PALEVO 1631-0683 Elsevier S1631-0683(19)30131-9 10.1016/j.crpv.2019.07.004 Research article General Palaeontology, Systematics, and Evolution (Vertebrate Palaeontology) General Palaeontomogy, Systematics, and Evolution/Paléontologie générale, systématique et évolution (Vertebrate Palaeontology/Paléontologie des Vertébrés) Stephanorhinus jeanvireti (Mammalia, Rhinocerotidae) from the early Pleistocene of Colțești (southwestern Romania) Stephanorhinus jeanvireti (Mammalia, Rhinocerotidae) du Pléistocène inférieur de Colteşti (Sud-Ouest de la Roumanie) Pandolfi Luca luca.pandolfi@unifi.it a Codrea Vlad A. b Popescu Aurelian c Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via Giorgio La Pira, 4, 50121 Firenze, Italy Dipartimento di Scienze della Terra, Università degli Studi di Firenze Via Giorgio La Pira, 4 Firenze 50121 Italy Babeș-Bolyai University of Cluj-Napoca, Laboratory of Palaeotheriology and Quaternary Geology, 1 Kogălniceanu Str., 400084 Cluj-Napoca, Romania Babeș-Bolyai University of Cluj-Napoca, Laboratory of Palaeotheriology and Quaternary Geology 1 Kogălniceanu Str. Cluj-Napoca 400084 Romania Museum of Oltenia Craiova, Department of Natural Sciences, 8 Popa Sapcă Str., 200424 Craiova, Romania Museum of Oltenia Craiova, Department of Natural Sciences 8 Popa Sapcă Str. Craiova 200424 Romania 18 8 S1631-0683(19)X0009-3 1041 1056 © 2019 Académie des sciences. Published by Elsevier B.V. All rights reserved. 2019 Académie des sciences Full (PDF)

The Rhinocerotidae material from the early Pleistocene Tetoiu Formation of Colțești (southwestern Romania) is here described for the first time. The rhinoceros is documented by calcaneus and second, third, and fourth metatarsals, probably belonging to the same individual. The morphology and the dimensions of these specimens enable us to record the presence of Stephanorhinus jeanvireti, a relatively rare rhinoceros usually reported from late Pliocene European localities and recently considered a junior synonym of S. elatus. Nevertheless, the taxon Rhinoceros elatus is here regarded as a nomen dubium and the name S. jeanvireti is retained in order to maintain nomenclatural stability, being it based on much more diagnostic material. In Romania, S. jeanvireti has been listed within a few late Pliocene faunal assemblages, but the rhinoceros remains are fragmentary and isolated bones. The Colțești find is among the best documented records of S. jeanvireti in Romania. The biochronological distribution of S. jeanvireti is mainly confined within the MNQ16 (early and early middle Villafranchian), and its first occurrence is doubtfully reported in late MNQ15 faunas. The record of Colțești (MNQ17/MNQ18) represents, instead, the last occurrence of this taxon in Europe.

Le matériel de Rhinocerotidae du Pléistocène inférieur de la formation Tetoiu de Colteşti (Sud-Ouest de la Roumanie) est ici décrit pour la première fois. Le rhinocéros est documenté par le calcanéum et les deuxième, troisième et quatrième métatarses, appartenant probablement au même individu. La morphologie et les dimensions de ces spécimens permettent d’enregistrer la présence de Stephanorhinus jeanvireti, un rhinocéros relativement rare, signalé d’ordinaire dans des localités européennes du Pliocène terminal et récemment considéré comme un synonyme juvénile de S. elatus. Néanmoins, le taxon Rhinoceros elatus est ici considéré comme un nomen dubium et la dénomination S. jeanvireti est retenue pour maintenir la stabilité de la nomenclature, basée sur beaucoup plus de matériel diagnostique. En Roumanie, S. jeanvireti a été listé dans plusieurs assemblages fauniques du Pliocène tardif, mais les restes de rhinocéros sont des os fragmentaires et isolés. La découverte faite à Colteşti figure parmi les enregistrements les mieux documentés de S. jeanvireti en Roumanie. La distribution biochronologique de S. jeanvireti est majoritairement limitée au MNSQ16 (Villafranchien inférieur et inférieur-moyen) et sa première occurrence est rapportée de manière douteuse dans les faunes du MNQ15 terminal. L’enregistrement de Colteşti (MNQ17/MNQ18) représente, au contraire, la dernière occurrence de ce taxon en Europe.

 Stephanorhinus, Morphology, Taxonomy, early Pleistocene, Paleobiogeography, Oltenia Province, southwestern Romania Stephanorhinus, Morphologie, Taxonomie, Pléistocène inférieur, Paléobiogéographie, Province d’Oltenia, Sud-Ouest de la Roumanie presented Handled by Lorenzo Rook Introduction

The occurrence of Rhinocerotidae during the Pliocene and early Pleistocene in eastern Europe is still poorly known, despite this area represents an important “bridge” between Western Europe and Asia. In this context, a revision of Plio-Pleistocene rhinoceroses from Eastern Europe has started in the last years and has contributed to improve knowledge of different fossil species such as ‘Dihoplus’ megarhinus (de Christol, 1834), Stephanorhinus etruscus (Falconer, 1868) and Stephanorhinus hundsheimensis (Toula, 1906) (Boulbes et al., 2014, Konidaris et al., 2015, Pandolfi and Erten, 2017, Pandolfi et al., 2017, Pandolfi et al., 2016, Pandolfi et al., 2015a and Spassov, 2005 and references therein). However, Pliocene European rhinoceroses are still poorly investigated, and relatively poorly documented with respect to the late Miocene and early Pleistocene ones. During the Pliocene (MNQ14, MNQ15, MNQ16), at least four species are documented in Europe: ‘D.’ megarhinus, considered a typical Pliocene species (Guérin, 1980 and Pandolfi, 2013), but also recorded during the latest Miocene (Pandolfi and Rook, 2017, Pandolfi et al., 2016 and Pandolfi et al., 2015a); ‘Stephanorhinus’ miguelcrusafonti (Guérin and Santafé-Llopis, 1978), reported at a few early Pliocene Spanish and French localities (Cerdeño, 1992, Guérin, 1980 and Guérin and Santafé-Llopis, 1978); Stephanorhinus jeanvireti (Guérin, 1972: see note in Systematic Paleontology; Campanino et al., 1994 and Pandolfi, 2013) documented from some late Pliocene localities; and S. etruscus that occurred in Europe at the end of the Pliocene and during the early Pleistocene (Cerdeño, 1993, Pandolfi et al., 2017 and Pandolfi et al., 2015b and references therein).

Starting with late Miocene in Romania as in other regions of Europe (Guérin, 1980 and Heissig, 1996), Rhinocerotidae decreased their presence in the faunal communities in both diversity and abundance (Rădulescu and Samson, 2001 and Rădulescu et al., 2003). ‘Dihoplus’ megarhinus (Rădulescu and Samson, 2001 and Rădulescu et al., 2003; therein as Stephanorhinus leptorhinus in partim), during the early Pliocene (Dacian; Andreescu et al., 2011, Andreescu et al., 2013 and Rădulescu and Samson, 2001), S. jeanvireti, during the late Pliocene (Rădulescu and Samson, 2001 and Rădulescu et al., 2003; therein as Stephanorhinus elatus in partim), and S. etruscus, during the early Pleistocene, (Pandolfi et al., 2017 and references therein) were listed within some faunal assemblages.

The aim of this paper is to describe postcranial rhinoceros remains collected from early Pleistocene deposits at Colțești (Oltenia Province, southwestern Romania; Fig. 1) and to discuss the biochronological implications of this finding.

Southwestern Romania is perhaps the most representative region where Pliocene and early Pleistocene sedimentary deposits are widely exposed in that country. These deposits have depositional histories related either to the peculiar sedimentary evolution of the western side of the Dacian Lake or to the fluvial system that replaced this lake after its warping, in the late Pliocene (Codrea, 1997, Codrea et al., 2006 and Jipa, 2006).

For a long time and even hitherto, the Pliocene stratigraphy of Oltenia has been mainly based on mollusks (Andreescu et al., 2011 and Andreescu et al., 2013 and references therein). Nevertheless, the fossil vertebrates—and overall mammal ones,—added useful details, leading to a better stratigraphy (Andreescu et al., 2013, Feru et al., 1983, Rădulescu and Samson, 1990, Rădulescu and Samson, 2001 and Rădulescu et al., 2003).

 Geological and stratigraphic background

The vertebrate locality Colțești was found in April 1999. Colțești is a village located 51 km southeast from Târgu Jiu, nearby Hurezani, in the Gorj District (Fig. 1). The fossils were recovered in an unusual way, due to the works connective to a drilling rig-up carried out by the drillers from Drilling Boreholes Craiova (Foraj Sonde S.A. Craiova) Society. More precisely, the workers dug a pit mud using a bulldozer. A part of the scattered bones was donated by the drilling staff to the “Al. Stefulescu” Museum in Târgu Jiu and another part to the Oltenia Museum, Natural Sciences Branch in Craiova.

From a structural viewpoint, Colțești is situated in the southern Carpathians Foredeep. This unit started its evolution next to the Latest Cretaceous (‘Laramian’) tectonic phase, finally thrusting the Moesian Platform (Săndulescu, 1984). The geological history began in the earliest Cenozoic. Between the Oligocene and the early Miocene, this region was part of the Proto- (Rusu, 1977) and Eo-Paratethys Sea realms, followed in the middle Miocene by the Meso-Paratethys (Popov et al., 2004). Paleogene and Miocene formations accumulated in marine and brackish realms until the Maeotian, when sectors of the riparian areas became emerged and large herbivores occurred on these terrestrial environments (Mihăilă, 1971). In Pontian, the marine and brackish waters were replaced by almost freshwater environments (Vasiliev et al., 2005).

At the beginning of the Pliocene (Dacian), this area was covered by the Dacian Lake, where a large diversity of mollusks lived, many of them endemic. In the late Pliocene (Romanian), the lake vanished in the Colțești area, replaced by a fluvial system with associated coal forming swamps. Various land vertebrates lived in these environments. In the Quaternary, this area was completely emerged.

Large herbivores, including rhinoceros remains, were also found mainly due to the coal mining works in the numerous open pits of this region. The vertebrate remains from Colțești were unearthed from yellowish, medium, and rough sand and sandstone concretions, belonging to the Tetoiu Formation (Andreescu et al., 2011 and Andreescu et al., 2013), in particular to the so-called T-1 complex. The fossil mammals delivered by the Tetoiu Fm. are to be considered among the richest, most diverse and interesting ones in the early Pleistocene of the eastern Paratethys. The sequences of the early Pleistocene deposits, represented by the Tetoiu Fm., containing several fossiliferous levels rich in larger mammals, are to be found at Tetoiu in the middle valley of the Olteţ River. Paleontological investigations carried out in this area have detected a number of mammal-bearing deposits that can be assigned to three successive fossiliferous complexes named T1, T2, and T3 (Feru et al., 1983 and Rădulescu and Samson, 1990), corresponding to: MNQ17 (final part) or first half of MNQ18, first half of MNQ19 and second half of MNQ19, respectively (Andreescu et al., 2011: fig. 4; Andreescu et al., 2013: fig. 4; Nomade et al., 2014: fig. 7). The fossil localities from T-1 have yielded the selected following mammalian species: Mammuthus meridionalis, Trogontherium dacicum, S. etruscus, Stephanorhinus sp., Allohippus athanasiui (= Equus stenonis or E. sussenbornensis see Forsten, 1999), Eucladoceros sp., Pliotragus ardeus, Nyctereutes megamastoides, Lynx issiodorensis, Mitilanotherium inexpectatum, Ursus etruscus, Pliocrocuta perrieri, Homotherium crenatidens, Megantereon cultridens, Manis cf. hungarica (Bolomey, 1965, Feru et al., 1983, Rădulescu and Samson, 1990, Rădulescu and Samson, 2001, Samson and Rǎdulescu, 1963, Samson and Rǎdulescu, 1966 and Samson and Rǎdulescu, 1973). An important biostratigraphic event in the Tetoiu area is indicated at ∼1.7–1.8 Ma by the possible appearance of a Homo erectus lineage documented through the discovery of a few flint pebbles (Rădulescu and Samson, 1990, Rădulescu and Samson, 1995 and Samson and Rǎdulescu, 1963).

 Material and methods

The revised Quaternary time scale of Gibbard et al. (2010) for chronological references is used in this text. The bottom and top boundaries of the Pliocene are placed at 5.4 Ma and 2.6 Ma, respectively.

The postcranial elements analyzed here include a left calcaneus and second, third, and fourth metatarsals belonging to a same individual. The material is currently housed at OMNSC. The specimens were morphologically compared with the rhinoceros material collected at several Pliocene and early Pleistocene localities of Eurasia and housed in several institutions, as well as with published data (Appendix 1). The anatomical descriptions follow Guérin (1980) and Antoine (2002), whereas the morphometric approach follows Guérin (1980). The comparative tables are reported in Appendix 1.

Institutional abbreviations

OMNSC, Oltenia Museum Natural Sciences Department Craiova; HNHM, Magyar Természettudományi Múzeum (Hungarian Natural History Museum), Budapest, Hungary; IGF, Museo di Storia Naturale, sezione di Geologia e Paleontologia, Florence, Italy; MNCN, Museo Nacional de Ciencias Naturales, Madrid, Spain; MPLBP, Museo di Paleontologia Luigi Boldrini di Pietrafitta, Perugia, Italy; MPUR, Museo di Paleontologia, Sapienza, Università di Roma, Rome, Italy; NHMW, Naturhistorisches Museum, Wien, Austria; NMB, Naturhistorisches Museum, Basel, Switzerland.

Other abbreviations

ca, circa (about); L, maximal length; M, upper molar; MC, metacarpal; MNQ, mammal zones; MT, metatarsal.

 Systematic paleontology

Order Perissodactyla Owen, 1848

Family Rhinocerotidae Gray, 1821

Tribe Rhinocerotini Gray, 1821

Genus Stephanorhinus Kretzoi, 1942

Type species: Stephanorhinus etruscus (Falconer, 1868).

Note. The debate on the taxonomic status of the European Pleistocene fossil rhinoceroses has been recently summarized by Pandolfi and Tagliacozzo (2015). Nevertheless, some issues concerning the paraphyly of the genus have been raised. Geraads (1988) suggested that Stephanorhinus is a paraphyletic group and Coelodonta represents the name for the monophyletic group that includes the species characterized by loss of anterior teeth, firmly closed external auditory pseudomeatus and backward-movement of the nasal notch. Heissig (1999) suggested a paraphyly of Stephanorhinus, recognizing two evolutionary lineages, one from Dihoplus schleiermacheri to S. jeanvireti through ‘D’. megarhinus, and the other one from Stephanorhinus pikermiensis to S. etruscus and other species. The Pikermi rhinoceros was later included into Stephanorhinus also by Antoine and Saraç (2005). Recent paleogenomic studies on this genus, however, revealed that Stephanorhinus is strictly related to Coelodonta (Cappellini et al., 2019 and Kirillova et al., 2017). In the light of these results, the diagnostic characters of Stephanorhinus need to be re-discussed, as well as the validity of this genus (Cappellini et al., 2019). These issues, however, are beyond the contribution of this work and, in this paper, we consider Stephanorhinus as a monophyletic group as defined by diagnostic characters yet reported in past literature.

Stephanorhinus jeanvireti (Guérin, 1972)

Synonymy list: 1895 Rhinoceros etruscus var. astensis Sacco, 1–31, pl. I–IV; nomen oblitum.

2003 Stephanorhinus elatus (non Croizet and Jobert, 1828); Rădulescu et al., 3, 4, 8.

2010 Stephanorhinus elatus (non Croizet and Jobert, 1828); Rook and Martínez-Navarro, 136.

2019 Stephanorhinus elatus (non Croizet and Jobert, 1828); Ballatore and Breda, 205, tab. 2, in partim.

See Guérin (1972; in partim), Guérin and Tsoukala (2013; in partim) and Ballatore and Breda (2016; in partim) for a complete synonymy list.

Note: llatore and Breda (2016) recently suggested that S. jeanvireti (Guérin, 1972) is a junior synonym of S. elatus (Croizet and Jobert, 1828). Ballatore and Breda (2016) reported that the name S. elatus is not a nomen oblitum (being used in a few papers during the past century) and that S. jeanvireti should be replaced in force of ICZN rules. Nevertheless, the specimens of Rhinoceros elatus figured and described by Croizet and Jobert (1828) do not display diagnostic morphological features to distinguish them among the Pliocene and early Pleistocene rhinoceroses (including ‘D.’ megarhinus and ‘S.’ miguelcrusafonti). Ballatore and Breda (2016) briefly re-described those specimens but did not find exhaustive diagnostic characters. The anterior leg erected as a lectotype has been not collected in articulation and, except for the radius and humerus (but with some doubts), the bones reported by Croizet and Jobert (1828) cannot be confidently interpreted as belonging to the same individual (necessary to be included together as holotype/lectotype). The humerus and radius, which possibly constitute the articulated partial leg, are poorly preserved and do not display diagnostic characters (as also reported by Ballatore and Breda, 2016). In summary, the type specimens on which Rhinoceros elatus was established by Croizet and Jobert (1828) cannot be distinguished from other Pliocene and early Pleistocene taxa, and the lectotype erected by Ballatore and Breda (2016) is not properly defined. The name Rhinoceros elatus is here regarded as a nomen dubium and therefore S. jeanvireti is retained in order to maintain nomenclatural stability; furthermore, S. jeanvireti is based on well-preserved material such as skull and mandible. Despite this, some remains recovered posteriorly to Croizet and Jobert's work from Les Étouaires do actually correspond to S. jeanvireti, as it is reflected in the synonymy list (see also Pandolfi et al., 2017).

Type material: skull and mandible (No. Vt 627) described and figured by Guérin (1972), stored at Natural History Museum Basel.

Type locality and horizon: Vialette (Haute-Loire, France), late Pliocene.

Diagnosis: see Guérin, 1972 and Guérin, 1980 and Guérin and Tsoukala (2013).

Referred material: associated left calcaneus (OMNSC 44635) and left second (OMNSC 44640), third (OMNSC 44616), and fourth (OMNSC 44624) metatarsals, fitting together, evidencing that they belong to the same individual (Fig. S1).

Description and comparison: Calcaneus (44635). In lateral view, the plantar border of the bone is rather straight, the beak protrudes a little bit beyond the tuber calcanei, the proximal profile shows a marked concavity, and the posterior part of the tuber calcanei is higher and more convex than the anterior part (Fig. 2A).

In plantar view, the sustentaculum talii is perpendicular to the main axis of the bone, forming a right angle with the corpus of the bone (Fig. 2B). The articular surface located on the sustentaculum talii is wide, but separated from the latero-proximal articular surface and connected with the distal articular surface for the astragalus (Fig. 2C). In distal view, the latero-proximal articular surface for the astragalus is relatively broad, proximo-distally convex in the proximal half and concave in the distal one (Fig. 2D). The distal border of this surface extends distally, joining the distal articular surface for the astragalus. The latter is elongated, relatively high and connected distally with the articular surface for the cuboid. The latter is sub-trapezoidal in shape; it is latero-medially depressed, with a slightly convex posterior border. This surface extends proximally, almost reaching the sustentaculum talii, and it is well evident in plantar view.

The studied calcaneus is larger than those of S. etruscus (Appendix 1; Fig. 3); in this species, the tuber calcanei protrudes well over the beak, the posterior border of the bone is slightly concave, and the sustentaculum talii shows an obtuse angle with the corpus of the bone. The calcaneus of ‘S.’ miguelcrusafonti differs from 44635 by being less massive and smaller and by having a concave posterior profile of the bone. The calcanei of S. jeanvireti and ‘D.’ megarhinus are rather similar in morphology, with a wide overlapping in size (Fig. 3). The main diagnostic feature in the calcaneus seems to be the development of the sustentaculum talii; S. jeanvireti is characterized by a long sustentaculum talii with a medial border wider than in ‘D.’ megarhinus. This feature is evident in the calcaneus 44635. In plantar view, in S. jeanvireti and in the studied specimen, the lateral border is slightly concave, while in ‘D.’ megarhinus it is irregular.

MTII (44640). The bone is rather slender (Fig. 4A). In proximal view, the articular surface is damaged on its dorso-medial side, but it seems rather elliptical, with a concave lateral border. The proximal epiphysis is more developed dorso-plantarly than the proximal articular surface (Fig. 5C). In lateral view, two articular surfaces are present (Fig. 4B): the dorsal one is elliptical and proximo-distally elongated, whereas the plantar one (poorly preserved) is sub-trapezoidal, but higher than the dorsal one. In posterior view (Fig. 4C), a poorly preserved articular surface for the first cuneiform is present. The section of the diaphysis is rather rounded with a marked and wide posterior carena (Fig. 4 and Fig. 5).

The bone is larger than the MTIIs of S. etruscus and ‘S.’ miguelcrusafonti. The specimen differs from ‘D.’ megarhinus, in which the proximal articular surface reaches the dorsal border of the proximal epiphysis and the latter is not much higher than the plantar border. In ‘D.’ megarhinus, the section of the diaphysis is more irregular than that of the studied specimen. The morphology of the MT 44640 resembles in its features the MTIIs of S. jeanvireti (Guérin, 1972: fig. 18; Guérin, 1980: fig. 80).

MTIII (44616). In anterior view, the medial and lateral borders of the diaphysis are rather straight (Fig. 6A). In proximal view, the MTIII displays a rather convex anterior border of the proximal epiphysis; the plantar articular surface for the MTIV is partially visible (Fig. 5B). In lateral view (Fig. 6B), the dorsal articular surface for MTII is partially preserved and it seems to be trapezoidal, whereas the plantar articular surface for MTII is elliptical. These two surfaces are rather similar in size and are separated by a wide groove. In plantar view, the diaphysis is rather flat (Fig. 6C), the plantar tuberosity is well evident. In medial view (Fig. 6D), the proximal epiphysis is poorly preserved, but traces of two small articular surfaces are present, with the dorsal one larger than the plantar one. In this view, the plantar tuberosity is evident, but it is not much developed. The section of the diaphysis is sub-triangular with rounded angles, a rather flat plantar border and a medial border much narrower than the lateral one (Fig. 5E).

The proximal epiphysis in 44616 is longer than wide, contrary to the case of ‘D.’ megarhinus, and similarly to S. jeanvireti. The anterior border of the proximal epiphysis in 44616 displays a little concavity in the medial side, but this feature is less marked than in ‘D.’ megarhinus (even though some specimens display a little concavity in the middle of the anterior border, e.g., NMB Mp59) and it is usually absent in S. jeanvireti. Contrary to the studied specimen, the dorsal articular surface for MTIV is wider than in ‘D.’ megarhinus. The section of the diaphysis in ‘D.’ megarhinus is medio-laterally longer and dorso-plantarly narrower than that of the studied specimen and S. jeanvireti; in addition, the plantar border in ‘D.’ megarhinus is usually concave or slightly concave. The studied bone is longer than S. etruscus and ‘S.’ miguelcrusafonti (Fig. 7).

MTIV (44624). In anterior view, the two proximo-medial articular surfaces are partially evident (Fig. 8A). In lateral view, the proximal tuberosity is little developed (Fig. 8B). In proximal view, the articular surface is pentagonal, with relatively marked angles. The surface is concave medio-laterally. Its dorso-plantar length is similar to its medio-lateral one. The plantar border of the proximal articular surface is rather sinuous and the proximal epiphysis is a little more plantarly developed than the articular surface. In proximal view, the medial tuberosity, on which the plantar articular surface for MTIII is located, is well developed and much more medially projected than the proximal articular surface (Fig. 5A). In posterior view (Fig. 8C), the articular surface projects proximally over the rest of the proximal epiphysis. On the medial side of the bone (Fig. 8D), two articular surfaces for the MTIII are present. The dorsal one is poorly preserved, but seems to be less developed than the posterior one, which is rather elliptical. The section of the diaphysis is sub-rectangular in shape, with rounded angles and the medial border shorter than the lateral one (Fig. 5D).

The proximal articular surface is more irregular in shape and medio-distally longer in ‘D.’ megarhinus than in 44624 and S. jeanvireti. The section of the diaphysis is much more rounded in ‘D.’ megarhinus than in S. jeanvireti and the studied specimen. The MTIV 44624 is longer than in S. etruscus and ‘S.’ miguelcrusafonti.

 Discussion

Stephanorhinus jeanvireti seems to be a relatively rare taxon with respect to other European Plio-Pleistocene rhinoceroses (except for ‘S.’ miguelcrusafonti). The species was defined at Vialette (ca 3 Ma, MNQ16a), in France, but also recorded at Perrier–Les Étouaires (ca 2.78 Ma, MNQ16b; Pandolfi et al., 2017), Vincent and Desnes (both MNQ16) (Guérin, 1972 and Guérin, 1980). Stephanorhinus jeanvireti was reported in only one locality in Spain, Camp dels Ninots, with an age around 3.2 Ma (MNQ15-MNQ16 transition, Soler et al., 2012; Fig. 9). In Italy, it was recorded in several localities (MNQ16 or generally referred as Pliocene), in particular from the northern area: in Piedmont (Dusino, Villafranca d’Asti, Roatto), Emilia-Romagna (Monte Pulgnasco, Pradalbino, Monte Pastore, Monte San Pietro), and Tuscany (Montopoli, Capannoli) (Guérin, 1980, Pandolfi, 2013 and Simonelli, 1897 and references therein). In Germany, S. jeanvireti was recorded in the Hambach lignite mine (MNQ16) (Lacombat and Mörs, 2008) and scanty material from the Netherlands was doubtfully referred to this species (De Vos et al., 1998, Hooijer, 1981, Hooijer, 1985 and Scager et al., 2017); our revision confirms that the tooth remains are too fragmentary for a confident determination. In Slovakia, S. jeanvireti was documented at least at Hajnáčka (MNQ16a), Nová Vieska (MNQ16b or 17a), and Strekov (MNQ16a) (Ďurišová, 2004, Šujan et al., 2013 and Vlačiky et al., 2008; Pandolfi, pers. observ. at NHMW; Fig. 9), and in Bulgaria it was recorded at Musselievo (second half of MNQ15; Spassov, 2005). In Hungary, S. jeanvireti was present at Rákoscsaba (early Villafranchian), where it is documented by isolated teeth partially published by Pandolfi et al. (2015a) and by an unpublished mandible. In Russia, S. ex gr. jeanvireti was mentioned in the Khapry faunal complex from the Sea Azov Region by Bajgusheva et al. (2001), but this record has been reported as Stephanorhinus sp. or S. ex gr. megarhinus-kirchbergensis by Titov (2008). Recently, S. jeanvireti was reported in Greece at Milia (MNQ16; Guérin and Tsoukala, 2013). According to Lacombat and Mörs (2008), the species could be present at Kvabebi (Georgia), Udunga and Nizhnavodyanoy (Russia). The material from Georgia represents a large and massive rhinoceros and it is at present under review by one of us (LP). The rhinoceros from Udunga has been referred by Fukuchi et al. (2009) as S. megarhinus (recte ‘D.’ megarhinus, Pandolfi et al., 2016 and Pandolfi et al., 2015a), whereas the material from the Azov Region (Alekseeva, 1977 and Bajgusheva et al., 2001) should be deeply revised.

In Romania, Rădulescu and Samson (1985 and references therein), Codrea (2000) and Rădulescu et al. (2003) reported S. jeanvireti (= Dicerorhinus elatus in Rădulescu and Samson, 1995 or Stephanorhinus elatus in Rădulescu and Samson, 2001) in the Covrigi fauna (MNQ16a), Cernăteşti (MNQ16a), Iarăş 2 and Araci–Fântâna Fagului (early Gauss, MNQ16), Cernatu (early–middle Villafranchian) and S. cf. S. jeanvireti at Ilieni “Basin” (in fact, a sub-basin of the Bârsei Basin; early Villafranchian, MNQ16a). Most of the rhinoceros remains collected from these localities are represented by fragmentary and isolated bones.

Apart from the rhinoceros, the faunal assemblage recovered from Colțești includes Ursus etruscus (Stiucă et al., 2000), Pliotragus ardeus, Allohippus cf. athanasiui (= E. stenonis or E. sussenbornensis) and Eucladoceros sp. (Costin Rădulescu, personal communication to one of us, AP), and it is correlated with the faunal assemblage of the T-1 faunal complex (Andreescu et al., 2011: fig. 4; Andreescu et al., 2013: fig. 4 and references therein), corresponding to the final part of MNQ17 or the first half of MNQ18. Thus, the Colțești record represents the youngest occurrence of S. jeanvireti in Eurasia.

Stephanorhinus jeanvireti is usually associated with Anancus arvernensis and Tapirus arvernensis, in a woodland environment with humid climate (Guérin, 1980 and Szabó et al., 2017), related to MNQ16 localities (Fig. 9). At Colțești, the presence of Pliotragus and an equid would be indicative of a savannah with small forested interleaved areas, whereas Ursus etruscus could indicate the proximity of higher altitudes.

According to Heissig (1996), S. jeanvireti derived from ‘D.’ megarhinus and replaced it during the late Pliocene. The two species have been never recorded from coeval localities. The cluster analysis (UPGMA) performed by Pandolfi and Maiorino (2016: fig. 9) placed the cranial shape of S. jeanvireti in an intermediate stage between ‘D.’ megarhinus and S. kirchbergensis, whereas the upper-tooth row shape clustered together with that of ‘D.’ megarhinus (Pandolfi and Maiorino, 2016: fig. 9B), with D. sumatrensis as their closest relative. The presence of vestigial incisors, which was considered as a distinctive character between ‘D.’ megarhinus and S. jeanvireti, has been recently verified also on the latter species (Guérin and Tsoukala, 2013). Guérin (1980) suggested an Asian origin of S. jeanvireti, but Pliocene records of South-East Asia and East Asia are at present not exhaustive enough to support this hypothesis.

 Conclusion

The study of the rhinoceros material collected from the locality of Colțești represents a contribution towards a revision of the mammal faunas from Romania.

The morphological comparison revealed that the studied specimens can be assigned to S. jeanvireti. The western limit of the geographic distribution of this species is at present located at the northeasternmost part of the Iberian Peninsula (Camp dels Ninots, Girona) whereas the Romanian findings represent the eastern limit. The biochronological distribution of S. jeanvireti is mainly confined to the MNQ16 (early and early middle Villafranchian; Rook and Martínez-Navarro, 2010), and its first occurrence is doubtfully reported in late MNQ15 faunas (Spassov, 2005). The record of Colțești (MNQ17/MNQ18) represents, at present, the last occurrence of this taxon in Eurasia (Fig. 9). A better understanding of late Pliocene and earliest Pleistocene western Asian and eastern European Rhinocerotidae (including the revision of the Balkan and Caucasian material) could deeply contribute to the investigation of the origin and phyletic affinities of S. jeanvireti, as well as of the temporal and spatial distribution of fossil rhinoceroses.

 Acknowledgments

We are grateful to S. Mayda for his useful suggestions; we are greatly indebted to E. Cerdeño for her constructive criticism and helpful comments on the manuscript. L.P. thanks E. Cioppi (IGF), P. Pérez Dios (MNCN), M.C. De Angelis (MPLBP), R. Manni (MPUR), and L. Costeur (NMB) for their help and assistance during his visits to the fossil rhinoceros collections. L.P. thanks the European Community Research Infrastructure Action, EU-SYNTHESYS projects AT-TAF-2550, DE-TAF-3049, GB-TAF-2825, HU-TAF-3593, ES-TAF-2997; part of this research received support from the SYNTHESYS Project http://www.synthesys.info/ which is financed by European Community Research Infrastructure Action under the FP7 “Capacities” Program. VAC thanks Babeș-Bolyai University of Cluj-Napoca (abbreviated UBB) grants AGC: 30850, 30851, 30852, 31509, 32164. We thank Marian Bordeianu (UBB) for his help in drawing Fig. 1 on the computer, and Nicolae Trif (UBB and Museum Brukenthal Sibiu) for the photos.

 Comparative tables of the rhinoceros material from Colțești (southwestern Romania)

Calcaneum Locality Museum Specimen Reference Range value Greatest length Minimum posterior transversal diameter Greatest width of the tuber Greatest breadth at sustentaculum tali Colțești OMNSC 44635 This paper 138 40.2 63 86.5 S. etruscus Pietrafitta MPLBP Pandolfi et al., 2017 116.72 38.27 46.02 73.24 S. etruscus Pietrafitta MPLBP Pandolfi et al., 2017 110.9 36.1 40.81 69.38 S. etruscus Valdarno NMB Pandolfi et al., 2017 119.2 25.4 39.3 61.9 S. etruscus Valdarno MNCN Pandolfi et al., 2017 116.28 27.6 39.5 60 S. etruscus Valdarno IGF Pandolfi et al., 2017 121.47 31.31 44 S. etruscus Valdarno IGF Pandolfi et al., 2017 32.71 45.9 63 S. etruscus Valdarno IGF Pandolfi et al., 2017 108.17 32.28 40.3 69.9 S. etruscus Valdarno IGF Pandolfi et al., 2017 122.86 35.13 44.8 66.08 S. etruscus Olivola Val di Magra IGF Pandolfi et al., 2017 120 30 44 S. etruscus Olivola Val di Magra IGF Pandolfi et al., 2017 31 41 74.8 S. etruscus Olivola Val di Magra IGF Pandolfi et al., 2017 120 31.3 46 68 S. etruscus Capitone MPUR Pandolfi et al., 2017 119.8 34.5 45 72 S. etruscus Capitone MPUR Pandolfi et al., 2017 119 37 46 73 S. etruscus Venta Micena Santafè-Llopis and Casanovas-Cladellas, 1987 45 S. etruscus El Rincon MNCN Pandolfi et al., 2017 122.62 33.5 76.19 S. etruscus Les Etouaries NMB Pandolfi et al., 2017 121.7 35.6 75.2 S. etruscus Les Etouaries NMB Pandolfi et al., 2017 39.2 S. etruscus Les Etouaries NMB Pandolfi et al., 2017 117.2 35.4 48 74.7 S. etruscus Senèze Lacombat, 2005 121.36 37.65 44.38 72.77 S. etruscus Senèze Lacombat, 2005 120.87 39.16 46.19 73.11 S. etruscus Saint Vallier Guérin, 2004 120.5 34.5 44 75.5 S. etruscus Saint Vallier Guérin, 2004 123 33 45 72 S. etruscus Guérin, 1980 Min 109 31 39.5 60 S. etruscus Guérin, 1980 Med 118.72 34.98 43.89 67.61 S. etruscus Guérin, 1980 Max 130 39 49 75 S. jeanvireti Milia Guérin and Tsoukala, 2013 139 43 56 80 S. jeanvireti Guérin, 1980 Min 128 35 48 75 S. jeanvireti Guérin, 1980 Med 139.08 38.15 53.61 81.36 S. jeanvireti Guérin, 1980 Max 149.5 43.5 61 89 “D.” megarhinus Guérin, 1980 Min 133 38.5 48 72 “D.” megarhinus Guérin, 1980 Med 148.54 43.72 56.14 84.4 “D.” megarhinus Guérin, 1980 Max 163 51 63 98 “S.” miguelcrusafonti Guérin, 1980 128.5 49 MTII Locality Museum Specimen Reference Range value Length Breadth of the proximal epiphysis Depth of the proximal epiphysis Breadth of the shaft Depth of the shaft Breadth of the distal epiphysis Breadth of the distal articular surface Depth of the distal epiphysis Colțești OMNSC 44640 This paper 187 33 45.3 26 23.5 42 34.3 41 S. etruscus Montopoli IGF Pandolfi et al., 2017 27.9 35.85 26 21.1 S. etruscus Valdarno NMB Pandolfi et al., 2017 193.8 33.1 17.4 48.1 37 33.1 S. etruscus Valdarno IGF Pandolfi et al., 2017 29.3 37.1 30.7 S. etruscus Valdarno IGF Pandolfi et al., 2017 27 39 30.5 S. etruscus Valdarno IGF Pandolfi et al., 2017 27.19 24.35 37.71 30.25 36.35 S. etruscus Valdarno IGF Pandolfi et al., 2017 145.71 29.74 36.15 24.42 20.85 33.67 26.12 32.66 S. etruscus Valdarno IGF Pandolfi et al., 2017 148 28.9 23.3 23.51 36.04 29.87 35.17 S. etruscus Capitone MPUR Pandolfi et al., 2017 156 27 40 22.5 23.5 34.7 34 S. etruscus La Puebla de Valverde MNCN Pandolfi et al., 2017 25 41 S. etruscus La Puebla de Valverde Guérin and Heintz, 1971 25 42 S. etruscus Saint Vallier NMB Pandolfi et al., 2017 28.08 39.44 22.78 24.97 S. etruscus Guérin, 1980 Min 145 21 32.5 22.5 21 34 31 32 S. etruscus Guérin, 1980 Med 155.41 27.1 37.9 25.91 24.38 37.1 32.87 35.15 S. etruscus Guérin, 1980 Max 162 31 42 29 27.5 41 35 39 S. jeanvireti Guérin, 1980 Min 179 30 45 22 23 39 36 41 S. jeanvireti Guérin, 1980 Med 182.43 35.12 46.21 30 28.81 43.07 38.72 43.44 S. jeanvireti Guérin, 1980 Max 189 40 50 33 34 46 40 47 “D.” megarhinus Guérin, 1980 Min 174.5 31.5 41 27 24 43 36.5 37 “D.” megarhinus Guérin, 1980 Med 182.92 35.66 47.56 33.93 26.56 46.92 40.71 42.64 “D.” megarhinus Guérin, 1980 Max 191 42.5 53 38.5 33 52.5 45 46 “D.” megarhinus Çalta Guérin and Sen, 1998 175.5 ca 43 ca 32 29 29.5 37.5 33 38.5 “S.” miguelcrusafonti Guérin and Santafé-Llopis, 1978 162 34.5 39 29.5 28.5 37 33 38.5 “S.” miguelcrusafonti Guérin and Santafé-Llopis, 1978 158.5 26.5 37.5 MTIII Locality Museum Specimen Reference Range value Length Breadth of the proximal epiphysis Depth of the proximal epiphysis Breadth of the shaft Depth of the shaft Breadth of the distal epiphysis Breadth of the distal articular surface Depth of the distal epiphysis Colțești OMNSC 44616 This paper 203 55.6 47 45.5 24.3 61.4 46.7 40 S. etruscus Valdarno IGF Pandolfi et al., 2017 182 51.5 42.5 55 42.9 S. etruscus Valdarno IGF Pandolfi et al., 2017 185 51.8 42.7 53 43.1 S. etruscus Valdarno IGF Pandolfi et al., 2017 182.5 47.96 44.02 42.28 23.11 53.77 41.3 37.57 S. etruscus Valdarno IGF Pandolfi et al., 2017 175.34 49.83 39.37 41.24 21.36 53.51 41.1 37.31 S. etruscus Capitone MPUR Pandolfi et al., 2017 182 47.5 39 25 51.75 45 S. etruscus Villanueva de Pitamo MNCN Pandolfi et al., 2017 152.18 40.36 32.92 16.68 41.01 36.66 27.71 S. etruscus La Puebla de Valverde Guérin and Heintz, 1971 51 46 S. etruscus La Puebla de Valverde Guérin and Heintz, 1971 39 24 49 40 32 S. etruscus Saint Vallier NMB Pandolfi et al., 2017 46.36 35.16 S. etruscus Saint Vallier NMB Pandolfi et al., 2017 49.11 40.46 40.3 22.8 S. etruscus Saint Vallier NMB Pandolfi et al., 2017 44.75 40.12 S. etruscus Várhegy HNHM Pandolfi et al., 2017 178 44 25 52 45.5 39 S. etruscus Guérin, 1980 Min 165 43 38 33.5 21.5 48 39 32 S. etruscus Guérin, 1980 Med 175.92 48.54 41.86 40.18 22.9 52.13 42.28 36.86 S. etruscus Guérin, 1980 Max 187 56 47 45 25 59 46 40 S. jeanvireti Milia Guérin and Tsoukala, 2013 200 58 48 51.5 27 59.5 50 46 S. jeanvireti Milia Guérin and Tsoukala, 2013 57.5 S. jeanvireti Guérin, 1980 Min 200 53 44 46 21 58 45 40 S. jeanvireti Guérin, 1980 Med 205.29 57 47.83 49.36 24.06 61.83 50.17 43.44 S. jeanvireti Guérin, 1980 Max 212 60.5 52 53 25 65.5 54 45 “D.” megarhinus Guérin, 1980 Min 190 53 42.5 45 22 57 46.5 37.5 “D.” megarhinus Guérin, 1980 Med 202.53 60.03 47.93 56.16 25.55 65.47 53.26 43.92 “D.” megarhinus Guérin, 1980 Max 213.5 65 55 63 34 72 59 48 “S.” miguelcrusafonti Layna Guérin and Santafé-Llopis, 1978 186 52 41 60 47.5 38.5 “S.” miguelcrusafonti Layna Guérin and Santafé-Llopis, 1978 187.5 51.5 45.5 25 58.5 47 40 “S ” miguelcrusafonti Layna Guérin and Santafé-Llopis, 1978 187 52 40.5 45 24.5 56 45 38 MTIV Locality Museum Specimen Reference Range value Length Breadth of the proximal epiphysis Depth of the proximal epiphysis Breadth of the shaft Depth of the shaft Breadth of the distal epiphysis Breadth of the distal articular surface Depth of the distal epiphysis Colțești OMNSC 44624 This paper 176.5 49 44.5 32 21.3 37.5 36 42 S. etruscus Capitone MPUR Pandolfi et al., 2017 145 37 34.5 23 24 34.5 37.5 S. etruscus Valdarno IGF Pandolfi et al., 2017 155 32 37.5 29 S. etruscus Valdarno IGF Pandolfi et al., 2017 154 37 S. etruscus Valdarno IGF Pandolfi et al., 2017 156.07 38.61 35.1 25.44 22.42 36.53 30.06 36.04 S. etruscus Valdarno IGF Pandolfi et al., 2017 38.77 35.18 29.24 19.5 S. etruscus Valdarno IGF Pandolfi et al., 2017 41.33 35.9 S. etruscus El Rincon MNCN Pandolfi et al., 2017 41.84 36.1 32.07 24.03 S. etruscus La Puebla de Valverde MNCN Pandolfi et al., 2017 42.44 35.01 S. etruscus La Puebla de Valverde Guérin and Heintz, 1971 153 40 35 24 25 S. etruscus Saint Vallier NMB Pandolfi et al., 2017 40.45 32.76 S. etruscus Saint Vallier NMB Pandolfi et al., 2017 38.96 33.8 S. etruscus Senèze NMB Pandolfi et al., 2017 165.6 33.3 19.9 40.8 37.9 S. etruscus Senèze Lacombat, 2005 151 40.92 35.37 28.3 23.34 32.34 30.2 35.37 S. etruscus Guérin, 1980 Min 146 30 33 22 20 29 28.5 33 S. etruscus Guérin, 1980 Med 153.69 37.58 36.32 26.25 24.03 33.32 32.1 35.93 S. etruscus Guérin, 1980 Max 163 42 41 31 28 41.5 35 39 S. jeanvireti Milia Guérin and Tsoukala, 2013 178.5 56 51.5 36.5 35.5 43 43 45 S. jeanvireti Hajnacka I Durisova, 2004 188 49 44.2 34 27 40 38.5 40.5 S. jeanvireti Hajnacka I Durisova, 2004 186 50 44 37 32 46 43 43 S. jeanvireti Guérin, 1980 Min 168 44 40.5 31 23 40 37 41 S. jeanvireti Guérin, 1980 Med 180.43 49 44.15 35 28.56 41.6 39.86 45 S. jeanvireti Guérin, 1980 Max 188.5 52 49 38 36 44 45 49.5 “D.” megarhinus Guérin, 1980 Min 164 42 38 32 22 39 35.5 40.5 “D.” megarhinus Guérin, 1980 Med 176.19 49.65 47.55 34.29 31.08 40.22 38.72 44.81 “D.” megarhinus Guérin, 1980 Max 182 55 53.5 37 35 42 42 47 “S.” miguelcrusafonti Layna MNCN This work 156.5 37 35 29 29.5 35 33.5 34.5 “S.” miguelcrusafonti Layna MNCN This work 38.5

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Location of the locality Colțești (southwestern Romania) on the geological map.

Situation de la localité de Colteşti (Sud-Ouest de la Roumanie) sur la carte géologique.

Stephanorhinus jeanvireti from Colțești, calcaneus: lateral view (A); plantar view (B); medial view (C); distal view (D). The scale bar equals 5 cm.

Stephanorhinus jeanvireti de Colteşti, calcaneum : vue latérale (A) ; vue plantaire (B) ; vue médiale (C) ; vue distale (D). La barre d’échelle représente 5 cm.

Scatterplot of greatest length (GL) and greatest width of the tuber calcanei (GWt) of the calcaneus from Colțești, S. etruscus, S. jeanvireti, ‘S.’ miguelcrusafonti and ‘D.’ megarhinus (data from the Appendix 1).

Nuage de points représentatifs de la plus grande longueur (GL) et de la plus grande largeur (GWt) du tubercule calcanéen du calcanéum de Colteşti, S. etruscus, S. jeanvireti, ‘S.’ miguelcrusafonti et ‘D.’ megarhinus (données de l’Annexe 1).

Stephanorhinus jeanvireti from Colțești, MTII: dorsal view (A); lateral view (B); plantar view (C); medial view (D). The scale bar equals 5 cm.

Stephanorhinus jeanvireti de Colteşti, MT II : vue dorsale (A) ; vue latérale (B) ; vue plantaire (C) ; vue médiale (D). La barre d’échelle représente 5 cm.

Stephanorhinus jeanvireti from Colțești, proximal articular surface of metatarsals in proximal view (A-C) and cross section of metatarsal diaphyses (D-F): MTII (A); MTIII (B,E); MTIV (C,F).

Stephanorhinus jeanvireti de Colteşti, vues proximale de métatarse (A-B) et section transversale de diaphyses de métatarse (D-F) : MTII (A) ; MTIII (B) ; MTIV (C).

Stephanorhinus jeanvireti from Colțești, MTIII: dorsal view (A); lateral view (B); plantar view (C); medial view (D). The scale bar equals 5 cm.

Stephanorhinus jeanvireti de Colteşti, MTIII : vue dorsale (A) ; vue latérale (B) ; vue plantaire (C) ; vue médiale (D). La barre d’échelle représente 5 cm.

Scatterplot of greatest length (GL) and width of the diaphysis (WD) of the MTIII from Colțești, S. etruscus, S. jeanvireti, ‘S.’ miguelcrusafonti and ‘D.’ megarhinus (data from Appendix 1).

Nuage de points représentatifs de la plus grande longueur (GL) et de la largeur (WD) de la diaphyse du MTIII de Colteşti. S. etruscus, S. jeanvireti, ‘S.’ miguelcrusafonti et ‘D’ megarhinus (données de l’Annexe 1).

Stephanorhinus jeanvireti from Colțești, MTIV: dorsal view (A); lateral view (B); plantar view (C); medial view (D). The scale bar equals 5 cm.

Stephanorhinus jeanvireti de Colteşti, MTIV : vue dorsale (A) ; vue latérale (B) ; vue plantaire (C) ; vue médiale (D). La barre d’échelle représente 5 cm.

Biochronological occurrence of Stephanorhinus jeanvireti from Colțești and chronological position of selected European localities (see text for details on localities and ages). Scheme modified from Rook et al. (2017). T-1 and T-2 refer to the first two T-faunal horizons reported by Andreescu et al. (2013).

Distribution biochronologique de Stephanorhinus jeanvireti de Colțești et position chronologique de localités européennes sélectionnées (voir le texte pour les localités et les âges). Schéma de Rook et al. (2017) modifié. T-1 et T-2 se réfèrent aux deux premiers horizons fauniques rapportés par Andreescu et al. (2013).