188; 6 p3 sin, SMF 1985/144-146, 160, 161, 163; 11 p3 dex, L Length; SMF 1985/143, 147-153, 159, 162, 164; 5 fragments of p3, SMF Ltrig Trigonid length; 1985/154-158; 9 p4/m1 sin, SMF 1985/189-196, 234; 6 p4/m1 dex, PH Hypoflexus width; SMF 1985/215-220; 6 trigonids of p4/m1 sin, SMF 1985/204-206, TH Distal hyperloph length; 209, 223, 235; 4 trigonids of p4/m1 dex, SMF 1985/221, 225, 236, W Width; 237; 3 talonids of p4/m1 sin, SMF 1985/207, 208, 238; 2 talonids of p4/m1 dex, SMF 1985/222, 224; 1 m2 sin, SMF 1985/210; Wtl Third lobe width of m2; 1 m2 dex, SMF 1985/226; 1 trigonid of m2 sin, SMF 1985/211; Wtal Talonid width; 1 trigonid of m2 dex, SMF 1985/228; 3 talonids of m2 sin, SMF Wtrig Trigonid width. 1985/212-214; 4 talonids of m2 dex, SMF 1985/227, 229, 230, 239; 2 ?m2 sin, SMF 1985/232, 233; 2 D3/4 sin, SMF 1985/268, Other abbreviations 269; 2 d3 sin, SMF 1985/256, 257; 5 d3 dex, SMF 1985/258-262; 4 d4 sin, SMF 1985/263-266; 1 d4 dex, SMF 1985/267. Raciszyn Dp-d (L of d3)-(L of p3) [mm]; 1: 1 mandible body dex with complete dentition, ZPAL M.10. Kd/p ((L of d3)/(L of p3))*100 [%]; MdL ((L of alveolar row)/(L of diastema))*100 [%]; MEASUREMENTS. — See Tables 1 and 2, Fejfar et al. (2006), and Čermák & Angelone (2013). MH (((H of mandible at p3+ H of mandible at m2)/2)/ L of alveolar row)*100 [%]; STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION. — Early Pliocene N Number of specimens; (late Ruscinian, MN15b) of central Europe: localities Gundersheim OR Observed range; (probably Gundersheim 1 sensu Tobien 1980; original material of F. Heller), Gundersheim-fissure 4, Gundersheim-Findling, possibly X̄ Arithmetic mean. Wölfersheim (all in Germany) and Raciszyn 1 (Poland). SYSTEMATIC PALAEONTOLOGY DESCRIPTION d3 (Fig. 2A, B) Class MAMMALIA Linnaeus, 1758 Three-lobed tooth; the anteroconid is always isolated (N = 7), Order LAGOMORPHA Brandt, 1855 shaped as an elongated triangle; it may bear an anterior and/or Family PROLAGIDAE Gureev, 1960 posterior undulation in the lingual side; in one case there is a Genus Prolagus Pomel, 1853 connection of the anteroconid and the accessory cusp, marked by a deep anteroflexid. A small, roundish cusp is present in Prolagus bilobus Heller, 1936 the antero-lingual side, between anteroconid and protoflexid, (Figs 2-4) in 86% of cases (N = 7). The trigonid is indented by an average-sized to deep centroflexid; the protoconulid is quite NAME-BEARING TYPE AND TYPE LOCALITY. — Lectotype – left p3 (SMF reduced, as well as the metaconid. A centroisthmus between 1996/136), illustrated in Heller (1936: fig. 3), designed by López trigonid and talonid is present in 40% of cases (N = 5). In COMPTES RENDUS PALEVOL • 2021 • 20 (28) 601 Čermák S. et al. TABLE 1. — Dental measurements of Prolagus bilobus Heller, 1936 from the early the anterior side of the entoconid, which is always quite Pliocene (MN15b) locality Gundersheim-Findling. See Material and methods for abbreviations. thick (length > 1/5 of total L; N = 11) and without enamel hiatus; the mesoflexid is very variable, from relatively short and U-shaped to quite long and J-shaped (N = 12) – in one Lower and upper teeth case, the mesoflexid connects to the centroflexid, isolating N x̄ OR CV the metaconid, and in 2 cases it hosts an additional enamel p3 L 10 2.47 2.20-2.77 7.96 W 9 2.30 2.11-2.57 6.34 islet in its lingual side (such islet in one case connects to the p4/m1 L 13 1.95 1.69-2.13 7.11 entoconid with wear, in the other case it remains isolated); Ltrig 22 1.12 0.90-1.32 9.41 the crochet is present in all adult specimens, predominantly Wtrig 23 2.13 1.88-2.47 8.64 Wtal 19 2.09 1.85-2.39 7.73 with a very large size (in 50%; large in 36%, small in 14%; m2 L 3 2.65 2.45-2.81 – N = 14); the centroflexid has undulated sides (71%, N = Wtrig 6 2.02 1.71-2.19 9.09 14), parallel in ¾ of its length, then widens in correspond- Wtal 7 1.99 1.82-2.25 7.36 ence of the crochet – apart for the sporadical above men- Wtl 7 1.38 1.26-1.54 8.46 d3 L 5 1.37 1.07-1.50 12.84 tioned connection with the mesoflexid, the centroflexid W 5 1.06 0.96-1.14 7.27 may be anteriorly “closed” by a connection of proto- and d4 L 5 1.50 1.29-1.79 12.30 metaconid; the metaisthmus varies from relatively wide (2 Wtrig 5 1.12 1.00-1.21 7.94 Wtal 5 1.16 1.05-1.25 6.88 cases) to very thin (N = 14), when it is thin, it may have a P2 L 5 1.69 1.60-1.82 5.52 bottlenecked appearance; the protoisthmus is always thin to W 5 2.64 2.50-2.87 6.97 very thin (N = 13) and its connection with the protoconid P3 L 10 2.04 1.71-2.21 7.06 W 9 3.45 2.85-3.84 8.92 has a “bottlenecked” appearance only in 23% (N = 13); the AA 9 2.36 1.70-2.76 12.77 hypoconid is triangular in shape and predominantly of aver- P4 L 16 1.65 1.36-1.85 8.73 age size (only in two cases it is quite large); the hypoflexid is TH 17 0.98 0.80-1.17 10.79 W 11 3.23 2.79-3.80 9.98 very deep, almost reaching the posterior side of the tooth; AA 11 2.87 2.42-3.31 11.01 the anterior part of the protoconid is quite protruding lin- PH 12 1.45 1.19-1.69 11.61 gually, and the passage to the thin posterior part is marked M1 L 19 1.66 1.12-1.95 13.64 W 18 3.07 2.30-3.51 12.39 by an abrupt narrowing; the protoconid is connected to the AA 19 2.70 1.94-3.04 11.76 protoconulid by a wide to very wide loph, at the junction PH 19 2.42 1.70-2.90 12.70 protoconid-protoconulid there may be a spur in about half M2 L 13 1.51 1.25-1.80 11.03 W 12 2.84 2.29-3.54 13.32 of the cases; the protoconulid is quite short, often very wide AA 13 2.44 1.94-2.98 13.33 with a bulky appearance. PH 13 2.21 1.80-2.69 12.99 The occlusal features of juvenile teeth (identified mainly by D3/4 L 2 – 1.21, 1.29 – W 2 – 1.75, 1.96 – their conical shaft; N = 6) are the following: the anteroconid, always displaced towards the lingual side, is medium-sized (smaller than metaconid in 67%), rather flattened oval in the talonid, the entoconid is much smaller than the globous shape, distally smooth (in one case lingually with fold); the hypoconid, which appears to be the largest cusp, sometimes metaconid is mainly triangular, in one case is isolated; the even larger than the anteroconid. crochet may be absent (1 case), very reduced or of average size, straight or inclined with respect to the antero-posterior d4 (Fig. 2C, D) axis of the tooth, and its position also varies from central to Trigonid and talonid are always well separated. The trigonid displaced (there is no univocal correspondence between its is of equal size or slightly smaller than the talonid; it bears a size, shape and position); the hypoconid is large compared quite developed, flattened anterior lobe, separated from the to the size of the tooth, triangular or with rounded lingual main body of the trigonid by a simple, straight paraflexid edge; the hypoflexid is deep, as in adult individuals; the (no centroflexid). The triangular talonid lacks or has a very mesoflexid is predominantly V-shaped and short; the cen- reduced anterior isthmus. troflexid shape is similar to that of adults, with undulated sides in 33%; the protoisthmus is thin to very thin, often p3 (Fig. 2E [in part], F-N) bottlenecked (50%); the metaisthmus varies from thin to The occlusal features of adult teeth (identified mainly by their average wide; the protoconid has the same shape as in adults prismatic shaft) are the following: large size; the anteroconid (developed anteriorly, reduced posteriorly), however the is as large as metaconid (92%, N = 12) – in one case, it is anterior part may be more or less developed towards the slightly smaller; the anteroconid is roughly shaped as a right- lingual side; the proportions protoconid-protoconulid are angled triangle and appears “tilted” (i.e., with the 90° angle variable, but in general the protoconulid is usually thin, pointing posteriorly); the main body of the anteroconid is though not very long. always displaced towards the lingual side (N = 13), its posterior and labial sides with undulation or notch in 69% (N = 13); p4-m2 (Fig. 2E [in part], O) the metaconid is quadrangular, with enamel undulated and/ The p4-m1 consist of two separate lobes of approximately or crenulated all around; the crenulation may interest also similar width fused together by cement; posteriorly, both 602 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe A E F G H O B C I J D K L M N FIG. 2. — Morphology of lower teeth of Prolagus bilobus Heller, 1936 from the early Pliocene (MN15b) locality Gundersheim-Findling: A, left d3, occlusal view, SMF 1985/256; B, right d3, occlusal view, reversed, SMF 1985/259; C, left d4, occlusal view, SMF 1985/264; D, right d4, occlusal view, reversed, SMF 1985/267; E, right p3-m2, occlusal view, reversed, SMF 1985/165; F, right p3, occlusal and root views, reversed, SMF 1985/152; G, right p3, occlusal and root views, re- versed, SMF 1985/151; H, right p3, occlusal and root views, reversed, SMF 1985/150; I, left p3, occlusal and root views, SMF 1985/145; J, right p3, occlusal view, reversed, SMF 1985/147; K, right p3, occlusal view, reversed, SMF 1985/159; L, right p3, occlusal view, reversed, SMF 1985/162; M, right p3, occlusal view, reversed, SMF 1985/164; N, left p3, occlusal and root views, SMF 1985/163; O, right p4-m2, occlusal view, reversed, SMF 1985/166. Scale bar: 1 mm. lobes have thickened enamel. The trigonids are of rhomboid of the mandible is convex, and becomes more prominent shape, longer than talonids, with an accentuated anterior and angular with increasing ontogenetic age. The mylohyoid step; the talonid shows an antero-labial flexid, and a shal- line is moderately developed, and in ventral view it appears low notch in the labial edge; the m2 are three-lobed, the located near the buccal edge of the mandibular body. The first two similar to p4-m1, whereas the last lobe is simple, coronoid process is weakly developed. The articular head is triangular-shaped. strongly convex in medio-lateral direction. Lower jaw (Fig. 4) D3-4 (Fig. 3X, Y) Eleven fragments of mandibular bodies in different ontoge- Three roots. The mesial hyperloph becomes longer with wear; netic stages are available. In adult individuals, the mandible in less worn specimens it covers the lagiloph, and then “grows” is rather robust. The diastema is long, but slightly shorter to cover the entire postcone. The mesial hyperloph has a than the p3-m2 alveolar row (Table 2). The posterior mental notch on the posterior edge. The mesial hypercone is much foramen is large, oval, located close to the ventral edge of less developed than the distal one, both in size and width. the mandibular body below m1 (in most cases, below the Consequently, the hypoflexus appears as a very open “V”. Both trigonid/talonid connection; c. 67%, N = 6), and remarkably para- and mesoflexus are very deep, and their relative depth maintains the same position both in juvenile and adult spec- can vary: the paraflexus can be deeper than the mesoflexus imens. The area below p3-p4 is richly fenestrated. The lower or viceversa. The lagicone is small and thin; its connection incisor extends below the p4 (in most cases below the trigo- with the very short lagiloph is bottlenecked. The postcone is nid/talonid connection; c. 50%, N = 6). The ventral margin rounded, with inclined axis and with a developed postlobule. COMPTES RENDUS PALEVOL • 2021 • 20 (28) 603 Čermák S. et al. A B C D E F G H I J K L M N O P Q R S T U V W X Y FIG. 3. — Occlusal morphology of upper teeth of Prolagus bilobus Heller, 1936 from the early Pliocene (MN15b) locality Gundersheim-Findling: A, left P2, SMF 1985/70; B, left P2, SMF 1985/71; C, right P2, reversed, SMF 1985/72; D, right P2, reversed, SMF 1985/73; E, right P2, reversed, SMF 1985/74; F, right P3, re- versed, SMF 1985/86; G, left P3, SMF 1985/77; H, right P3, reversed, SMF 1985/78; I, right P3, reversed, SMF 1985/79; J, left P3, SMF 1985/82; K, left P4, SMF 1985/89; L, left P4, SMF 1985/91; M, left P4, SMF 1985/92; N, right P4, reversed, SMF 1985/99; O, right P4, reversed, SMF 1985/101; P, left M1, SMF 1985/109; Q, left M1, SMF 1985/110; R, left M1, SMF 1985/112; S, left M1, SMF 1985/111; T, right M1, reversed, SMF 1985/113; U, right M2, reversed, SMF 1985/127; V, right M2, reversed, SMF 1985/128; W, left M2, SMF 1985/129; X, left D3/4, SMF 1985/268; Y, left D3/4, SMF 1985/269. Scale bar: 1 mm. P2 (Fig. 3A-E) P4 (Fig. 3K-O) Quite flattened in shape. The mesial hyperloph is long, The mesial hyperloph is more protruding lingually than the covering 67-78% (N = 6) of the tooth width, often robust, distal one, but it is much shorter (TH, i.e., its average length, and always without enamel hiatus. The lagicone is straight is 40% of tooth L). The hypoflexus reaches about 45% of or very slightly inclined lingually, variable in thickness, tooth W (50% of AA) and never merged with parafossette and slightly longer than postcone in 83% (N = 6), in one (N = 21), it is V-shaped in its lingual part, then becomes case with an undulation in its postero-lingual side. Para- narrow U-shaped in 62% (N = 21); the U-shaped part also and mesoflexus are deep and curved, with the latter longer has a different inclination with respect to the other part, i.e., than the former. pointing anteriorly. The parafossette is long to extremely long in its anterior part, slightly inclined towards the anterior side P3 (Fig. 3F-J) of the tooth in most cases, and its width remains constant The mesial hyperloph is of average thickness, slightly widened all along, except for a few cases, in which the anterior end is at the distal end, always without enamel hiatus (N = 10). slightly widened; sometimes the anterior end of the parafos- The mesial hypercone is lingually less protruding than the settes covers and surpasses the mesofossette (18%; N = 22). distal one. The hypoflexus is V-shaped, from very shallow The mesofossette is variable in size from large to very large, to shallow. The para- and mesoflexus are deep, curved, of and in shape from V-shaped (predominant), to U-shaped equal depth or sometimes with the paraflexus as the deepest and C-shaped. of the two. The lagicone-lagiloph complex is very thin in most cases, the lagicone-lagiloph connection is marked by M1 (Fig. 3P-T) a sharp change of the curvature. The lagiloph covers half or The mesial hypercone is more protruding than the distal one. the entire postcone (in any case, never reaching the labial The hypoflexus is very long (c. 79% of W; c. 90% of AA), side of the tooth). The postcone is rounded or quadrangular, curved posteriorly or straight. The fossettes are predominantly connected posterolabially to a postlobule of very variable size. present; in 92% of cases (N = 26) there is one fossette, in 4% 604 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe A B C D E FIG. 4. — Mandibles of Prolagus bilobus Heller, 1936: A, juvenile right mandible with i1, p3-m2 (SMF 1985/165) from Gundersheim-Findling; B, fragment of right mandible with p4-m2 (SMF 1985/166) from Gundersheim-Findling; C, fragment of right mandible with p4-m2 (SMF 1994/895, paralectotype of P. bilobus) from Gundersheim (probably Gundersheim 1 sensu Tobien 1980); D, right mandible with p4-m2 (SMF 1996/156-157) from Gundersheim (unpublished material by F. Heller probably from Gundersheim 1 sensu Tobien 1980); E, right mandible with i1, p3-m2 (ZPAL M.10.) from Raciszyn 1. Scale bar: 10 mm. of cases two fossettes, and in the remaining cases (4%) fos- 5) P. michauxi and P. sorbinii in having a triangular indented settes are absent. The fossettes are positioned just under the anteroconid, vertically aligned with the entoconid. distal end of the hypoflexus; they vary in shape from round The availability of additional materials allowed us to highlight to slightly oval (42%; N = 24) to elongated (58%). some peculiarities of the species also in other tooth positions: – d3 with its isolated labial cusp and predominance of M2 (Fig. 3U-W) separated talonid also in advanced stages of wear. This fea- The precone is well-developed, the postlobule reduced. The ture, which recalls P. crusafonti (see Angelone 2007: table 2, mesial hypercone is slightly more protruding than the distal p. 413), can be considered as quite primitive. In fact, latest one. As with M1, the hypoflexus is very long (c. 78% of W; Miocene and Pliocene species (as far as we know, taking into c. 91% of AA) and curved posteriorly or straight. The fossettes account the scarce availability of fossil lagomorphs deciduous are predominantly absent (93%; N = 15); when present, they teeth and of detailed studies on the subject) are characterized are round and rather small. by a connected talonid. – d4 with a very deep paraflexid but not a centroflexid, as TAXONOMIC COMPARISON AND REMARKS in late Miocene congeneric populations of western Europe. Prolagus bilobus has recently been partly revised by Čermák & – P2 with generally flattened appearance and very long Angelone (2013). The species is clearly differentiated by its mesial hyperloph. A flattened P2 is a common and stable very large p3 (Fig. 5) with lingually shifted, large, triangular, feature of P. bilobus, but it is instead quite variable in other posteriorly, and buccally indented anteroconid; undulated Pliocene species. The mesial hyperloph is remarkably longer sides of centroflexid; long crochet; metaconid of similar size as compared to the Pliocene species P. pannonicus, P. latiunci- anteroconid; thick entoconid lacking enamel hiatus. The p3 of natus, and P. ibericus. P. bilobus has a very homogeneous and stable morphology, as – P3 with centrocone not reaching the labial edge and far as the anteroconid, crochet and entoconid are considered: P4 with long, upturned parafossette. Both characters are in all the adult individuals have a lingually displaced anteroconid common with the sorbinii group, providing a hint about the with a unique “tilted” appearance, a voluminous crochet, and possible phylogenetic affinity of P. bilobus. a thick, crenulated entoconid. These features seem unaffected – P4 with mesial hyperloph remarkably thin. by the intraspecific variability range. The p3 differs from that The very thin dentine bridge between parafossette and in: 1) P. pannonicus, P. sorbinii, P. latiuncinatus, P. caucasicus, hypoflexus in P4 of P. bilobus can be well observed in P. pan- P. michauxi, P. ibericus, and P. savagei in having a significantly nonicus and P. latiuncinatus; in Prolagus sp. from Beremend larger size (Fig. 5); 2) P. pannonicus and P. caucasicus in having 39 these structures may even be confluent. a larger and wider p3 anteroconid; 3) P. sorbinii, P. ibericus, Tesakov & Averianov (2002) attributed to P. bilobus scanty P. depereti, P. savagei, and P. italicus in having a larger crochet; materials of isolated p3s from Tanatary (MD, MN15; 1 4) P. caucasicus in having a quadrangular metaconid; and p3, formerly attributed to P. cf. oeningensis by Erbajeva & COMPTES RENDUS PALEVOL • 2021 • 20 (28) 605 Čermák S. et al. Prolagus bilobus Gundersheim (probably 1) - DE (?MN15b), Lt L Ht Lt Gundersheim-Findling - DE (MN15b) Raciszyn 1 - PL (MN15b) 2.5 W Ht Prolagus sp. Beremend 26 - HU (MN15b) Ht Prolagus italicus (Montagnola Senese - IT, MN17) L [mm] Ht Ht Prolagus savagei (Arondelli - IT, MN16a) 2.0 Prolagus depereti (Perpignan - FR, MN15) Ht Prolagus ibericus (Layna - ES, MN15) Ht Prolagus michauxi (Sète - FR, MN15a) Prolagus caucasicus (Kosyakino - RU, MN14) Ht Prolagus sorbinii (Monte Castellaro - IT, MN13) 1.5 Ht - holotype Prolagus latiuncinatus (Polgárdi - HU, MN13) Lt - lectotype Prolagus pannonicus (Sümeg - HU, MN10/11) 1.5 2.0 2.5 W [mm] FIG. 5. — Comparison of p3 size of Prolagus bilobus Heller, 1936 and Prolagus sp. from Beremend 26 with materials from type localities (type materials or addi- tional materials) of selected late Miocene-Pleistocene Prolagus species of Europe. TABLE 2. — Mandibular measurements of Prolagus bilobus Heller, 1936: 1, Gun- dersheim-Findling, SMF 1985/165; 2, Gundersheim (probably Gundersheim 1 For the sake of completeness, it is worth to mention that sensu Tobien 1980), SMF 1996/156, unpublished material of F. Heller; 3, Rac- Tesakov & Averianov (2002) tentatively referred to P. bilobus iszyn 1, ZPAL M.10. the population from Polgárdi (HU, MN13), a hypothesis refuted by Angelone & Čermák (2015) who assigned the Measurements 1 2 3 sample to the new species P. latiuncinatus. Lingual height of mandible at p3 4.89 7.13 9.09 To conclude, at present we include only the localities of Lingual height of mandible at p4 6.50 9.01 12.98 Gunderhseim and Raciszyn 1 in the proven P. bilobus distri- Lingual height of mandible at m2 6.23 8.56 12.25 bution, as the remains from Wölfersheim and Kisláng are at Length of diastema 5.75 – 11.14 Alveolar length of p3-m3 7.95 9.39 12.04 present not available (see above for details). Mandible width at p4 3.31 4.48 5.82 Mandible width at m2 2.53 3.56 4.29 Prolagus sp. Shushpanov (1988) and Erbajeva (1988)), Tatareshty (MD, (Figs 6; 7) MN15; 1 p3), Kotlovina (UA, probably MN15 or MN16 according to Nesin & Nadachowski (2001); 1 p3, formerly MATERIAL. — Beremend 26: 1 incomplete lower jaw sin with p3- m2, IG Lag/Be26/01. Beremend 39: 2 incomplete upper jaws sin attributed to P. cf. oeningensis by Agadjanian & Erbajeva (1983)), with P3-M2 and P4-M2, IG Lag/Be39/02, IG Lag/Be39/03; 1 m1 and Kamenskoe (UA, age unclear, probably MN15; 1 p3, dex, IG Lag/Be39/04. formerly attributed to Prolagus sp. by Topachevsky (1962)). These p3 are characterized by: 1) a small, not tilted anteroconid MEASUREMENTS. — See Table 3. shifted towards the lingual side of the tooth; 2) a reduced, STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION. — Early Pliocene diamond-shaped metaconid with unfolded enamel, and 3) (late Ruscinian, MN15b); Beremend 26 and 39. a reduced entoconid. Thus, they are quite different from P. bilobus. In the specimens from Tatareshty and Kotlovina, the DESCRIPTION crochet is absent or quite reduced, contrarily to adult specimes p3 (Fig. 6E [in part]) of P. bilobus. The measurements, available only for Tatareshty Medium-size. The anteroconid is large, triangular in shape, (L × W = 1.95 × 1.93; Tesakov & Averianov 2002) and Tanatary not tilted (i.e., with posterior side perpendicular to the (L × W = 1.80 × 1.70; Erbajeva & Shushpanov 1988), indicate antero-posterior axis of the tooth), not perfectly central, that the Moldavian specimens are sensibly smaller in size than but slightly displaced towards the lingual side of the tooth. P. bilobus. The overall morphology of the p3 of Ukrainian and There are no indentations, undulations, nor convexities on Moldavian materials described above reminds the appearance the labial and posterior side of the anteroconid. The meta- of some populations of P. sorbinii (see Angelone 2007), a conid is as large as the anteroconid, hatchet-shaped, with species distributed in Greece and Italy in the late Miocene undulated labial and posterior sides. The metaisthmus is and earliest Pliocene (Angelone & Rook 2012). Thus, those narrow, bottleneck-shaped. The entoconid is quite thick remains are tentatively attributed here to P. aff. sorbinii. (length > 1/5 L), without enamel hiatus. The protoisthmus 606 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe C B D A E FIG. 6. — Mandible of Prolagus sp. from the early Pliocene (MN15b) locality Beremend 26: A-E, left mandible (IG Lag/Be26/01) with p3-m2 in (A) ventral, (C) lingual, and (D) buccal views; B, rostral view of the mandibular ramus; E, occlusal morphology of p3-m2. Scale bars: A-D, 10 mm; E, 1 mm. is very narrow, bottleneck-shaped. The crochet is extremely lower incisor extends below the trigonid of p4. The ventral feeble, in central position, symmetrical in shape. The cen- margin of the mandible is convex. The mylohyoid line is long, troflexid is straight, parallel to the antero-posterior axis of well-developed, and in ventral view appears located near the the tooth, with undulated sides, and enlarged at its distal lingual edge of the mandibular body. The coronoid process is end, in correspondence of the crochet. The hypoconid is moderately developed. The articular head is strongly convex small, triangular in shape, with flattened labial end. The in medio-lateral direction. protoconid is wide in its anterior part, narrowing towards the posterior part of the tooth in its distal part. The con- P3 (Fig. 7E [in part]) nection protoconid-protoconulid is marked by a spur. The The mesial hyperloph is thin, of constant width, without enamel protoconulid is relatively small, of average width. hiatus, and covers part of the postcone. The mesial hypercone is less protruding than the distal one. The distal hyperloph, p4-m2 (Fig. 6E [in part]) however, is very reduced in length. The V-shaped hypoflexus In p4-m1 trigonid and talonid have similar width and both is short. Para- and mesoflexus have the same depth, and reach show a thickening of the enamel on the posterior side. The posteriorly almost the edge of the tooth. The lagicone is of trigonids are of rhomboid shape, longer than the talonids. average size and rounded, the lagiloph has no indentations In p4 the trigonid has an anterior step. The talonid shows an or steps, and the centrocone does not reach the labial end. antero-labial flexid, more marked in p4, less marked in m1 and even less in m2. The three-lobed m2 have trigonid and P4 (Fig. 7D [in part], E [in part]) talonid similar to other molariform teeth, plus a triangular- The mesial hyperloph is slightly longer than the distal one; shaped posterior lobe. the length of mesial- and distal hyperloph is similar. The hypoflexus is V-shaped in its lingual part, then takes a very Lower jaw (Fig. 6A-D) narrow U-shape; in one individual the hypoflexus is con- The outline is slender in both lateral and ventral views; the nected to the parafossette, in the other, only a very thin diastema is long, and its length is almost equal to the alveolar enamel bridge separates the hypoflexus and the parafossette. one. The posterior mental foramen is large, oval, located close The parafossette is very long, thin, and its anterior part may the ventral edge of mandibular body in the area below m1/ be upturned anteriorly; the mesofossette is C- or V-shaped, m2. Several accessory foramina are present below p3-p4. The with the anterior tip slightly longer than the posterior one. COMPTES RENDUS PALEVOL • 2021 • 20 (28) 607 Čermák S. et al. TABLE 3. — Cranial and dental measurements of Prolagus sp. from the early differences can be observed with respect to some late Miocene Pliocene (MN15b) localities Beremend 26 and 39. See Material and methods for abbreviations. congeneric species (P. sorbinii / P. latiuncinatus), namely in its very large triangular anteroconid of p3. Prolagus sp. from Beremend differs from the Pliocene-Holocene species Mandible and maxillae Lower teeth Upper teeth P. caucasicus, P. michauxi, P. ibericus, P. depereti, P. figaro, Total height of mandible 21.37 P3 L 1.89 P. bilobus, P. savagei, P. italicus, and P. sardus, as its anteroconid Lingual height 6.46 p3 L 2.04 W 3.30 of mandible at p3 is not displaced towards the lingual side. Moreover, the Lingual height of 8.18 W 1.91 AA 2.45 p3 of Prolagus sp. from Beremend 26 differs from that in: mandible at p4 1) P. pannonicus, P. latiuncinatus, P. caucasicus, and P. ibericus Lingual height of 7.92 p4 L 1.76 P4 L 1.50, in having a larger size; (Fig. 5); 2) P. depereti, P. savagei, and mandible at m2 1.59 Length of diastema 8.18 Ltrig 1.05 TH 0.83, P. bilobus in having a smaller size (Fig. 5); 3) P. pannonicus and 0.84 P. caucasicus in having a larger and wider p3 anteroconid; and Coronar length of p4-m2 6.08 Wtrig 1.93 W 3.13, 4) P. pannonicus, P. latiuncinatus, and P. bilobus in having a 3.18 Alveolar length of p3-m2 9.16 Wtal 1.96 AA 2.68, significantly less developed crochet. 2.68 Surprisingly, Prolagus sp. from Beremend 26 is morphologi- Mandible width at m2 3.20 m1 L1.79, PH 1.15, cally and dimensionally incompatible with the coeval P. bilobus. 1.91 1.33 The p3 of Prolagus bilobus has an extremely stable morphol- Mandible width at p4 4.34 Ltrig 1.05, M1 L 1.49, 1.15 1.58 ogy (see section “Taxonomic comparison and remarks” about Width of articular head 3.95 Wtrig 2.01, W 2.99, of P. bilobus). This characteristic makes impossible to “fit” in 2.01 3.08 P. bilobus the lower jaw from Beremend 26, which p3 has a Wtal 1.98, AA 2.65, central, not tilted anteroconid, an extremely small crochet, 2.01 2.68 m2 L 2.33 PH 2.35, and a medium-sized entoconid. Moreover, the lower jaw of 2.36 Prolagus sp. from Beremend 26 has a different outline with Wtrig 1.92 M2 L 1.43, respect to P. bilobus, especially in the ventral part (which is 1.50 Alveolar length of P2-M2 9.54, Wtal 1.83 W 2.85 regularly convex and not angular), and in the size and posi- 9.68 tion of the posterior foramen (smaller and positioned further Length of P2 alveolus 1.95 Wtl 1.31 AA 2.54 posteriorly). Also the measurements of the p3 and of the lower Width of P2 alveolus 2.86 PH 2.16, jaw of Prolagus sp. from Beremend 26 are incompatible with 2.33 an attribution to P. bilobus. In fact, though pertaining to an old individual, the measurements of p3 and lower jaw of Pro- M1 (Fig. 7D [in part], E [in part]) lagus sp. from Beremend 26 fall below the distribution of adult The mesial hyperloph is rather thin, longer than the distal P. bilobus (Fig. 5; Table 3). However, the upper teeth pertaining one. The hypoflexus is rather long (c. 78% of W, c. 88% to a jaw excavated from Beremend 39, another karst filling of of AA; N = 2), posteriorly curved; in 1 of 2 individuals, a the Beremend karst complex, which apparently is coeval to round, small fossette is present posteriorly to the labial tip Beremend 26, at least judging from the accompanying fauna of the hypoflexus. (Pongrácz in litt.; Čermák & Wagner 2013), do not show substantial differences in morphology and measures compared M2 (Fig. 7D [in part], E [in part]) to P. bilobus (cf. Tables 1; 3). This taxonomic incongruence The general features are the same as M1, with a relatively between materials from coeval and neighbouring fissure fillings longer hypoflexus (c. 82% of W, c. 92% of AA; N = 1) and does not find an explanation for the moment and, though in no fossette. our opinion Prolagus from Beremend 26 represents a distinct species, we provisionally keep all the available material from Upper jaw (Fig. 7A-C) Beremend in open nomenclature as Prolagus sp. The hard palate is of average width. The posterior edge of the It is really tempting to suppose that the p3 and only Prolagus incisive foramen reaches the posterior edge of P2. The ante- specimen from Kisláng (Kretzoi 1954) may be related to Prola- rior edge of the choanae reaches the half of P4. The premolar gus sp. from Beremend 26. Indeed, the geographical proximity foramen is large, oval, positioned medially to P4. Next to of the fossil sites, and the age range of the mixed assemblage of the maxillopalatine suture, there is a large palatine foramen, Kisláng (MN15-MQ1; Mayhew 2012) are compatible with posteriorly accompanied by several smaller foramina. The this hypothesis. However, the morpho-dimensional features masseteric spine is short and triangular in shape. The infraor- of the p3 remain from Kisláng described in literature are too bital foramen is oval, prolonged only slightly in dorsoventral unclear to make any taxonomic speculation. Kretzoi (1954: 247) direction. The infraorbital canal is short; in relation to the provided only indications about the presence of a crochet, and wall of P2 alveolus, placed rather dorsally. hypothesized a taxonomic proximity of the Kisláng specimen to Prolagus from Gundersheim. This is in contrast with our TAXONOMIC COMPARISON AND REMARKS observation about the lower series from Beremend 26 (totally The sole Prolagus specimen from Beremend 26, represented by different from Gundersheim). We envisage the retrieval of the an almost complete mandible, is extremely peculiar. Evident Prolagus materials from Kisláng, in order to solve this open issue. 608 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe A B C D E FIG. 7. — Maxillae of Prolagus sp. from the early Pliocene (MN15b) locality Beremend 39: A-D, left maxilla (IG Lag/Be39/03) with P4-M2 in (A) ventral, (B) buccal, and (C) rostral views; D, occlusal morphology of P4-M2; E, occlusal morphology of P3-M2 in left maxilla (IG Lag/Be39/02). Scale bars: A-C, 5 mm ; D, E, 1 mm. RESULTS AND DISCUSSION Teeth In the genus Prolagus, the morphology of juvenile permanent NOTES ON THE ONTOGENY OF P. BILOBUS teeth is quite different from the adults of the same species. Mandibles This does not just depend by the fact that the cusps may be The available jaws of P. bilobus cover various ontogenetic stages unworn: in fact, especially in p3, the relative size of the cusps, (Fig. 4) and show a noticeable size range (Table 2), typical of their connection, their position, and also their presence, are lagomorphs. Several allometric changes distinguish juvenile prone to noticeable variations during the ontogeny. Due to and adult individuals (see Fig. 4): the progressive inclination of the wear surface (W) and the – the diastema is relatively shorter (compared to the alveolar conical shape of the juvenile tooth shaft (L and W), also the row length) in the juvenile mandible (SMF1985/165) than size ranges of the teeth show a wide variation if juveniles are in the unbroken aged specimen (ZPAL M.10). The values of included in the measurements. It has even been observed, MdL are 138% vs 108%, respectively. since the early studies about this taxon, that in some tooth – the mandibular body is relatively lower (compared to positions the measurements cluster in two very distinct clouds, alveolar row length) in the juvenile mandible (SMF1985/165) one for permanent juvenile teeth and one for adults (López in contrast to that in the adult (SMF 1996/156) and aged Martínez 1974). (ZPAL M.10) ones. The values of MH are 70 vs 84 and 89%, Also in the case of P. bilobus, it is possible to notice the respectively (to exclude the influence of ventral curvature of remarkable ontogenetic changes in p3. Moreover, in this the mandibular body, the height was calculated as the mean case, thanks to the stable occlusal surface morphology which value of measurements at p3 and m2). characterizes this species (see “Taxonomic comparison and – a relative height increase of the mandibular body is asso- remarks” of P. bilobus) and thanks to a sufficient number of ciated with the curvature increase of its ventral margin; this specimens for both ontogenetic categories (6 conical p3 in the phenomenon was also observed in the well-known and docu- initial stage of abrasion = permanent juvenile teeth; and 14 mented species Prolagus sardus (Tobien 1935: Fig. 2, p. 272). p3 with prismatic shafts = permanent adult teeth), we were The above described ontogenetic variations observed in the able to observe in the occlusal and “radical” parts of the teeth jaws of P. bilobus are quite standard for the genus. In Prolagus, the following pattern (young to old): though, also the position of the jaw posterior foramen slightly – the enlargement, deformation, and tilting of the antero- varies during the ontogeny, but in P. bilobus this feature remains conid; unchanged in all the available specimens. – the deformation of the metaconid; COMPTES RENDUS PALEVOL • 2021 • 20 (28) 609 Čermák S. et al. – the noticeable thickening of the protoconulid; MN13 (Angelone 2007; Angelone & Čermák 2015 with – the thickening of the entoconid; references). The only known exception is the MN15 P. ibericus, – the appearance or the enlargement of the crochet. an extremely peculiar species whose phylogenetic history is The ontogenetic changes observed in P. bilobus can not quite mysterious and characterized by several “incongruent” represent a pattern perfectly applicable to other congeneric features compared to coeval ones (first of all the very small species. However, it would be worth to verify if the patterns size), known only from the type locality Layna (NE Spain; of ontogenetic trajectories are similar in phylogenetically López Martínez 1989). It must be remarked, though, that in close species. spite of all its peculiarities, the Kd/p of P. ibericus is congruent It is very interesting to notice the extreme ontogenetic dimen- with that of coeval species. sional change between the deciduous and the permanent teeth of P. bilobus (Fig. 8). Indeed, López Martínez (1974) noticed EVOLUTIONARY TRENDS IN PROLAGUS P3: that the relative proportion of the length of d3 and p3 varies CENTRAL VS WESTERN EUROPEAN PLIOCENE SPECIES through time in the species of the genus Prolagus, but this Size observation had no further development in literature. Thirteen The dental size (p3) of European prolagids shows a general selected fossil localities ranging in age from the early Miocene increase trough time, with a few countertrending episodes (MN2b) to the Late Pleistocene with sufficient materials of (e.g. P. oeningensis P. crusafonti; López Martínez 1989) d3/p3 were selected to verify and further investigate López and an interesting sudden enlargement in correspondence Martínez’s (1974) remark. Materials from different ages, but of the MN13 (Angelone 2007). Until Miocene, though, the also from different geographical areas were selected, in order to size range of the p3 of coeval Prolagus species is comparable exclude possible biases. It appears (Fig. 8) that in early Miocene (with punctual exception as P. major in the MN6 of Spain; to early middle Miocene species, Ld3 is larger than Lp3 (mean López Martínez 1989). During the Pliocene, instead, Pro- value of the Kd/p: 110-123%), as occurs in Prolagus’ ancestor lagus shows a wide range of size (Fig. 5): from quite small Piezodus Viret, 1929 (see López Martínez 1974). Since the late (e.g. P. ibericus; Layna, MN15) to very large (e.g. P. bilobus; middle Miocene, the size ratio is reversed, i.e., Lp3 is larger Gundersheim-Findling, Raciszyn 1, both MN15b, which is than Ld3. Moreover, it is remarkable that after the reversal of the largest continental species of the genus). size proportions, the size of d3 not only remains small, but also relatively stable. The size of p3, on the contrary, though Enamel folding and entoconid thickness showing a general increase, is variable, consequently the size Morphological patterns corresponding to temporal trends were differences between d3 and p3 are also quite variable. The mean individuated in the p3 of western European Plio-Pleistocene value of Kd/p ranges from 98% in P. tobieni from the MN7/8 species, namely the simplification of the enamel folding pat- of Escobosa to 55% in P. bilobus from the MN15b of Gun- tern (particularly the loss of the crochet and the smoothing dersheim-Findling, which is the species with the largest d3/p3 of anteroconid and metaconid folds) and the reduction of size difference among the studied taxa (Fig. 8). The percentage the entoconid (Angelone 2008a). of size difference between d3 and p3 in the analyzed taxa does The detailed study of P. bilobus and Prolagus sp. from not evidence a gradual trend depending on the geologic age, Beremend 26 allowed to verify if those trends are shared as hypothesized by López Martínez (1974: 142). Of particular also by Pliocene Prolagus of central Europe. The answer is interest in this sense, is to observe that two distinct species, negative. Prolagus bilobus and Prolagus sp. from Beremend P. tobieni and P. oeningensis, recorded from the same locality 26 do not follow the above listed morphological trends, (Escobosa, Spain), show a quite different Kd/p. It can not be possessing a folded enamel and a thicker entoconid (this excluded that, within a mosaic evolutionary pattern, which is feature is particularly evident in the former species). In this commonly observed in Prolagus, the size differences between sense, they are similar to P. italicus, whose peculiarities were d3 and p3 may carry a signal of phylogenetic closeness of spe- explained by Angelone (2008a) as a consequence of its status cies within distinct clades. At any rate, this parameter may be of peninsular Italian endemic species. Hypothesizing that the useful for species discrimination (cf. Álvarez-Sierra et al. 1990: morphological analogies between P. italicus, P. bilobus, and 16), and it may have a biochronological significance, being Prolagus sp. from Beremend 26 are common to continental a real turning point in the evolution of Prolagus similarly to: isolated species, however, is quite simplistic in our view. In – the appearance of the centroflexid. The presence of a fact, the conditions of isolation (geographical extent, dura- long centroflexid became a stable feature of the genus after tion, dynamics, and climate) of these species are too different the P. vasconiensis/oeningensis transition, near the early/mid- to justify a shared common morphological trend in the teeth dle Miocene boundary. The only exception is a conservative occlusal surface. Possibly, some of the similarities shared by clade endemic to Spain, whose last representant was P. tobieni P. bilobus, P. italicus, and Prolagus sp. from Beremend 26 (still possessing a very incipient centroflexid), a species which and, consequently, their divergence from Pliocene western survived until the early late Miocene (according to Hordijk & European species, may derive from a common origin from van der Meulen 2010). an eastern European stock. In fact, P. italicus, though known – the appearance of the mesial hyperloph in P2 coupled only from the MN17 of central Italy, stemmed from the with the sudden increase of Lp3. These features make a pan- eastern European species P. sorbinii just after its dispersal European, “simultaneous” appearance between MN12 and into Italy, an event occurred in the Messinian (see cladistics 610 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe Monte Tuttavista Navarrete San Juan La Galocha 5 Sandelzhausen Escobosa Escobosa Richardhof Wald Sümeg Brisighella Sète Gundersheim-F. VII-2 XI-r MN2b MN2b MN3a MN5 MN7/8 MN7/8 early MN10 MN10/11 MN13 MN15a MN15b Middle-Late Pleistocene 3.0 P. pannonicus P. aff oeningensis P. aff. crusafonti P. fortis P. sardus P. sardus P. tobieni P. aff fortis P. vasconiensis P. oeningensis P. michauxi P. bilobus P. sorbinii 10 2.5 48 10 2 10 8 L [mm] 6 2.0 6 18 103 1 17 8 N=1 20 20 18 16 2 6 5 1 2 5 1.5 2 16 L L VS 1.0 Kd/p = 112 119 123 110 0 98 88 77 88 76 78 55 81 63 FIG. 8. — Comparison of d3 and p3 lengths (indicated respectively by lighter blue bars and points and darker blue bars and points) in selected early Miocene to Late Pleistocene Prolagus species of Europe. Red/green bars show negative/positive values of Dp-d. Values above or below bars are relative to Kd/p values. See Material and methods for abbreviations. analysis in Angelone et al. 2015; palaeobiogeographical and be isolated endemic populations, relicts left behind after the taxonomic details in Angelone et al. 2019, 2020). There is no reduction of the formerly homogenous Miocene distribution available analysis at present to support the origin of P. bilobus of the genus, a phenomenon that became quite evident since and Prolagus sp. from Beremend 26 from eastern European the Pliocene (López Martínez 2001). Nevertheless, in our species, though it can be a palaeobiogeographically sensible view, there may be an alternative explanation for the Prolagus hypothesis. A definitive answer can be obtained only after occurrences in the Pliocene of central Europe. the revision of Pliocene Prolagus species from eastern Europe and Anatolia. Palaeoclimatical background The genus Prolagus appeared during the early Miocene Entoconid enamel hiatus (MN2b), i.e., towards the end of a period of stable climatic As stated above, an eastern origin for P. bilobus and Prolagus sp. conditions (late Oligocene-early Miocene, MP28-MN3, c. from Beremend 26 can only be hypothesized, but an encour- 27-17 Ma; Maridet et al. 2007). In this period, Europe was aging hint in that sense (waiting for a phylogenetic analysis) characterized by a homogeneous mammal fauna with high can be the presence of an enamel hiatus in the entoconid of interregional taxonomic affinities (sensu Raup & Crick 1979). p3. Angelone (2008a) hypothesized a delayed acquisition of The situation became increasingly heterogeneous since the the enamel hiatus in the Italian P. sorbinii-italicus stock (in end of the early Miocene (MN4, c. 17-16.5 Ma). At a larger P. sorbinii the hiatus is absent, whereas in P. italicus only a scale, we assist at a general global cooling trend started about partial hiatus may be present) in contrast to “the European 14 Ma (Zachos et al. 2001). In Europe, an abrupt end of the lineage” i.e., western European species, which indeed repre- warm period has been recorded at 14-13.5 Ma (Böhme 2003), sent a separate clade; see Angelone et al. 2015), and in which and since the middle Miocene, several cooling and warming the presence of the enamel hiatus can be observed since the pulses, as well as several fluctuations in the precipitation rate latest Miocene (MN13, Messinian). However, other studies occurred (Böhme et al. 2008 with references). In particular, (Čermák & Angelone 2013; Angelone & Čermák 2015; this the transition to a drier climate and a concomitant cooling study) documented the absence of entoconid enamel hiatus episode (c. 9.7-9.5 Ma) determined the biotic crisis known also in the central European taxa P. latiuncinatus (Polgárdi 2, as the Vallesian crisis (but see Casanovas-Vilar et al. 2014). MN13), P. bilobus (Gundersheim-Findling, Raciszyn 1, both The climatic trend induced a strong latitudinal differentia- dated to MN15b), and Prolagus sp. (Beremend 26, MN15b). tion in the European climates. In the precipitation regime, a This evidence reinforces the hypothesis of a south-eastern quite evident latitudinal differentiation between northern and European phylogenetic affinity of P. bilobus and Prolagus sp. southern Europe occurred at c. 8 Ma (Böhme et al. 2008). from Beremend 26. The study of European faunal assemblages evidences the strong isolation of some regions during the late Miocene. EXPLAINING THE CENTRAL EUROPEAN Since the latest Miocene and until the first part of the early DISTRIBUTION OF PROLAGUS DURING THE PLIOCENE Pliocene (MN13-MN14, c. 6.5-5 Ma), a renewed faunal The Pliocene remains of Prolagus recorded at the northern homogeneity shortly occurred in the peri-Mediterranean border of its geographical range were generally considered to area of Europe (Maridet et al. 2007). COMPTES RENDUS PALEVOL • 2021 • 20 (28) 611 Čermák S. et al. The early Pliocene is considered to be a period of globally continental and arid climate (ecogeographical group 2 sensu warm climate (De Schepper et al. 2014 with references), though Popov 2004), inhabiting edge habitats such as dry open for- interrupted by short-lived glacial events (c. 4.9/4.8 Ma, and ests, bushes, dry meadows, etc. In the Mediterranean area, the c. 4 Ma). Since the early/late Pliocene transition (3.6 Ma), genera of group 1 are absent or are very poorly represented, the relatively stable humid conditions which characterized whereas those of group 2 are predominant, together with Europe during the early Pliocene came to an end. The increase genera characteristic of dry open habitats (mostly dry savanna in coolness and dryness culminated with a global glaciation dwellers ecogeographical group 3 sensu Popov 2004). See at c. 3.3 Ma (MIS M2; see De Schepper et al. 2009 and Tan Kretzoi (1956, 1962), Sulimski (1959, 1962, 1964), Kowal- et al. 2017, both with references). The glaciation, though ski (1960), Repenning (1967), Michaux (1971), Jánossy (1972, intense (comparable to early Quaternary glaciations) was 1986), Van de Weerd & Daams (1978), Van de Weerd (1979), short-lived (50 ky) and followed by the mid-Piacenzian Warm Skoczen (1980), Van de Weerd et al. (1982), Reumer (1984, Period (c. 3.3-3.0 Ma) (Tan et al. 2017). Though characterized 1989), Van der Meulen & Van Kolfschoten (1986), Fejfar & by a climate deterioration if compared to the early Pliocene, Storch (1990), Terzea (1997), Şen (1998), Şen et al. (1998), the interval 3.6- c. 3.0 Ma was “the last sustained interval in Ünay & de Bruijn (1998), Popov (2001, 2003, 2004), Fejfar Earth’s history when global climate was warmer than today” et al. (2006) for details. (De Schepper et al. 2009). Between c. 3.0-2.7 Ma is recorded The lagomorph diversity in the late Ruscinian localities the onset of the Northern Hemisphere glaciation. Such event, (MN15b, i.e., coeval with the Prolagus samples under study) contrarily to the quite ephemeral late Eocene to early Plio- of central and south-eastern Europe is quite high (see Kret- cene glaciations (Lunt et al. 2008), is the forerunner of the zoi 1962; Sych 1965; Daxner & Fejfar 1967; Fladerer & glacial-interglacial cycles of the Quaternary (Ganopolski & Reiner 1996; Terzea 1997; Dahlmann 2001; Popov 2004; Calov 2011). Čermák 2007, 2009; Čermák & Angelone 2013; Čermák & The Pliocene environmental changes impacted on mam- Wagner 2013). Five species of lagomorphs pertaining to five malian evolution and dynamics, driving extensive rearrange- distinct genera (Ochotonoma Şen, 1998, Prolagus, Hypolagus ments of community and faunal structures, including large Dice, 1917, Pliopentalagus Gureev & Konkova in Gureev, scale migrations and rearrangements of the selection pres- 1964, Trischizolagus Radulesco & Samson, 1967) are recorded. sure upon the phenotypic design of particular taxa. A short The palaeobiogeographical trajectory and the biochronologi- period near the early/late Pliocene boundary c. MN15-16) cal importance as MN15b markers of some of them is well represents one of the most dramatic global turning points of documented (Angelone 2008b; Čermák & Wagner 2013). The the late Cenozoic faunal history. Nearly a half of the Neogene most common lagomorph was H. petenyii, which occurred mammalian taxa disappeared at that time, and almost all the in almost all central and south-eastern MN15b European elements constituting the modern Quaternary communities localities. This species is usually regarded as a forest dweller appeared in that stage as well (Bernor et al. 1996; Rössner & (Fladerer & Reiner 1996). Trischizolagus dumitrescuae Rad- Heissig 1999; Reumer & Wessels 2003). The aridisation and ulesco & Samson, 1967 and O. csarnotana generally occur spread of open ground habitats grew particularly pronounced together in the Pliocene of Europe. Their presence indicates at the MN16/17 boundary (c. 2.5 Ma), as attested by the a rather dry and hot climate, and the presence of vast areas of loess deposits recognized throughout the northern hemi- savanna-steppes (Popov 2004). Most of the available remains sphere (Rabeder 1981; Rabeder & Verginis 1987; Tedford of T. dumitrescuae are known from the northern part of the et al. 1991; Shi 1994; Xue et al. 2006). peri-Paratethyan area, in fossil sites of early Ruscinian age (MN14). Western occurrences of T. dumitrescuae are rather A focus on Pliocene palaeoecology of central Europe rare and exclusively limited to the late Ruscinian (MN15b). with emphasis on the distribution of lagomorphs Ochotonoma csarnotana from Beremend 26 (Čermák & Wag- In the Pliocene of Europe, the climatic changes resulted in a ner 2013) so far represents the westernmost proven record decoupling of the climatic regime of northern and southern of the species. European occurrences of O. csarnotana are Europe. Relatively warmer and arid conditions characterized limited to south-eastern regions and are closely related to the the Mediterranean area, in contrast to the onset of cooler and distribution of Ochotonoma in Anatolia (see Čermák 2007 humid conditions in the northern parts of central Europe for details). The distribution range of T. dumitrescuae and (Popov 2004). The plant record indicates that, in the early O. csarnotana in central Europe does not exceed latitude Pliocene (before c. 4 Ma), mixed mesophytic forests and N46° (occurences in Beremend 26/39 and Csarnóta 2). On broad-leaved deciduous forest prevailed in the studied area the contrary, the distribution range of Pliopentalagus dietrichi (Kovar-Eder et al. 2008). Indeed, Pliocene faunas of central clearly exceeds this latitude and reaches the Rhine Graben Europe (documented particularly from the localities Gun- (western Germany), together with P. bilobus (co-occurrences dersheim-Findling, Węże 1, and Csarnóta 2) contain a large in Gundersheim-Findling and probably in Wölfersheim). proportion of mesophilous lypotyphlan and rodent species Pliopentalagus dietrichi may be considered, by analogy with (>70%; ecogeographical group 1 sensu Popov 2004). This is its extant relative Pentalagus furnessi (Stone, 1900), a forest indicative for an even and humid climate. The other 30% of dweller inhabiting dense subtropical forests characterized by a the assemblages is represented by species with relatively wide low cursorial ability. In this framework, it is difficult to guess tolerances, adapted to a wide range of biotopes under more the environmental requirements of P. bilobus and Prolagus sp. 612 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe from Beremend. Possibly the latter was more related to a dry in the Pliocene of central Europe except for MN15b: c. 30 and open habitat, given the ecological requirements of the vertebrate-bearing fossiliferous localities are known between other lagomorphs found in the same assemblage, but the MN14-16 (Wagner et al. 2009) and the occurrences of Prolagus former may have pertained to either ecogeographical group are exclusively limited to the 6 localities (out of a total of the 1 or 2 (sensu Popov 2004). 11 available for MN15b). Such chronologically limited occur- It is evident that in MN15b a conspicuous number of the rences are indicative of a dispersal, rather than of a progressive lagomorph genera reached central Europe, most of them Pliocene shrinking of the Miocene distribution area that left from eastern or south-eastern regions. Such appearance of isolated endemic populations. This dispersal most probably lagomorph taxa with very different ecological requirements, occurred from south-eastern regions of Europe (e.g. the NW all together in a short time span, suggests that the onset of peri-Black Sea area), as the main morphological characters favourable climatic conditions was not the only and/or pri- of central European Pliocene Prolagus here analyzed is quite mary driving factor of their coeval dispersal. different from those of western European ones. Moreover, a dispersal wave from eastern regions fits quite well in the The post-Miocene fate of Prolagus: framework of the above discussed extensive faunal rearrange- climate change, palaeogeography, competitive pressure ments occurred during the early Pliocene. It is clear from the climatic and ecological frameworks above The subsequent, relatively sudden, disappearance of Prolagus illustrated that, except for the earliest stages (MN2b-MN3), from central Europe after MN15b can be explained by the Prolagus developed and passed unscathed through periods of increase of the competitive pressure related to the continuous profound climatic/environmental changes for at least 14 of global climatic degradation during the late Pliocene. In this the c. 20 my of its evolutionary history. Actually, Prolagus sense, Prolagus met the same fate of Pliopentalagus dietrichi in species had a wide range of environmental adaptations. Seve- Europe (see Čermák & Wagner 2013). In fact, the Pliocene ral species lived in subtropical forest or wetland lacustrine climatic deterioration caused an increase of the aridisation habitats, nevertheless other species, i.e., P. ibericus from Layna and the spreading of open ground habitats in central Europe. (MN15) or Prolagus spp. from the Gargano palaeoarchipelago, This opened the way for Ochotona Link, 1795 and Lepus Lin- inhabited arid environments (López Martínez 2001). It is naeus, 1758, very well adapted for those landscape settings, also possible that several of those species did not have strict and the Prolagus “story” in central Europe was over and was ecological requirements. Anyway, Prolagus was quite adapt- never repeated again. able to different and variable climatic conditions. Thus, it is reasonable to suggest that climate change sensu stricto was not the primary reason for the limited distribution of Prolagus CONCLUSIONS in the Pliocene of central Europe. A different perspective on the distribution of the genus Pro- This paper fills a gap in the study of Pliocene prolagids, until lagus in Europe comes from the palaeogeographical approach. now mainly centered on western European species, performing Until the middle Miocene, the geographic distribution of a detailed analysis of all available published and unpublished Prolagus was arranged in a roughly latitudinal band (as far Pliocene materials of Prolagus from central Europe. as the distribution of emerged lands permitted; cf. López Most of the record of central European Pliocene Prolagus Martínez 2001 for Prolagus distribution data, and Popov pertains to Prolagus bilobus. This species is reported here et al. 2004 for updated palaeogeographic setting). During the from the MN15b localities of Gundersheim and Raciszyn 1. late Miocene, the area in which Prolagus was distributed was Some materials of P. bilobus had already been revised and its deformed by the Alpine arc in its central part (the radial axis diagnosis emended (Čermák & Angelone 2013), however the of the arc, coinciding with the Rhine graben, was “pushed” availability of relatively rich materials allowed to define here northwards; see Sue et al. 2007). The bending movement may some additional peculiarities in tooth positions other than p3. have had an effect on isolation (the question is how much) The extremely low intraspecific variability of the specimens between western and eastern Prolagus populations, with the of P. bilobus is striking. This characteristic allowed to easily Alpine arc and the Bohemian Massif acting as physiographic describe the ontogenetic trajectory of the p3 of P. bilobus. barriers. Indeed, significantly increasing differences between From juvenile to adult, the p3 undergoes the enlargement, western European Prolagus species and central-eastern European deformation, and tilting of the anteroconid, the deformation ones start to be noticed at least since MN10 (Angelone & of the metaconid, a noticeable thickening of the protoconulid, Veitschegger 2015). the thickening of the entoconid, and the enlargement of the The decline in Prolagus abundance in northern central Europe crochet. In the mandibles of adult specimens, compared to appears evident already since the late Miocene (López Mar- the juveniles, it is possible to observe an enlargement of the tínez 2001). Although the mammalian fossil record around diastema in relation to the alveolar row length, the heighten- the Miocene/Pliocene boundary (late MN13 to MN14) is ing of the mandibular body, and an increase of the curvature extremely rare north of the Alps, Prolagus had probably disap- in the outline of the ventral margin. peared there since the latest Miocene. Further south, in the The study of the ontogenetic development of P. bilobus was Pannonian Basin, the genus was still present in the MN13 extended to the deciduous teeth (d3), and the dimensional (Angelone & Čermák 2015). For sure, Prolagus is absent results were compared with Prolagus species from several fossil COMPTES RENDUS PALEVOL • 2021 • 20 (28) 613 Čermák S. et al. localities ranging in age from the early Miocene (MN2b) to Acknowledgements the Middle-Late Pleistocene. In early Miocene-early middle We would like to express our thanks to László Pongrácz (Györ), Miocene taxa, the d3 is larger than p3, as occurs in Piezodus Katrin Krohmann (SMF), and Adam T. Halamski (ZPAL) and (López Martínez 1974). However, since late middle Miocene, M. Borsuk-Białynicka (ZPAL) for providing us the fossil mate- the size relationship appears reversed, and the values of d3 rial in their care. We are grateful to two anonymous reviewers remain quite stable. The reversal is not gradual: the d3/p3 for their valuable comments and critical remarks. The study relative length proportion is one of the characters that, in was supported by institutional support RVO67985831 of the the genus Prolagus, change their state at a certain geological Institute of Geology of the Czech Academy of Sciences. CA moment and in a trans-specific way, and, in this sense, it may was supported by: Visiting Professor grant of the President’s have a biochronological value. At any rate, the size difference International Fellowship Initiative of the Chinese Academy between d3/p3 in P. bilobus is clearly the largest among the of Science; Spanish Agencia Estatal de Investigación and the studied taxa. European Regional Development Fund of the European New materials from Beremend 26 and 39 (MN15b) revealed Union (CGL2016-76431-P); CERCA Program, Generalitat the presence of a possible new species of Prolagus. A lower de Catalunya; Grant to Department of Science, Roma Tre jaw of an adult individual from Beremend 26 appears clearly University (MIUR-Italy Dipartimenti di Eccellenza, ART. 1, distinct from other Pliocene-Holocene congeneric species. In C. 314-337 L. 232/2016). BMS was supported by Xunta de particular, the differences with the coeval P. bilobus are striking. Galicia (ED481B 2018/046, Axudas á etapa postdoutoral da Teeth and mandible dimensions fall below the lower range Xunta de Galicia 2018-Modalidade A). of P. bilobus, and also the morphology of p3 is incompatible with P. bilobus. Nevertheless, an upper jaw excavated from the neighbouring fissure filling Beremend 39 does not show sub- REFERENCES stantial differences in morphology and measures with P. bilobus. The two fillings seem coeval, basing on the biochronological AGADJANIAN A. K. & ERBAJEVA M. A. 1983. — [Late Cenozoic rodents and lagomorphs of the USSR]. Nauka, Moscow, 189 p. (in Russian). indications given by the faunal content. These contradictory ÁLVAREZ-SIERRA M. A., DAAMS R., LACOMBA J. I., LÓPEZ-MARTÍNEZ N., taxonomic evidences can be solved only by the retrieval of VAN DER MEULEN A. J., SESÉ C. & DE VISSER J. 1990. — Palae- additional material, and the prolagids from Beremend 26 and ontology and biostratigraphy (micromammals) of the continental 39 are provisionally left in open nomenclature as Prolagus sp. Oligocene-Miocene deposits of the North-Central Ebro basin Ukrainian and Moldavian MN15-?16 materials attributed (Huesca, Spain). Scripta Geologica 94: 1-77. ANGELONE C. 2007. — Messinian Prolagus (Ochotonidae, Lago- by Tesakov & Averianov (2002) to P. bilobus do not match morpha, Mammalia) of Italy. Geobios 40: 407-421. https://doi. the diagnosis of the species and, in our opinion, should be org/10.1016/j.geobios.2006.04.004 provisionally referred to P. aff. sorbinii. The material from ANGELONE C. 2008a. — Prolagus italicus n. sp. (Ochotonidae, Lago- Kisláng (mentioned by Kretzoi 1954) could have shed light morpha, Mammalia) a new Pliocene species of peninsular Italy. Geo- on the subject. Unfortunately, the material is currently una- bios 41: 445-453. https://doi.org/10.1016/j.geobios.2007.12.001 ANGELONE C. 2008b. — Family Ochotonidae (Lagomorpha) and vailable. The same fate was suffered by the Prolagus remains its application in biochronology: some case studies from the Plio- from Wölfersheim (faunal list in Tobien 1977; eventually Quaternary of Eurasia. Quaternary International 179: 5-8. https:// reported also by Dahlmann 2001). doi.org/10.1016/j.quaint.2007.08.019 Contrarily to the “traditional” view (López Martínez 2001), ANGELONE C. & ČERMÁK S. 2015. — Two new species of Prolagus we hypothesize that central European Prolagus are not relict (Lagomorpha, Mammalia) from the Late Miocene of Hungary: taxonomy, biochronology, and palaeobiogeography. Paläon- species “left behind” after a progressive reduction of the geo- tologische Zeitschrift 89: 1023-1038. https://doi.org/10.1007/ graphic distribution of the genus due to climate deterioration s12542-014-0247-z s.s. occurred during the Pliocene. Prolagus disappeared from ANGELONE C. & ROOK L. 2012. — Late Neogene and Quaternary northern central Europe since the latest Miocene to reappear lagomorphs from Tuscany: a revision based on specimens in Basel briefly in the late early Pliocene (MN15b). This evidence rather Naturhistorisches Museum and Florence University collections. indicates a sudden dispersal, most probably from south-eastern Swiss Journal of Geosciences 131: 127-145. https://doi.org/10.1007/ s13358-011-0035-2 regions of Europe, as the morphology of P. bilobus and Prola- ANGELONE C. & SESÉ C. 2009. — New characters for species dis- gus sp. studied herein exclude close phylogenetic affinities with crimination within the genus Prolagus (Ochotonidae, Lagomor- western European species. Prolagus is not the only lagomorph pha, Mammalia). Journal of Paleontology 83: 80-88. https://doi. to appear in central Europe in MN15b: others are e.g. Pli- org/10.1666/07-067R2.1 https://doi.org/10.1666/07-067R2.1 opentalagus and Ochotonoma, two taxa of clear eastern origin. ANGELONE C. & VEITSCHEGGER K. 2015. — MN10 Prolagus (Ochotonidae, Lagomorpha) from Austria: a new tile for the The MN15b central and south-eastern Europe lagomorph central European palaeogeography of the genus. Neues Jahrbuch palaeobiogeographic dynamics represent only a tile of the extensive für Geologie und Paläontologie 275: 1-10. https://doi.org/10.1127/ faunal rearrangements that characterize the entire Europe during njgpa/2015/0444 the Pliocene. The post-Miocene fate of Prolagus in central ANGELONE C., ČERMÁK S. & KOTSAKIS T. 2015. — The most ancient Europe was determined by a combination of palaeogeographic lagomorphs of Sardinia: an overview. Geobios 48: 287-296. https:// doi.org/10.1016/j.geobios.2015.06.002 and climatic changes which caused extensive rearrangements of ANGELONE C., MONCUNILL-SOLÉ B. & KOTSAKIS T. 2019. — Con- community and faunal structures that increased the competitive tribution of fossil Lagomorpha (Mammalia) to the refinement pressure on the genus. of the late Miocene-Quaternary palaeobiogeographical setting 614 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe of Italy. Comptes Rendus Palevol 18: 1025-1040. https://doi. ERGHEM C., H ENNISSEN J., H EAD M. J., L OUWYE S. & org/10.1016/j.crpv.2019.10.002 FABIAN K. 2009. — Northern Hemisphere Glaciation during ANGELONE C., MONCUNILL-SOLÉ B. & KOTSAKIS T. 2020. — Fossil the Globally Warm Early Late Pliocene. PLoS ONE 8(12): e81508. Lagomorpha (Mammalia) of Italy: systematics and biochronol- https://doi.org/10.1371/journal.pone.0081508 ogy. Rivista Italiana di Paleontologia e Stratigrafia 126: 157-187. DE SCHEPPER S., GIBBARD P. L., SALZMANN U. & EHLERS J. 2014. https://doi.org/10.13130/2039-4942/13014 — A global synthesis of the marine and terrestrial evidence for AVERIANOV A. O. & TESAKOV A. S. 1998. — Lagomorphs (Mam- glaciation during the Pliocene Epoch. Earth-Science Reviews malia, Lagomorpha) from early Pliocene locality of Kosyakino 135: 83-102. https://doi.org/10.1016/j.earscirev.2014.04.003 of the Northern Caucasus. Paleontological Journal 32: 305-309. DICE L. R. 1917. — Systematic position of several tertiary lago- BERNOR R. L., KOUFOS G. G., WOODBURNE M. & FORTE- morphs. University of California Publications, Bulletin of the LIUS M. 1996. — The evolutionary history and biochronology Department of Geology 10: 179-183. of european and southeastern asian late Miocene and Pliocene ERBAJEVA M. A. 1988. — [Cenozoic Pikas (Taxonomy, Systematics, hipparionine horses, in BERNOR R. L., FAHLBUSCH V. & MITT- Phylogeny)]. Nauka, Moscow, 224 p. (in Russian). MANN H. W. (eds), The evolution of Western Eurasian Neogene ERBAJEVA M. A. & SHUSHPANOV K. I. 1988. — [Pliocene pikas of mammal faunas. Columbia University Press, New York: 307-338. Moldavia]. Vestnik Zoologii 4: 55-60. (in Russian). BERZI A. 1967. — Lagomorphs from the type Villafranchian of FEJFAR O. 1961. — Die plio-pleistozänen Wirbeltierfaunen von Villafranca d’Asti (Italy). Preliminary note. Giornale di Geologia Hajnáčka und Ivanovce (Slowakei), ČSSR. III. Lagomorpha. Neues 35: 137-150. Jahrbuch für Geologie und Paläontologie, Monatshefte 5: 267-282. BÖHME M. 2003. — The Miocene Climatic Optimum: evidence FEJFAR O. & HEINRICH W.-D. 1983. — Arvicoliden-Sukzession from ectothermic vertebrates of Central Europe. Palaeogeogra- und Biostratigraphie des Oberpliozäns und Quartärs in Europa. phy, Palaeoclimatology, Palaeoecology 195: 389-401. https://doi. Schriftenreihe für geologische Wissenschaften 19/20: 61-109. org/10.1016/S0031-0182(03)00367-5 FEJFAR O. & STORCH G. 1990. — Eine pliozäne (ober-ruscinische) BÖHME M., ILG A. & WINKLHOFER M. 2008. — Late Miocene Kleinsäugerfauna aus Gundersheim, Rheinhessen: 1. Nagetiere: “washhouse” climate in Europe. Earth and Planetary Science Let- Mammalia, Rodentia. Senckenbergiana lethaea 71: 139-184. ters 275: 393-401. https://doi.org/10.1016/j.epsl.2008.09.011 FEJFAR O., HEINRICH W.-D. & LINDAY E. H. 1998. — Updating BRANDT J. F. 1855. — Beiträge zur näheren Kenntniss der Säugeth- the Neogene Rodent biochronology in Europe. Mededelingen iere Russlands. Mémoires de l’Académie impériale des sciences de Nederlands Instituut voor Toegepaste Geowetenschappen TNO 60: St.-Pétersbourg 9: 1-365. 533-554. CASANOVAS-VILAR I., VAN DEN HOEK OSTENDE L. W., FURIÓ M. & FEJFAR O., STORCH G. & TOBIEN, H. 2006. — Gundersheim 4, a MADERN A. P. 2014. — The range and extent of the Vallesian third Ruscinian micromammalian assemblage from Germany. Crisis (Late Miocene): new prospects based on the micromam- Palaeontographica A 278: 97-111. https://doi.org/10.1127/ mal record from the Vallès-Penedès basin (Catalonia, Spain). pala/278/2006/97 Journal of Iberian Geology 40: 29-48. https://doi.org/10.5209/ FLADERER F. A. & REINER G. 1996. — Evolutionary shifts in the rev_JIGE.2014.v40.n1.44086 first premolar pattern of Hypolagus beremendensis (Petényi, 1864) ČERMÁK S. 2007. — New finds of Ochotonoma csarnotana (Lago- (Lagomorpha, Mammalia) in the Plio-Pleistocene of Central morpha, Ochotonidae) from the Pliocene of Hungary: a new look Europe. Acta Zoologica Cracoviensia 39: 147-160. on the species. Neues Jahrbuch für Geologie und Paläontologie 246: FOSTOWICZ-FRELIK Ł. 2010. — A new species of Pliocene Prolagus 247-256. https://doi.org/10.1127/0077-7749/2007/0246-0247 (Lagomorpha: Ochotonidae) from Poland in the northermost ČERMÁK S. 2009. — The Plio-Pleistocene record of Hypolagus record of the genus. Journal of Vertebrate Paleontology 30: 609- (Lagomorpha, Leporidae) from the Czech and Slovak Republics 612. https://doi.org/10.1080/02724631003621789 with comments on systematics and classification of the genus. GANOPOLSKI A. & CALOV R. 2011. — The role of orbital forcing, Bulletin of Geosciences 84: 497-524. https://doi.org/10.3140/ carbon dioxide and regolith in 100 kyr glacial cycles. Climate of bull.geosci.1104 the Past 7: 1415-1425. https://doi.org/10.5194/cp-7-1415-2011 ČERMÁK S. & ANGELONE C. 2013. — Revision of the type material GUREEV A. A. 1960. — [Lagomorphs (Lagomorpha) from the Oli- of the Pliocene species Prolagus bilobus Heller, 1936 (Mamma- gocene of Mongolia and Kazakhstan]. Trudy Paleontologicheskogo lia, Lagomorpha), with comments on the taxonomic validity of Instituta Akademii Nauk SSSR 77: 5-34. (in Russian). P. osmolskae Fostowicz-Frelik, 2010. Bulletin of Geosciences 88: GUREEV A. A. 1964. — [Fauna of the USSR (Lagomorpha), Vol. 3 45-50. https://doi.org/10.3140/bull.geosci.1369 (10)]. Nauka, Moscow & Leningrad. 276 p. (in Russian). ČERMÁK S. & WAGNER J. 2013. — The Pliocene record of Trischiz- HELLER F. 1936. — Eine oberpliozäne Wirbeltierfauna aus Rheinhes- olagus and Pliopentalagus (Leporidae, Lagomorpha, Mammalia) in sen. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, Central Europe, with comments on taxonomy and evolutionary Abteilung B 76: 99-160. history of Leporinae. Neues Jahrbuch für Geologie und Paläontologie HORDIJK K. & VAN DER MEULEN A. J. 2010. — Systematics of 268: 97-111. https://doi.org/10.1127/0077-7749/2013/0321 resident species of Lagopsis and Prolagus (Ochotonidae, Lago- CZÁSZÁR G. & KORDOS L. 2004. — Beremend, Kőfejtő. Pro- morpha, Mammalia) from the late early and middle Miocene of gram, előadáskivonatok, kirándulásvezető, 7. Magyar Őslénytani northeastern Central Spain. Geologica Ultraiectina 333: 17-85. Vándorgyűlés 2004: 51-57. JÁNOSSY D. 1972. — Middle Pliocene microvertebrate fauna from DAHLMANN T. 2001. — Die Kleinsäuger der unter-pliozänen Fund- the Osztramos Loc. 1. (Northern Hungary). Annales historico- stelle Wölfersheim in der Wetterau (Mammalia: Lipotyphla, naturales Musei nationalis hungarici 64: 27-50. Chiroptera, Rodentia). Courier Forschungsinstitut Senckenberg JÁNOSSY D. 1986. — Pleistocene vertebrate faunas of Hungary. Aka- 227: 1-129. démiai Kiadó, Budapest, 208 p. DAHLMANN T. & STORCH G. 1996. — Eine pliozäne (oberruschi- KOENIGSWALD W. VON & TOBIEN H. 1990. — Important arvi- nische) Kleinsäugerfauna aus Gundersheim, Rheinhessen. 2. colid faunas from the Late Pliocene to Early Holocene in West- Insektenfresser: Mammalia, Lipotyphla. Senckenbergiana lethaea ern Germany (FRG), in FEJFAR O. & HEINRICH W.-D. (eds), 76: 181-191. https://doi.org/10.1007/BF03042849 International symposium evolution, phylogeny and biostratigraphy DAXNER G. & FEJFAR O. 1967. — Über die gattung Alilepus Dice, of arvicolids (Rodentia, Mammalia). Czech Geological Survey, 1931 und Pliopentalagus Gureev, 1964 (Lagomorpha, Mamma- Prague: 231-254. lia). Annalen des Naturhistorischen Museums in Wien 71: 37-55. KÖNIG C. D. E. 1825. — Icones Fossilium Sectiles, Centuria Prima. DE SCHEPPER S., GROENEVELD J., NAAFS B. D. A., VAN RENT- G.B. Sowerby, London, 4 p. COMPTES RENDUS PALEVOL • 2021 • 20 (28) 615 Čermák S. et al. KORDOS L. 1991. — Upper Pliocene paleovertebrate localities, NADACHOWSKI A. 1990. — Review of fossil Rodentia from Poland Beremend, Villány Mts. Magyarország geológiai alapszevényei (Mamalia). Senckenbergiana Biologica 70: 229-250. 1991: 1-6. NADACHOWSKI A., MARCISZAK A., RZEBIK-KOWALSKA B. & KORMOS T. 1930. — Beiträge zur Präglazialfauna des Somlyóberges GORNIG W. 2015. — Sesja terenowa C. Raciszyn. Fauna kopalna- bei Püspökfürdo. Állattani Közlemények 27: 40-62. Raciszyn 1 (RN1), in KICIŃSKA D., STEFANIAK K. & SZYNK- KOVAR-EDER J., JECHOREK H., KVAČEK Z. & PARASCHIV V. 2008. — IEWICZ A. (eds), Przewodnik sesji terenowych. Materiały 49. The Integrated Plant Record: An Essential Tool For Reconstruct- Sympozjum Speleologicznego, Załęcze Wielkie 22-25.10.2015. ing Neogene Zonal Vegetation In Europe. Palaios 23: 97-111. Sekcja Speleologiczna Polskiego Towarzystwa Przyrodników im. https://doi.org/10.2110/palo.2006.p06-039r Kopernika, Kraków: 50-51. KOWALSKI K. 1960. — Cricetidae and Microtidae from the Pliocene NESIN V. A. & NADACHOWSKI A. 2001. — Late Miocene and Plio- of Weze (Poland). Acta Zoologica Cracoviensia 5: 447-506. cene small mammal faunas (Insectivora, Lagomorpha, Rodentia) KOWALSKI K. 1990. — Stratigraphy of Neogene mammals in Poland, of southeastern Europe. Acta Zoologica Cracoviensia 44: 107-135. in LINDAY E. H., FAHLBUSCH V. & MEIN P. (eds), European Neo- PETÉNYI S. J. 1864. — A beremendi mészkőbánya természetrajz-és gene Mammal Chronology. Plenum Press, New York: 193-209. őslénytanilag Petényi Salamon által leírva, in KUBINYI F. (ed.), KRETZOI M. 1954. — Bericht über die calabrische (villafranchische) Petényi S. János Hátrahagyott munkái. Magyar Tudományos Fauna von Kislang, Kom. Fejér. Állami Földtani Intézet Évijjelen- Akademia, Pest: 35-81. tése az 1953 1: 239-265. POMEL M. 1853. — Catalogue méthodique et descriptif des vertébrés KRETZOI M. 1956. — Die altpleistozänen Wirbeltierfaunen des fossiles découverts dans le bassin hydrographique supérieur de la Villányer Gebirges. Geologica hungarica, Series palaeontologica Loire, et surtout dans la vallée de son affluent principal, l’Allier. 27: 1-264. J.-B. Baillière, Paris, 193 p. KRETZOI M. 1959. — Insectivoren, Nagetiere und Lagomorphen PONGRÁCZ L. 1999. — A beremendi Szőlő-hegy természettudományi der jüngstpliozänen Fauna von Csarnóta im Villáyer Gebirge kutatásának 150 éve. Petényi-emlékkönyv, Beremend, 149 p. (Südungarn). Vertebrata Hungarica 2: 237-246. POPOV V. V. 2001. — Late Pliocene voles (Mammalia: Arvicolidae) KRETZOI M. 1962. — Fauna und Faunenhorizont von Csarnóta. from Varshets (North Bulgaria). Acta Zoologica Cracoviensia 44: Jahresbericht der Ungarischen Geologischen Anstalt 1959: 297-395. 143-172. LINK D. H. F. 1795. — Beiträge zur Naturgeschichte. Rostock und POPOV V. V. 2003. — Late Pliocene Soricidae (Insectivora, Mam- Leipzig 2: 1-126. malia) from Varshets (North Bulgaria). Acta Zoologica Cracov- LINNAEUS C. 1758. — Systema naturae per Regna tria naturae, secun- iensia 46: 43-72. dum Classes, Ordines, Genera, Species. Tomus I. Editio decima, POPOV V. V. 2004. — Pliocene small mammals (Mammalia, Lipo- reformata. Laurentii Salvii, Holmiæ, 824 p. typhla, Chiroptera, Lagomorpha, Rodentia) from Muselievo LÓPEZ MARTÍNEZ N. 1974. — Evolution de la lignée Piezodus- (North Bulgaria). Geodiversitas 26: 403-491. Prolagus (Lagmorpha, Ochotonidae) dans le Cenozoïque d’Europe P OPOV S. V., R ÖGEL F., R OYANOV A. Y., S TEININGER F. F., Sud-Occidentale. Unpublished PhD Thesis. Université des Science SHCHERBA I. G. & KOVÁČ M. 2004. — Lithological–paleogeo- et techniques du Languedoc, Académie de Montpellier, 165 p. graphic maps of Paratethys. 10 Maps Late Eocene to Pliocene. LÓPEZ MARTÍNEZ N. 1989. — Revisión sistemática y bioestratigrá- Courier Forschungsinstitut Senckenberg 250: 1-46. fica de los Lagomorpha (Mammalia) del Terciario y Cuaternario RABEDER G. 1981. — Die Arvicoliden (Rodentia, Mammalia) aus inferior de España. Memorias del Museo Paleontológico de la Uni- dem Pliozän und dem ältesten Pleistozän von Niederösterreich. versidad de Zaragoza 3: 1-350. Beiträge zur Paläontologie von Österreich 10: 1-373. LÓPEZ MARTÍNEZ N. 2001. — Paleobiogeographical history of Prolagus, RABEDER G. & VERGINIS S. 1987. — Die plio-pleistozänen löesspro- an European ochotonid (Lagomorpha). Lynx (n.s.) 32: 215-231. file von Stranzendorf und Krems (Niederösterreich). Griechische LÓPEZ MARTÍNEZ N. & THALER L. 1975. — Biogéographie, évolu- Geographische Gesellschaft 1. Panhellenische Geographen-Tagung tion et compléments à la systématique du groupe d’Ochotonides Athen. 9: 285-306. Piezodus-Prolagus (Mammalia, Lagomorpha). Bulletin de la Société RADULESCO C. & SAMSON P. 1967. — Contribution à la connais- géologique de France 17: 850-866. https://doi.org/10.2113/gss- sance du complexe faunique de Măluşteni–Bereşti (Pléistocène gfbull.S7-XVII.5.850 inférieur), Roumanie I. Ord. Lagomorpha, Fam. Leporidae. Neues LÓPEZ MARTÍNEZ N., SESÉ BENITO C. & SANZ GARCÍA J. L. 1977. — Jahrbuch für Geologie und Paläontologie, Monatshefte 9: 544-563. La microfauna (Rodentia, Insectivora y Reptilia) de las fisuras del RAUP D. M. & CRICK R. E. 1979. — Measurement of faunal simi- Mioceno medio de Escobosa de Calatañazor (Soria, España). Acta larity in paleontology. Journal of Paleontology 53: 1213-1227. Geologica Hispanica 12, 60-68. http://hdl.handle.net/10261/3393 REPENNING CH. A. 1967. — Subfamilies and Genera of the Soricidae. LUNT D. J., FOSTER G. L., HAYWOOD A. M. & STONE E. J. 2008. — Classification, historical zoogeography and temporal correlation Late Pliocene Greenland glaciation controlled by a decline in of the shrews. United States Survey Professional Paper 565: 1-74. atmospheric CO2 levels. Nature 454: 1102-1105. https://doi. https://doi.org/10.3133/pp565 org/10.1038/nature07223 REUMER J. W. F. 1984. — Ruscinian and early Pleistocene Soricidae MARIDET O., ESCARGUEL G., COSTEUR L., MEIN P., HUGUENEY M. & (Insectivora, Mammalia) from Tegelen (The Nederlands) and LEGENDRE S. 2007. — Small mammal (rodents and lagomorphs) Hungary. Scripta Geologica 73: 1-173. European biogeography from the Late Oligocene to the mid REUMER J. W. F. 1989. — Speciation and evolution in the Soricidae Pliocene. Global Ecology and Biogeography 16: 529-544. https:// (Mammalia: Insectivora) in relation with the paleoclimate. Revue doi.org/10.1111/j.1466-8238.2006.00306.x Suisse Zoology 96: 81-90. https://doi.org/10.5962/bhl.part.117758 MASINI F. 1989. — Prolagus sorbinii, a new ochotonid (Mammalia, REUMER J. W. F. & WESSELS W. 2003. — Distribution and migra- Lagomorpha) from the Messinian of Italy. Bollettino della Società tion of Tertiary mammals in Eurasia. A volume in honour of Paleontologica Italiana 28: 295-306. Hans de Bruijn. Deinsea 10, 576 p. MAYHEW D. F. 2012. — Revision of the fossil vole assemblage ROMAN F. & VIRET J. 1930. — Le Miocène continental de l’Armagnac (Mammalia, Rodentia, Arvicolidae) from Pleistocene Depos- et le gisement burdigalien de La Romieu (Gers), in DE MAR- its at Kisláng, Hungary. Palaeontology 55: 11-29. https://doi. GERIE E., BARROIS CH., BERTRAND L., BERTRAND P. & BOUR- org/10.1111/j.1475-4983.2011.01115.x CART J. (eds), Livre Jubilaire de la Société géologique de France 2. MICHAUX J. 1971. — Arvicolinae (Rodentia) du Pliocéne terminal Société Géologique de France, Paris: 577-604. et du Quatemaire ancien de France et d’Espagne. Palaeovertebrata RÖSSNER G. E. & HEISSIG K. 1999. — The Miocene Land Mammals 4: 137-214. https://doi.org/10.18563/pv.4.5.137-214 of Europe. Verlag Dr. Friedrich Pfeil, Munich, 516 p. 616 COMPTES RENDUS PALEVOL • 2021 • 20 (28) Prolagus in the framework of the Pliocene faunal rearrangements in central Europe ŞEN Ş. 1998. — Pliocene vertebrate locality of Çalta, Ankara, Turkey. TOBIEN H. 1935. — Über die pleistozänen und postpleistozänen 4. Rodentia and Lagomorpha. Geodiversitas 20: 359-378. Prolagusformen Korsikas und Sardiniens. Berichte der naturfor- ŞEN Ş., BOUVRAIN G. & GERAADS D. 1998. — Pliocene vertebrate schenden Gesellschaft zu Freiburg im Breisgau 34: 253-344. locality of Çalta, Ankara, Turkey. 12. Paleoecology, biogeography TOBIEN H. 1977. — 7. Fauna, in BOENIGK W., BRELIE G., BRUN- and biochronology. Geodiversitas 20: 497-510. NACKER K., KEMPF E. K., KOČÍ A., SCHIRMER W., STADLER G., SHI N. 1994. — The Late Cenozoic Stratigraphy, Chronology, Palynol- STREIT R. & TOBIEN H. (eds), Jungtertiär und Quartär im Horloff- ogy and Environmental Development in the Yushe Basin, North Graben, Vogelsberg. Geologische Abhandlungen Hessen 75: 65-68. China. Striae 36: 1-90. TOBIEN H. 1980. — Säugerfaunen von der Grenze Pliozän/Pleis- SKOCZEN S. 1980. — Scaptomychini Van Valen, 1967, Urotrichini and tozän in Rheinhessen. 1. Die Spaltfüllungen von Gundersheim Scalopini Dobson, 1883 (Insectivora, Mammalia) in the Pliocene bei Worms. Mainzer geowissenschaftliche Mitteilungen 8: 209-218. and Pleistocene of Poland. Acta Zoologica Cracoviensia 24: 411-448. TOPACHEVSKY V. O. 1962. — [On the Geological Age of Early STONE W. 1900. — Descriptions of a New Rabbit from the Liu Kiu Alluvial deposits of the Lower Dnieper Valley]. Geologicheskii Islands and a New Flying Squirrel from Borneo. Proceedings of the zhurnal 22: 106-109 (in Ukrainian). Academy of Natural Sciences of Philadelphia 52: 460-463. ÜNAY E. & DE BRUIJN H. 1998. — Plio-Pleistocene rodents and SUE C., DELACOU B., CHAMPAGNAC J.-D., ALLANIC C., TRICART P. & lagomorphs from Anatolia. Mededelingen Nederlands Instituut BURKHARD M. 2007. — Extensional neotectonics around the voor Toegepaste Geowetenschappen TNO 60: 431-466. bend of the Western/Central Alps: An overview. International VAN DER MEULEN A. & VAN KOLFSCHOTEN T. 1986. — Review of Journal of Earth Sciences 96: 1101-1129. https://doi.org/10.1007/ the Late Turolian to Early Biharian mammal faunas from Greece s00531-007-0181-3 and Turkey. Memoria Società Geologia Italiana 31: 201-211. SULIMSKI A. 1959. — Pliocene Insectivores from Węże (Owadożerne VAN DE WEERD A. 1979. — Early Ruscinian rodents and lago- z plioceńskiej brekcji kostnej z Wężów). Acta Palaeontologica morphs (Mammalia) from the lignites near Ptolemais (Macedonia, Polonica 4: 119-179. Greece). Proceedings of the Koninklijke Nederlandse Akademie van SULIMSKI A. 1962. — O nowym znalezisku kopalnej fauny kregow- Wetenschappen, Series B Physical Sciences 82: 127-168. ców w okolicy Działoszyna. Przegląd Geologiczny 10: 219-223. VAN DE WEERD A. & DAAMS R. 1978. — Quantitative composition SULIMSKI A. 1964. — Pliocene Lagomorpha and Rodentia from Węże of rodent faunas in the Spanish Neogene and palaeoecological 1 (Poland). Acta Palaeontologica Polonica 9: 149-262. implications. Proceedings of the Koninklijke Nederlandse Akademie SULIMSKI A. & SZYNKIEWICZ A. 1994. — Pliocene small mammals van Wetenschappen, Series B Physical Sciences 81: 448-473. from Raciszyn 1 (Cracow-Wieluń Upland, Central Poland), in VAN DE WEERD A., REUMER J. W. F. & DE VOS J. 1982. — Pliocene Neogene and Quaternary Mammals of the Palearctic: Abstracts of the mammals from Apolakkia Formation (Rhodes, Greece). Proceed- Conference in Honour of Prof. Kazimierz Kowalski. Polish Academy ings of the Koninklijke Nederlandse Akademie van Wetenschappen, of Sciences, Kraków: 69. Series B Physical Sciences 85: 89-112. SYCH L. 1965. — Fossil Leporidae from the Pliocene and the Pleis- VIRET J. 1929. — Les faunes de mammifères de l’Oligocène supé- tocene of Poland. Acta Zoologica Cracoviensia 10: 1-88. rieur de la Limagne bourbonnaise. Annales de l‘Université de Lyon, nouvelle série 1 47: 1-327. SZYNKIEWICZ A. & JAGIEŁŁO M. 2015. — Zjawiska krasowe w rejonie WAGNER R. 1829. — Über die Knochenbrekzie in Sardinien und die Raciszyna, in KICIŃSKA D., STEFANIAK K., SZYNKIEWICZ A. (eds), darin gefundenen Thiere, so wie über einige andere hieher gehörige Przewodnik sesji terenowych. Materiały 49. Sympozjum Speleologicznego, Erscheinungen. Archiv für die gesammte Naturlehre 15: 10-31. Załęcze Wielkie 22-25.10.2015. Sekcja Speleologiczna Polskiego WAGNER J., ČERMÁK S., HORÁČEK I., FEJFAR O. & MIHEVC A. 2009. — Towarzystwa Przyrodników im. M. Kopernika, Kraków: 43-50. New mammalian fossil records refining a view on Early/Late TAN N., RAMSTEIN G., DUMAS C., CONTOUX C., LADANT J.-B., SEP- Pliocene faunal turnover in Central Europe. Journal of Vertebrate ULCHRE P., ZHANG Z. S. & DE SCHEPPER S. 2017. — Exploring the Paleontology 29 (Supl. 3): 197A. MIS M2 glaciation occurring during a warm and high atmospheric WIBLE J. R. 2007. — On the Cranial Osteology of the Lagomor- CO2 Pliocene background climate. Earth and Planetary Science pha. Bulletin of Carnegie Museum of Natural History 39: 213- Letters 472: 266-276. https://doi.org/10.1016/j.epsl.2017.04.050 234. https://doi.org/10.2992/0145-9058(2007)39[213:OTC T EDFORD R. H., F LYNN J. L., Q IU Z., O PDYKE H. & OOT]2.0.CO;2 DOENS W. R. 1991. — Yushe basin, China, Paleomagnetically XUE X., ZHANG Y. & YUE L. 2006. — Paleoenvironments indi- calibrated mammalian biostratigraphic standard for the Late Neo- cated by the fossil mammalian assemblages from red clay-loess gene of eastern Asia. Journal of Vertebrate Paleontology 11: 519-526. sequence in the Chinese Loess Plateau since 8.0 Ma B.P. Sci- https://doi.org/10.1080/02724634.1991.10011420 ence in China Series D 49: 518-530. https://doi.org/10.1007/ TERZEA E. 1997. — Biochronologie du pliocène du bord Méridional s11430-006-0518-y du bassin Dacique (Roumanie). Mémoires et Travaux de l’Institut ZACHOS J. C., PAGANI M., SLOAN L. C., THOMAS E. & BIL- de Montpellier 21: 649-660. LUPS K. 2001. — Trends, rhythms, and aberrations in global TESAKOV A. S. & AVERIANOV A. O. 2002. — Prolagus (Lagomorpha, climate 65 Ma to present. Science 292: 686-693. https://doi. Prolagidae) from the Pliocene of Moldova and Ukraine. Paleonto- org/10.1126/science.1059412 logical Journal 36: 80-86. Submitted on 21 February 2020; accepted on 25 May 2020; published on 28 June 2021. COMPTES RENDUS PALEVOL • 2021 • 20 (28) 617