palevol comptes rendus 2021  20  29 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 : Audrina Neveu (Muséum national d’Histoire naturelle ; audrina.neveu@mnhn.fr) Révisions linguistiques des textes anglais / English language revisions : Kevin Padian (University of California at Berkeley) Rédacteurs associés / Associate editors (*, took charge of the editorial process of the article/a pris en charge le suivi éditorial de l’article) : Micropaléontologie/Micropalaeontology Maria Rose Petrizzo (Università di Milano, Milano) Paléobotanique/Palaeobotany Cyrille Prestianni (Royal Belgian Institute of Natural Sciences, Brussels) Métazoaires/Metazoa Annalisa Ferretti (Università di Modena e Reggio Emilia, Modena) Paléoichthyologie/Palaeoichthyology Philippe Janvier (Muséum national d’Histoire naturelle, Académie des sciences, Paris) Amniotes du Mésozoïque/Mesozoic amniotes Hans-Dieter Sues (Smithsonian National Museum of Natural History, Washington) Tortues/Turtles Juliana Sterli (CONICET, Museo Paleontológico Egidio Feruglio, Trelew) Lépidosauromorphes/Lepidosauromorphs Hussam Zaher (Universidade de São Paulo) Oiseaux/Birds Eric Buffetaut (CNRS, École Normale Supérieure, Paris) Paléomammalogie (mammifères de moyenne et grande taille)/Palaeomammalogy (large and mid-sized mammals) Lorenzo Rook* (Università degli Studi di Firenze, Firenze) Paléomammalogie (petits mammifères sauf Euarchontoglires)/Palaeomammalogy (small mammals except for Euarchontoglires) Robert Asher (Cambridge University, Cambridge) Paléomammalogie (Euarchontoglires)/Palaeomammalogy (Euarchontoglires) K. Christopher Beard (University of Kansas, Lawrence) Paléoanthropologie/Palaeoanthropology Roberto Macchiarelli (Université de Poitiers, Poitiers) Archéologie préhistorique/Prehistoric archaeology Marcel Otte (Université de Liège, Liège) Référés / Reviewers : https://sciencepress.mnhn.fr/fr/periodiques/comptes-rendus-palevol/referes-du-journal Couverture / Cover : Made from the Figures of the article. 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 Ancestral foxes at the gates of Europe: the Pliocene fox from Çalta-1 (Turkey) and their relationships with Asian and European Plio-Pleistocene foxes Saverio BARTOLINI-LUCENTI Earth Science Department, Paleo[Fab]Lab, University of Florence, Via La Pira 4, 50121 Firenze (Italy) and Natural History Museum, Geology and Paleontology Section, University of Florence, Via La Pira 4, 50121 Firenze (Italy) saverio.bartolinilucenti@unifi.it (corresponding author) Joan MADURELL-MALAPEIRA Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, c/ de les columnes s/n Campus de la UAB, Cerdanyola del Vallès, 08193 Barcelona (Spain) Submitted on 2 May 2020 | Accepted on 14 June 2020 | Published on 13 July 2021 urn:lsid:zoobank.org:pub:1ACE41D1-37C8-4110-ABDA-AD5F57DB74B8 Bartolini-Lucenti S. & Madurell-Malapeira J. 2021. — Ancestral foxes at the gates of Europe: the Pliocene fox from Çalta-1 (Turkey) and their relationships with Asian and European Plio-Pleistocene foxes. Comptes Rendus Palevol 20 (29): 619-626. https://doi.org/10.5852/cr-palevol2021v20a29 ABSTRACT The Pliocene record of genus Vulpes Frisch, 1775 in Eurasia is scarce, coming from few sparse locali- ties. The lack of a comprehensive and integrated revision led to the description of numerous different taxa, often only tentatively related to extant species but not with one another. Çalta-1 is an important Pliocene site located in the Anatolian region of Turkey, dated to 4.0 Ma. In the present review, we reappraise the interesting record of Vulpes galatica Ginsburg, 1998. Morphological and ­morphometric evidence suggests a strong similarity between this taxon and the early Late Pliocene V. beihaiensis KEY WORDS Qiu & Tedford, 1990, recovered from the Chinese Yushe Basin. Such evidence favors the parsimonious Carnivora, interpretation of synonymy between the two species, under the name V. beihaiensis. This hypothesis Vulpes, opens a new interpretation on the biogeography of the Pliocene-Early Pleistocene foxes of Eurasia. Pliocene, Eurasia, Vulpes beihaiensis links the Asian and European records, with its affinity to V. ­alopecoides (Del Cam- biogeography. pana, 1913) and, eventually, to the extant red fox (Vulpes vulpes (Linnaeus, 1758)). COMPTES RENDUS PALEVOL • 2021 • 20 (29) © Publications scientifiques du Muséum et/and Académie des sciences, Paris. www.cr-palevol.fr 619 Bartolini-Lucenti S. & Madurell-Malapeira J. RÉSUMÉ Anciens renards aux portes de l’Europe : le renard pliocène de Calta-1 (Turquie) et ses relations avec les renards européens et asiatiques du Plio-Pléistocène. Le signalement pliocène du genre Vulpes Frisch, 1775 en Eurasie est peu abondant, du fait de quelques localités éparses. L’absence d’une révision intégrée et exhaustive a conduit à la description de nombre de taxons souvent reliés, à titre d’essai, à des espèces actuelles, mais non l’un à l’autre. Calta-1 est un important site pliocène localisé dans la région anatolienne de Turquie, daté de 4 Ma. Dans cet article, nous réexaminons l’intéressant signalement de Vulpes galatica Ginsburg, 1998. Des preuves morphologiques et morphométriques suggèrent une importante similitude entre ce taxon et V. ­beihaiensis Qiu & Tedford, 1990 du début du Pliocène supérieur, récolté dans le bassin d’Yushe, en MOTS CLÉS Chine. De telles preuves plaident en faveur de l’interprétation parcimonieuse d’une synonymie entre Carnivora, les deux espèces, sous le nom de V. beihaiensis. Cette hypothèse permet une nouvelle interprétation Vulpes, des renards d’Eurasie au Plio-Pléistocène-Pléistocène inférieur. Vulpes beihaiensis relie les registres Pliocène, Eurasie, européens et asiatiques, avec son affinité à V. alopecoides (Del Campana, 1913) et, éventuellement, biogéographie. au renard rouge actuel (Vulpes vulpes (Linnaeus, 1758)). INTRODUCTION chikushanensis Young, 1930 from several Early-Middle Pleis- tocene Asian localities, e.g. Longdan (Qiu et al. 2004); Vulpes Fossil record of the genus Vulpes Frisch, 1775 is often frag- galatica Ginsburg, 1998 from the Early Pliocene site of Çalta-1 mentary and scattered, making it difficult to reconstruct (c. 4.0 Ma; Turkey; Fig. 1); and the hypercarnivorous and phylogenetic affinities among them, which often leads to large-sized Vulpes qiuzhudingi Wang, Tseng, Li, Takeuchi & the proliferation of numerous specific names with prob- Xie, 2014 from the Himalayan-Tibetan Plateau (China, lematic taxonomy. The European Plio-Pleistocene record c. 5.1-3.6 Ma, Wang et al. 2014; Fig. 1). The relationships was historically characterized by the existence of three taxa: between these extinct species are currently unclear, although Vulpes alopecoides (Del Campana, 1913), Vulpes praeglacialis some hypotheses have been made. Qiu et al. (2004) suggested (­Kormos, 1932) and V. praecorsac Kormos, 1932. Additionally, a link between the Odessa sample and V. chikushanensis. The Spassov (2000) ascribed to Vulpes sp. a single right P4 from Chinese V. ­beihaiensis is considered the closest relative of the Pliocene site of Musselievo (Bulgaria, MN15; Fig. 1). The V. corsac (Qiu & Tedford 1990), whereas V. qiuzhudingi might first is a typical form of the Late Pliocene-Early Pleistocene, be the ancestor of V. lagopus (Wang et al. 2014). Ginsburg whereas the latter two species are described from the Early (1998) described V. galatica (note that “galaticus” violates the Pleistocene, although the sample from Odessa Catacombs International Code on Zoological Nomenclature, 1999, as (Ukraine, MN15, Early Pliocene; Fig. 1) was referred to reported in Bartolini-Lucenti & Madurell-Malapeira 2020) as V. praecorsac (Odintzov 1965). The virtual absence of clear a highly carnivorous species, the earliest member of the genus diagnostic features between these three similar-sized species has Vulpes and the ancestor of modern V. vulpes and V. lagopus. led to confused taxonomy of samples from Western European According to this view, V. alopecoides would represent the localities. Generally, older samples were ascribed to V. alope- intermediate, both ecologically and phylogenetically, between coides, whereas younger large-sized remains were attributed to V. galatica and V. vulpes (Ginsburg 1998). In the Ginsburg V. praeglacialis and small-sized ones to V. praecorsac. Recently, original publication (Ginsburg 1998), V. alopecoides was the Bartolini-Lucenti & Madurell-Malapeira (2020) proposed to only fossil species considered as a comparison. synonymize the Early Pleistocene samples under the specific Here, we restudy the material of V. galatica from Çalta-1 in name of V. alopecoides, based on the low interspecific variance the light of the diversity of Plio-Pleistocene Vulpes. The position of the fossil compared to the elevated degree of intraspecific of this site, both in time and space (see the following section), variability typical of extant foxes [Vulpes corsac (Linnaeus, and the systematic position of this taxon might offer crucial 1768), Gimranov et al. 2015; Gimranov 2017; Vulpes lagopus insights on the evolution of Vulpes species in the Pliocene (Linnaeus, 1758), Daitch & Guralnick 2007; Szuma 2008a; and Early Pleistocene times, as the first well-chronologically Vulpes vulpes (Linnaeus, 1758), Szuma 2008b, 2008c]. This constrained record of Vulpes in Eurasia. with the notable exception of V. “praecorsac” from Odessa, whose features contrast significantly with the morphologies of V. alopecoides, therefore the former taxon could not be GEOLOGICAL included in alopecoid fox (Bartolini-­Lucenti & Madurell-­ AND PALEONTOLOGICAL SETTINGS Malapeira 2020). The Plio-Pleistocene Asian record includes four species: the Çalta-1 is an important Early Pliocene site located in Cen- small-sized Early Pliocene Vulpes beihaiensis Qiu & ­Tedford, tral Anatolia, northwest of Ankara (Turkey; Fig. 1), in the 1990 from Yushe Basin sites (c. 3.3-3.0 Ma; Fig. 1); Vulpes Kazan Basin (Lunkka et al. 1998). The rich record of verte- 620 COMPTES RENDUS PALEVOL • 2021 • 20 (29) Ancestral foxes at the gates of Europe Odessa Catacombs Musselievo Kvabebi Yushe Basin Çalta-1 Kunlun Basin Zanda Basin Zanclean Piacenzian V. alopecoides V. galatica V. qiuzhudingi V. beihaiensis V. praecorsac Vulpes sp. Fig. 1. — Geographic localization of the Pliocene localities with Vulpes Frisch, 1775 record. brates recovered from this site was described in an extensive MG-GNM, MHNL, MNHN, UCBL) and inspected all monography published in Geodiversitas (Sen 1998). The fossil-­ the relevant literature on Eurasian fossil Vulpes (Odintzov bearing succession of Çalta-1 is located in the Çalta member, 1965; Qiu & Tedford 1990; Qiu et al. 2004; Wang et al. the youngest of the Sinap Formation (Lunkka et al. 1998, 2014; Bartolini-Lucenti 2021). Fossil species considered as 2003). A second site, named Çalta-2, is related to the upper comparison include V. alopecoides (sensu Bartolini-Lucenti & part of the Kavakdere member (underlying Çalta member in Madurell-­Malapeira 2020) from selected site of Europe the scheme of Lunkka et al. 1998; Sen & Saraç 2018). The (i.e., the Pliocene record of Kvabebi, Georgia; and the Early Sinap Formation is characterized by fluviolacustrine depos- Pleistocene one of Pirro Nord, Italy); V. beihaiensis from its mainly characterized by the alternance of conglomerates, Zhaozhuang and Zhangwagou (Yushe Basin, China; Qiu & sandstone and mudstone of alluvial system. The Çalta mem- Tedford 1990); V. qiuzhudingi from the Himalayan-Tibetan ber, at the top of the succession, is composed of floodplain Plateau (Zanda and Kunlun Pass Basins, China, Wang et al. deposits, represented by massive mudstones, channeled by 2014); V. “praecorsac” from Odessa Catacombs (Ukraine; thin conglomerates and sandstones. Upper portion of these Odintzov 1965); Vulpes sp. from Musselievo (Bulgaria; massive mudstones contained the fossil specimens (Lunkka Spassov 2000). Extant specimens of V. corsac, V. lagopus and et al. 1998). Along with a rich associated fauna (Sen 1998), V. ­vulpes from the AMNH, HNHM, and MZUF were used five carnivoran taxa were reported by Ginsburg (1998) from for morphological and morphometrical comparisons. Log ratio this site: the hyenid Chasmaporthetes kani anatolicus Ginsburg, diagrams (Simpson 1941; Simpson et al. 1960) on selected 1998; the felids Machairodus giganteus (Wagner, 1848) (syn- cranial and dental variables were used to assess the degree of onym of Amphimachairodus giganteus following Beaumont, similarity or difference between Vulpes from Çalta-1 other 1978) and Lynx issiodorensis (Croitzer & Jobert, 1828); and fossil and extant species. the canids Nyctereutes donnezani (Depéret, 1890) and V. gala- tica. Chronologically the site is related to the late Ruscinian Abbreviations (MN15), c. 4 Ma (Sen 1998; Bernor & Sen 2017). Institutions AMNH American Museum of Natural History, New York; DST-UNIFI Earth Science Department of the University of Florence, Florence; MATERIAL AND METHODS HNHM Hungarian Natural History Museum, Budapest; IGF Geological and Paleontological Section of the Natural This study is based on the comparative morphological and History Museum of the University of Florence, Florence; morphometric analyses of Vulpes galatica from Çalta-1 (Turkey) MG-GNM S. Janashia Museum of Georgia, Georgian National in comparison to other Pliocene to Early Pleistocene species Museum, Tbilisi; of Eurasia. The studied fossils come from the collections of MGPT-PU Geology and Paleontology Museum of the University of Turin, Turin; the MNHN (see abbreviations below). As comparative fossil MZUF La Specola, Zoological Section of the Natural History material, we studied the collections of several European and Museum of the University of Florence, Florence; Chinese localities (AMNH, DST-UNIFI, HNHM, IGF, UCBL Université Claude-Bernard Lyon-1, Lyon. COMPTES RENDUS PALEVOL • 2021 • 20 (29) 621 Bartolini-Lucenti S. & Madurell-Malapeira J. Table 1. — Measurements of the Vulpes beihaiensis Qiu & Tedford, 1990 from (MNHN.F.ACA-293b) and right i3-m3 (MNHN.F.ACA-293a); Çalta (Turkey). Abbreviations: H, height; L, greatest mesiodistal length in case MNHN.F.ACA-302, right astragalus. of teeth and rostrocaudal one in case of the astragalus; Mm1H, mandibular height distal to the m1; Mm1W, mandibular width at level of the m1; Mp4H, man- Measurements. — See Table 1. dibular height distal to the p4; W, buccolingual width. Description Specimens L W H Upper teeth MNHN.F.ACA-293 c 5.2 3.7 – The P4 is rather stout, with a large protocone a high paracone p1 3.2 1.9 – and modestly elongated metastyle, slightly shorter mesiodis- p2 6.5 2.4 – tally compared to the paracone. The protocone is advanced p3 7.5 2.7 – p4 8.3 3.3 – compared to the mesial margin of the P4. A cingulum bounds m1 11.9 4.8 – the lingual side of the tooth and another one is evident on trm1 7.9 – – the mesially, reaching the buccal margin of the paracone, tdm1 – 4.5 – m2 5.5 4.3 – at level of the feeble parastyle. The cingulum is prominent m3 3.1 2.7 – around the protocone. Mp4H – – 11.0 Mm1H – – 11.6 Mm1W – 5.6 – Mandible The mandible corpus is shallow with a scaphoid ventral MNHN.F.ACA-357 P4 14.3 6.0 – margin. The mesial mental foramen is located under the p1 MNHN.F.ACA-302 astragalus 22.9 – – and a smaller one under p3 in MNHN.F.ACA-293a whereas MNHN.F.ACA-293b apparently has only the mesial one. The right mandible is broken distal to the m3 whereas the left one Anatomy at level of the distal root of m1. c lower canine; M upper molar; Lower teeth m lower molar; P upper molar; Incisors are reduced, flattened teeth. The canine is buccolingually p lower molar. compressed but rather stout. The diastema between c1 and p1 is considerably wide. Premolars have upright protoconids Measurements and enlarged distal portions compared to the mesial ones, in L length; lateral and occlusal views. The p1 is monocuspid with a distal H height; cingulid. The p2 does not have distal accessory cuspulids. The W width. p3 and p4 display high, sharp and pointed protoconids. On the distal side of the p3 protoconid, a feeble accessory cuspulid is SYSTEMATIC PALAEONTOLOGY visible. Distally, a cingulid arises from the margin of the tooth. The p4 possesses a mesial shelf-like morphology with an acces- Order CARNIVORA Bowdich, 1821 sory cuspulid. Distal to the protocone MNHN.F.ACA-293a Family Canidae Fisher, 1817 has two cuspulids, whereas in MNHN.F.ACA-293b there is a Tribe Vulpini Hemprich & Ehrenberg, 1832 single larger one. The distal cingulid is fairly marked. The m1 Genus Vulpes Frisch, 1775 paraconid is considerably lower than the p4 protoconid. The m1 protoconid is high, sharp and stout, in buccal view. The Vulpes beihaiensis Qiu & Tedford, 1990 metaconid is relatively reduced and attached to the protoconid. (Fig. 2) Although smaller than the hypoconid, the entoconid is not so reduced. Talonid basin is round and rather wide, partially Vulpes beihaiensis Qiu & Tedford, 1990: 28. closed lingually by an accessory cuspulid, mesial to the entoco- Vulpes galaticus Ginsburg, 1998: 380-382, figs 1-2. — Sen & Saraç nid, and distally by a prominent cingulid. The later continues 2018: 536. on the buccal side, reaching the level of the hypoflexid. The m2 is rectangular in occlusal view. The protoconid and the Vulpes galatica – Bartolini-Lucenti & Madurell-Malapeira 2020: 2. metaconid are similar in size. On the talonid the hypoconid is Holotype. — AMNH F:AM 97062, cranium laterally compressed large but the entoconid is not reduced. The buccal cingulid is in Qiu & Tedford (1990). prominently and enlarged till the distal side of the tooth. The m3 is oval-shaped, with two equal-sized cuspulids and a cin- Type locality. — Zhanwagou (Yushe Basin, Shanxi, China). gulid bounding the distal margin. Chronostratigraphic distribution. — Early Pliocene (c. 4.0-3.0 Ma) Geographic distribution. — Asian continent, from Central DISCUSSION China to Turkey. Referred material. — MNHN.F.ACA-357, left P4 (L: 14.3 mm; W: In the original description by Ginsburg (1998), a P3 and M2 6.0 mm); MNHN.F.ACA-293, mandibular corpora with left ­i1-m1 were included in the sample. Nevertheless, their m ­ orphology 622 COMPTES RENDUS PALEVOL • 2021 • 20 (29) Ancestral foxes at the gates of Europe A D B E C F G H I Fig. 2. — Vulpes beihaiensis Qiu & Tedford, 1990 from Çalta-1. A-F, MNHN.F.ACA-293, mandibles: A-C, right hemimandible in buccal (A), lingual (B) and ­occlusal (C) views; D-F, left hemimandible in buccal (D), lingual (E) and occlusal (F) views. G-H, MNHN.F.ACA-357, left P4 in buccal (G) and occlusal (H) views. I, MNHN.F.ACA-302, right astragalus in proximal view. Scale bar: 1 cm. (e.g. lobed buccal cingulum on M2) does not fit with g­ eneral one Some fox remains were also unearthed from the western of Vulpes, probably belonging to a different canid (­Nyctereutes Asian site of Kuruksay (Tajikistan; MN17; Sharapov 1986). Temminck, 1838 or other). We therefore excluded them from These former dentognathic remains, ascribed to Vulpes aff. cor- the sample here described. sac, differs from the Çalta-1 record for several features: the proportionally longer lower premolars, with more prominent Comparison with Asian Plio-Pleistocene Vulpes distal cingulid. Çalta-1 specimen possesses a reduced mesial In dental morphology, the specimens from Çalta-1 do not differ portion of the p4, whereas in Kuruksay ones such portion greatly from Yushe Basin, as visible Fig. 3. There are only slight is evident. Some other differences worth to be mentioned differences, among which: 1) V. beihaiensis from Yushe Basin has between the Kuruksay taxon and Çalta-1 specimens are: the a more compressed buccolingually P4 at the level of the metastyle larger m1 protoconid and a stouter hypoconid, in buccal (Fig. 3A-C); 2) a cingulum does not girdle the P4 protocone; view, in the former compared to the latter and the metaconid 3) more evident P4 parastyle; 4) more evident distal cingulid is more developed in Kuruksay taxon compared to Çalta-1. on the lower premolars (Fig. 3I-M); and 5) generally lower m1 protoconid (although F:AM 97071 has the same morphology Comparison with European Pliocene of Çalta-1 specimens). In size and dental proportions, the two and Early Pleistocene Vulpes taxa differ only slightly (Fig. 4). On the contrary, the hypercar- The P4 from the Pliocene Bulgarian site of Musselievo differs nivorous V. qiuzhudingi greatly differs from Çalta-1 Vulpes in considerably from that of Çalta-1. For instance, the P4 of the many respects, morphological and metrical ones (Figs 3; 4) such former is buccolingually compressed, with a longer paracone as: 1) the corpus of V. qiuzhudingi is considerably deeper and compared to metastyle; the P4 protocone lies at the level stouter compared to Vulpes from Çalta-1; 2) the p2 protoconid of the mesial margin of the tooth; and a prominent buccal of V. qiuzhudingi is lowered compared to the upright one of cingulid is visible (Fig. 3A and D). In size, it is considerably Vulpes from ­Çalta-1; 3) V. qiuzhudingi does not show the high smaller and its proportions resemble those of V. “praecorsac” m1 protoconid (Fig. 3I, J, T, U); 4) the hypercarnivorous char- from Odessa (Fig. 4). acteristics of V. qiuzhudingi (e.g. enlarged buccal cuspulid on Vulpes “praecorsac” in the sample from Odessa display m1-m2; reduced metaconid; cristid-like entoconid) cannot be evident differences as compared with Çalta-1 specimens, found in the specimens of Çalta-1 (Fig. 3I, J, T, U); and 5) the namely: 1) the P4 of the former is stouter compared to biometrical dimensions of V. qiuzhudingi are considerably larger Çalta-1, with a longer paracone compared to the metastyle as compared to the taxon from Çalta-1 and its saw-toothed and more prominent buccal cingulum (Fig. 3E); 2) the P4 pattern contrast with the latter (Fig. 4). protocone in V. “praecorsac” from Odessa is reduced in height COMPTES RENDUS PALEVOL • 2021 • 20 (29) 623 Bartolini-Lucenti S. & Madurell-Malapeira J. A B C D E F G H I J K L M N O P Q R S T U Fig. 3. — Comparison of dental morphologies (A-H, occlusal view of P4; I-U, occlusal morphology of lower teeth) of various Vulpes spp. from Eurasia: A, Vulpes Frisch, 1775 from Çalta-1 (Turkey), MNHN.F.ACA-357, left P4 (reversed); B, C, V. beihaiensis Qiu & Tedford, 1990 from Yushe Basin (China), AMNH F:AM 97062, right (B) and left reversed P4 (C); D, Vulpes sp. from Musselievo (Bulgaria; Spassov 2000), right P4; E, V. “praecorsac” Kormos, 1932 from Odessa Catacombs (Ukraine; Odintzov, 1965), O-7764, right P4; F-H, V. alopecoides (Del Campana, 1913) from Pirro Nord (Italy): F, PP789, left P4 (reversed); G, MGPT-PU104618, right P4; H, MGPT-PU104805, left P4 (reversed); I, J, Vulpes from Çalta-1: I, MNHN.F.ACA-293b, left m1-m2; J, MNHN.F.ACA-293a, right p4-m3 (reversed); K-M, V. beihaiensis from Yushe Basin (China): K, AMNH F:AM 97068, right m1-m3 (reversed); L, AMNH F:AM 97071, left m1 (reversed); M, AMNH F:AM 97070, left p4-m1; N-Q, V. alopecoides from Europe: N, MG-29-2013/461, right m1-m2 (reversed) from Kvabebi (Georgia); O, MGPT-PU106227, right m1 (reversed) from Pirro Nord (Italy); P, PN 28, left p4-m1 from Pirro Nord (Italy); Q, MGPT-PU104721, right m1 (reversed) from Pirro Nord (Italy); R, S, V. “praecorsac” from Odessa Catacombs (Ukraine, Odintzov 1965): R, O-1519, left m1-m2; S, O-1520, right p4-m1 (reversed); T, U, V. qiuzhudingi Wang, Tseng, Li, Takeuchi & Xie, 2014 from Himalayan-Tibetan Plateau (southwestern China): T, IVPP V19060, left m2 from IVPP locality KL0605, Kunlun Pass Basin (China); U, IVPP V18923, left m1 from Zanda Basin (China). Scale bar: 1 cm. (Fig. 3A, E); 3) the mandible V. “praecorsac” is stouter com- of V. ­alopecoides, suggesting a possible different attribution pared to that of Çalta-1; 4) on the p4, V. “praecorsac” possess of the specimens from Muhkai II. a developed shelf-like portion with an accessory cuspulid, In comparison with Early Pleistocene V. alopecoides from which is reduced in MNHN.F.ACA-293 (Fig. 3I, J, R, S); 5) Europe (sensu Bartolini-Lucenti & Madurell-Malapeira 2020), the m1 of V. “praecorsac” differs from MNHN.F.ACA-293 the Çalta-1 specimens display many differences such as: for its elongation and buccolingual compression; the rounded 1) the studied material is stouter as compared to the average lingual margin of the talonid; the reduced entoconid com- of V. ­alopecoides (Fig. 4); 2) in the European species the P4 pared to the hypoconid (Fig. 3I, J, R, S); and 6) the m2 of lingual cingulum is less prominent whereas the P4 metastyle V. “praecorsac” differs from that of Çalta-1 for the peculiar appear enlarged at its base compared to the latter; 3) the corpus straight morphology of the mesial-mesiolingual side; the of the mandible from Çalta-1 shares with some specimens of mesially expanded buccal cingulid and for the narrower V. alopecoides the high position of the distal mental foramen talonid. It is also proportionally longer compared to that of (close to the alveolar plane); 4) the m1 trigonid of Vulpes MNHN.F.ACA-293 (Fig. 4). from Çalta-1 resemble that of earlier forms of V. alopecoides, Few isolated remains of foxes were recently found in the e.g. Kvabebi (Fig. 3I, N), in which it is slender compared Oldowan north-Caucasian site of Muhkai II (Amirkhanov the stouter morphology of Early Pleistocene forms, e.g. from et al. 2016). The former fragmentary remains were tenta- Pirro Nord (Fig. 3P); 5) the m1 transverse cristid of Çalta-1 tively included in Vulpes alopecoides without describing it is incomplete like in V. beihaiensis (Fig. 3U), whereas in in detail. Only one partial mandibular corpus with p2-p4 V. alopecoides it is generally complete (Fig. 3N-Q); 6) the were figured by Amorkhanov et al. (2016), which displays m2 of V. alopecoides from Kvabebi (Fig. 3N) differs from narrow premolars and a proportionally enlarged distal that of Çalta-1 for its more mesiodistally elongated occlusal accessory cuspulids as compared with Çalta-1 specimens. morphology, the larger trigonid cuspulids, the reduced lin- Such features also contrast with the usual morphology gual side, including the entoconid; and 7) with the exception 624 COMPTES RENDUS PALEVOL • 2021 • 20 (29) Ancestral foxes at the gates of Europe 0.10 0.05 0 -0.05 -0.10 -0.15 P3 L P4 L P4 W M1 L M1 W M2 L M2 W p3 L p4 L p4 W m1 L m1 W trm1 L tdm1 W m2 L m2 W Mp4H Mm1H Mm1B Vulpes from Çalta V. alopecoides V. beihaiensis V. “paecorsac” from Odessa V. qiuzhudingi V. Vulpes Vulpes sp. from Musselievo Fig. 4. — Log-ratio diagram based on selected dentognathic variables in Vulpes Frisch, 1775 from Çalta-1 and other Pliocene-Early Pleistocene species (V.­­ ­alopecoides (Del Campana, 1913) from Europe sensu Bartolini-Lucenti & Madurell-Malapeira 2020; V. “praecorsac” Kormos, 1932 from Odessa; V. beihaiensis Qiu & Tedford, 1990 from Yushe Basin; V. qiuzhudingi Wang, Tseng, Li, Takeuchi & Xie, 2014 from Zanda Basin; and Vulpes sp. from Musselievo). The average values of V. vulpes (Linnaeus, 1758) are used as standard reference. of the P4, V. alopecoides is generally larger than Vulpes from constrained records of small- to medium-sized foxes is that Çalta-1 (Fig. 4). Their pattern of proportions does not differ of the described specimens of V. beihaiensis from Çalta-1 considerably one another. (c. 4.0 Ma). This species configures as one of the key species of the Pliocene of Eurasia, ranging across the Asian continent until the middle Late Pliocene, as testified to by the Mazeg- CONCLUSIONS ouan remains from Yushe Basin (Qiu & Tedford 1990). Sub- sequently, around 3.1 Ma, the first V. alopecoides-like fox – the In the framework of the Pliocene record of Vulpes across Eurasia, species that characterized the Late Pliocene-Early Pleistocene characterized by numerous taxa of uncertain affinities (Fig. 1), record of Europe – appeared, in the eastern Georgian site of the record of Çalta-1 has a significant relevance. Firstly, for its Kvabebi (Rook et al. 2017). Considering the morphological geographical position, at the gates of Europe but close to the similarity between V. beihaiensis and V. alopecoides, like the junction between three continents (Sen 1998). Secondly, for Kvabebi Vulpes, it seems plausible that the latter species derives its chronology, representing one of the earliest well-calibrated from a western population of V. beihaiensis, diverging from occurrences of Vulpes in the Eurasian continent. As shown it between 4.0 and 3.1 Ma. In the Early Pleistocene V. alope- above, the similarity between the specimens from Çalta-1 and coides spread across Europe and became a common element those of V. beihaiensis from Yushe Basin, both morphologi- of European Early Pleistocene faunas (Bartolini-Lucenti & cal and biometric, especially in comparison to that of other Madurell-Malapeira 2020). Finally, during the Middle Pleis- Pliocene localities, supports the parsimonious and plausible tocene, as documented in western European site, the extant hypothesis of subjective synonymy between these taxa under red fox (Vulpes vulpes) might have arisen from a yet unknown the specific name of V. beihaiensis (following priority rule of population of V. alopecoides. the International Commission on Zoological Nomenclature 1999). Such a result contrasts with the interpretation by Ginsburg (1998). Nevertheless, this author did not compare Acknowledgements the specimens from Çalta-1 to the Chinese material described The authors are thankful to the kindness and availability of fol- by Qiu & Tedford (1990). Despite his specific attribution, lowing curators of universities and institutions: E. Cioppi and we agree with Ginsburg (1998)’s interpretation of a plausible P. Agnelli, curators of the IGF and of the MZUF, respectively; link between the Vulpes from Çalta-1 and the European Pleis- M. Pavia of the Paleontology collection of the MGPT-PU; tocene V. alopecoides, yet considering them as distinct species. M. Bukhsianidze and D. Lordkipanidze of the MG-GNM; Following the results described above, a new paleobiogeo- C. Argot, curator of the Paleovertebrate collection of the graphic scenario for the Old-World foxes is here suggested. MNHN; J. Galkin and J. Men, curator of the fossil mammals After their arise in the late Miocene of North America (c. 9.0- collection of AMNH; M. Gasparik, curator of the Paleoverte- 8.0 Ma), the center of origin and radiation of all Canidae brate collection of the HNHM. The MNHN gives access to (Wang & Tedford 2008; Tedford et al. 2009), members of the the collections in the framework of the RECOLNAT national genus Vulpes reached the Old World rather early, as the oldest Research Infrastructure. We also thank S. Sen for the fruitful record is in Chad in the Tortonian (late Miocene, c. 6.8 Ma; discussion on the site of Çalta-1 and its fauna and Dr X. Wang de Bonis et al. 2007). Among the earliest occurrences of Vulpes for kindly providing us the pictures of V. qiuzhudingi. The in Eurasia, the first stratigraphically and chronologically well- authors thank Prof. G. Koufos and an anonymous reviewer COMPTES RENDUS PALEVOL • 2021 • 20 (29) 625 Bartolini-Lucenti S. & Madurell-Malapeira J. for their suggestions that improved the first version of the Kormos T. 1932. — Die Füchse des ungarischen Oberpliozän. manuscript. This work has been funded by the Agencia Estatal Folia Zoologica hydrobiologica 4: 167-188. de Investigación – European Regional Development Fund Lunkka J. P., Kappelman J., Ekart D. & Sen S. 1998. — Sedi- mentation and lithostratigraphy, in Sen S. (ed), Pliocene verte- of the European Union (CGL2017-82654-P, AEI/FEDER- brate locality of Çalta, Ankara, Turkey. Geodiversitas 20: 329-338. UE) and the Generalitat de Catalunya (CERCA Program). Lunkka J. P., Kappelman J., Ekart D., Crabaugh J. & Gibbard P. S.B.L. study visits of to AMNH and MG-GNM have been 2003. — Geology, in Fortelius M., Kappelman J., Sen S. & supported by the Earth Sciences Department, University of Bernor R. L. (eds), Geology and Paleontology of the Miocene Sinap Florence (Ateneo funds, and International mobility funds) Formation, Turkey. Columbia University Press, Columbia: 25-40. Odintzov I. A. 1965. — Vulpes praecorsac Kormos from Pliocene and by the Italian Ministry of Foreign Affairs (DGSP-VI deposits of Odessa. Paleontologicheskyi Zbornik, L’vov University funds). J.M.-M. is member of consolidated research group 2: 57-64 (in Russian with English summary). 2017 SGR 116 (AGAUR, Generalitat de Catalunya). This Qiu Z. & Tedford R. H. 1990. — A Pliocene species of Vulpes work is part of the “Augusto Azzaroli (1921-2015) centennial from Yushe, Shanxi. Vertebrata Palasiatica 28: 245-258. papers” by the Vertebrate Paleontology Group at the Earth Qiu Z., Deng T. & Wang X. 2004. — Early Pleistocene mammalian fauna from Longdan, Dongxiang, Gansu, China. Paleontologia Sciences Department, University of Florence. Sinica, New Series C 27: 1-198. Rook L., Bartolini-Lucenti S., Bukhsianidze M. & Lordki- panidze D. 2017. — The Kvabebi Canidae record revisited (late REFERENCES Pliocene, Sighnaghi, eastern Georgia). Journal of Paleontology 91: 1258-1271. https://doi.org/10.1017/jpa.2017.73 Amirkhanov H. A., Ozherelyev D. V., Sablin M. V. & Aga- Sen S. (ed.) 1998. — Pliocene vertebrate locality of Çalta, Ankara, dzhanyan A. K. 2016. — Faunal remains from the Oldowan Turkey. Geodiversitas 20 (3), arts 7-19. site of Muhkai II in the North Caucasus: Potential for dating Sen S. & Saraç G. 2018. — Hyaenidae (Carnivora, Mammalia) and palaeolandscape reconstruction. Quaternary International from late Miocene and Pliocene of Çalta (Ankara, Turkey). Revue 395: 233-241. https://doi.org/10.1016/j.quaint.2014.12.061 de Paléobiologie 37 (2): 561-575. Bartolini-Lucenti S. 2021. — A new large-sized Pliocene fox Sharapov S. 1986. — The Kuruksay Complex of Upper Pliocene (Carnivora, Canidae) from Yushe Basin (Shanxi, China). Rivista Mammalian of Afghan-Tadjik Depression. Tadzhik Academy of Italiana di Paleontologia e Stratigrafia 127: 133-147. https://doi. Sciences, Dushambe, 251 p. org/10.13130/2039-4942/15206 Simpson G. G. 1941. — Large Pleistocene felines of North ­America. Bartolini-Lucenti S. & Madurell-Malapeira J. 2020. — Unra- American Museum Novitates 1136: 1-27. http://hdl.handle. veling the fossil record of foxes: An updated review on the Plio- net/2246/2262 Pleistocene Vulpes spp. from Europe. Quaternary Science Reviews Simpson G. G., Roe A. & Lewontin R. C. 1960. — Quantitative 236: 106296. https://doi.org/10.1016/j.quascirev.2020.106296 Zoology. Harcourt Brace, New York, 470 p. Beaumont G. 1978. — Notes complémentaires sur quelques Spassov N. 2000. — Biochronology and zoogeographical affinities félidés (Carnivores). Archives des Sciences - Université de Genève of the Villafranchian faunas of Bulgaria and south Europe. Historia 31: 219-227. Naturalis Bulgarica 12: 89-128. https://www.biodiversitylibrary. Bernor R. L. & Sen S. 2017. — The Early Pliocene Plesiohippar- org/page/31093288 ion and Proboscidipparion (Equidae, Hipparionini) from Çalta, Szuma E. 2008a. — Geographic variation of tooth and skull sizes Turkey (Ruscinian Age, c. 4.0 Ma). Geodiversitas 39: 285-314. in the arctic fox Vulpes (Alopex) lagopus. Annales Zoologici Fennici https://doi.org/10.5252/g2017n2a7 45:185-199. https://doi.org/10.5735/086.045.0304 Croizet J. B. & Jobert A. C. G. 1828. — Recherches sur les osse- Szuma E. 2008b. — Geography of sexual dimorphism in the tooth ments fossiles du département du Puy-de-Dôme. Chez les principaux size of the red fox Vulpes vulpes (Mammalia, Carnivora). Journal libraires, Paris, 225 p. of Zoological Systematics and Evolutionary Research 46: 73-81. Daitch D. J. & Guralnick R. P. 2007. — Geographic variation https://doi.org/10.1111/j.1439-0469.2007.00418.x in tooth morphology of the arctic fox, Vulpes (Alopex) lagopus. Szuma E. 2008c. — Evolutionary and climatic factors affecting Journal of Mammalogy 88: 384-393. https://doi.org/10.1644/06- tooth size in the red fox Vulpes vulpes in the Holarctic. Mamma- MAMM-A-139R1.1 lian Research 53: 289-332. https://doi.org/10.1007/BF03195193 de Bonis L., Peigné S., Likius A., Mackaye H. T., Vignaud P. & Tedford R. H., Wang X. & Taylor B. E. 2009. — Phylogenetic Brunet M. 2007. — The oldest African fox (Vulpes riffautae n. sp., systematics of The North American fossil Caninae (Carnivora: Canidae, Carnivora) recovered in late Miocene deposits of the Canidae). Bulletin of the American Museum of Natural History Djurab desert, Chad. Naturwissenschaften 94: 575-580. https:// 325: 1-218. https://doi.org/10.1206/574.1 doi.org/10.1007/s00114-007-0230-6 Wagner J. A. 1848. — Urweltliche Säugthier-Ueberreste aus Del Campana D. 1913. — I cani Pliocenici di Toscana. Palaeon- Griechenland (Vol. 5). Königlich Akademie der Wissenschaften tographia Italica 19: 189-254. 5: 335-378. Depéret C. 1890. — Les Animaux Pliocènes du Roussillon. Wang X. & Tedford R. H. 2008. — Dogs: Their Fossil Relatives Mémoires de la Société géologique de France 3: 5-195. and Evolutionary History. Columbia University Press, Colum- Gimranov D. O., Kosintsev P. A. & Gasilin V. V. 2015. — Spe- bia, 240 p. cies diagnostics of corsac (Vulpes corsac), fox (Vulpes vulpes), and Wang X., Tseng Z. J., Li Q., Takeuchi G. T. & Xie G. 2014. — arctic fox (Vulpes lagopus) according to ontological characteristics From ‘third pole’ to north pole: a Himalayan origin for the arctic of mandible teeth. Zoological Journal 94: 1338-1350 (in Russian). fox. Proceedings of the Royal Society B: Biological Sciences 281 Ginsburg L. 1998. — Le gisement de vertébrés pliocènes de Çalta, (1787): 20140893. https://doi.org/10.1098/rspb.2014.0893 Ankara, Turquie. 5. Carnivores. Geodiversitas 20: 379-396. Gimranov D. O. 2017. — Species diagnostics of the corsac (Vulpes corsac), fox (Vulpes vulpes) and arctic fox (Vulpes lagopus, Car- Submitted on 2 May 2020; nivora, Canidae) using the upper teeth. Zoological Journal 96: accepted on 14 June 2020; 684-697 (in Russian). published on 13 July 2021. 626 COMPTES RENDUS PALEVOL • 2021 • 20 (29)