FOSS I L I MP RI N T • vo l. 75 • 2019 • no. 3–4 • p p. 3 9 7 –4 1 1 (for mer ly AC TA MUSEI NATIONALIS PR AGAE, S er ies B – H i stor ia Naturalis) A NEW THAUMASTOCYONINAE (AMPHICYONIDAE, CARNIVORA) FROM THE EARLY MIOCENE OF TUCHOŘICE, THE CZECH REPUBLIC JORGE MORALES1,*, OLDŘICH FEJFAR2, ELMAR HEIZMANN3, JAN WAGNER2, ALBERTO VALENCIANO4,5, JUAN ABELLA6,7 1 Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC, C/José Gutiérrez Abascal 2, 28006, Madrid, Spain; e-mail: jorge.morales@mncn.csic.es. 2 Department of Palaeontology, National Museum, Václavské nám. 68, 110 00 Prague 1, the Czech Republic; e-mail: oldrich.fejfar@natur.cuni.cz; jan_wagner@nm.cz. 3 Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany; e-mail: elmar.heizmann@smns-bw.de. 4 Research and Exhibitions Department, Iziko Museums of South Africa, P.O. Box 61, Cape Town, South Africa; e-mail: alb3rtovv@gmail.com. 5 Department of Biological Sciences, University of Cape Town, Palaeobiological Research Group, Private Bag X3, Rhodes Gift 7701, Cape Town, South Africa. 6 Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain; e-mail: juan.abella@icp.cat. 7 Instituto Nacional de Biodiversidad, Pje. Rumipamba N. 341 y Av. de los Shyris (Parque La Carolina) Quito, Ecuador. * corresponding author Morales, J., Fejfar, O., Heizmann, E., Wagner, J., Valenciano, A., Abella, J. (2019): A new Thaumastocyoninae (Amphicyonidae, Carnivora) from the early Miocene of Tuchořice, the Czech Republic. – Fossil Imprint, 75(3-4): 397–411, Praha. ISSN 2533- 4050 (print), ISSN 2533-4069 (on-line). Abstract: New Amphicyonidae fossil remains from the early Miocene site of Tuchořice (the Czech Republic) confirm the presence of a new Thaumastocyoninae taxon: Peignecyon felinoides n. gen. et n. sp. It is characterized by a peculiar combination of plesiomorphic and derived morphological traits. The new genus can be defined by a long and sharp mandible diastema, loss of mesial premolars (p2–p3), p4 with an inclined distally high main cuspid, moderate sectorial carnassial teeth, m1 with relict metaconid, and talonid and trigonid of similar width, and reduced M2 and m2. In the phylogenetic analysis the Thaumastocyoninae form a monophyletic group characterized by the start of the m2/M2 reduction, still moderate in Crassidia intermedia (von Meyer, 1849), but remarkable in the other species of the clade. Peignecyon felinoides already shows the advanced features defining the Thaumastocyoninae, and constitutes the sister group of the most specialized genera Tomocyon Viret, 1929b and Thaumastocyon Sthelin et Helbing, 1925. Consequently, it can be considered an excellent link between this group and the more primitive members of the tribe Ysengrini (Ysengrinia Ginsburg, 1966 and Crassidia Heizmannn et Kordikova, 2000). Peignecyon felinoides shows that the trend towards hypercarnivory had already emerged in the European early Miocene fauna, thus helping to understand the complex evolution of the Amphicyonidae during the Miocene. Key words: Thaumastocyoninae, Amphicyonidae, systematics, Miocene, Europe Received: April 8, 2019 | Accepted: June 21, 2019 | Issued: December 30, 2019 Introduction m2–m3, m1-talonid), and the sharp and robust carnassial teeth (m1 and P4). However, despite the constancy of this The Amphicyonidae family represented one of the most dental pattern, certain divergent adaptations took place. characteristic groups of carnivorans in the European faunas Some involved an increase in the molar dental surface, for approximately 35 million years, from its first appearance tending towards a less specialized diet, others exhibited a during the Eocene (ca. 40 Ma) to its extinction in the late reduction of this grinding surface, while mesio-distally Miocene (ca. 5 Ma). Despite the success of the group and the stretching the carnassials, by way of an adaption to a large temporal range registered, they generally maintained more carnivorous diet. This hypercarnivory has been seen a very homogeneous dental pattern, characterized by the to recur throughout the amphicyonid fossil record (Hunt large size (similar to that of the largest extant terrestrial 1998), first during the Oligocene and early Miocene in carnivores), robustness of the crushing molars (M1–M2, and the North American Daphoenidae Hough, 1948 and its 398 DOI 10.2478/if-2019-0025 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC Eurasian relatives, the Haplocyoninae de Bonis, 1966, and subsequently, with the Thaumastocyoninae Hürzeler, 1940, which appear at the same time as the last haplocyonines (Hunt 1998, Peigné and Heizmann 2003, Wang et al. 2016). The phylogenetic relationships within the Amphicyonidae species present notorious difficulties, as can be seen in some previous studies (Viranta 1996, Peigné et al. 2008, Morales et al. 2016). Several problems in this regard involve the proliferation of species, some of them defined with few diagnostic characters, the coexistence of different species in the same locality, the broad interspecific variability, and the perception of a parallel evolution to the amphicyonid lineages, with the multiplication of homoplasic characters. The present paper attempts to describe the hypercarnivorous Amphicyonidae from Tuchořice, placing particular emphasis upon the systematic revision of the Thaumastocyoninae subfamily. Text-fig. 1. Dental nomenclature. Peignecyon felinoides n. gen. et n. sp. a) TU 73916 upper right M1 in occlusal view; b) TU 7391147 lower right m1, b1) lingual view, b2) occlusal view. Abbreviations: bde, basal distobuccal expansion*; bc, buccal Tuchořice fossil locality cingulum; Ec, ectoflexus; dc, distal cingulum; dlc, distolingual The fossiliferous locality of Tuchořice is located in cristid; end, entoconid; hyd, hypoconid; lc, lingual cingulum; North Bohemia, the Czech Republic, close to the south- ME, metacone; MEC, metaconule; mc, mesial cingulum; med, metaconid; mlc, mesiolingual cristid; mpc, mesial protoconid west margin of the Most brown coal basin, near the villages cristid; MS, metastyle; PA, paracone; PAC, paraconule; pad, of Tuchořice and Lipno (Fejfar and Heizmann 2016). The paraconid; PR, protocone; prd, protoconid; PS, parastyle. *Not freshwater limestones of Tuchořice were believed to be developed in Peignecyon felinoides. the early Miocene equivalent of the underlying marls of the main brown coal seam located in the North Bohemian Tertiary basin. The mammal remains from Tuchořice have Material and methods been known since the nineteenth century. The first study of mammalian fossils from there was of fragmentary rhinoceros teeth reported by Suess (1861). Much richer finds from the Nomenclature and measurements travertines near Tuchořice were collected many years later Dental nomenclature (Text-fig. 1) follows Ginsburg by O. Fejfar. The small mammalian fauna has extremely (1999). Anatomical descriptions are based on Schaller low diversity in comparison with the nearby locality of (2007). Measurements (Tab. 2) were made using Mitutoyo Merkur-North (= Ahníkov 1) (Fejfar et al. 2003). Remains Absolute digital calliper to the nearest 0.1 mm. of large mammals are well preserved and diversified, especially the Rhinocerotidae (Heissig and Fejfar 2007) and Institutional abbreviations carnivores (Fejfar and Heizmann 2016). The abundance of MNHL Musée des Confluences, Lyon, France carnivore predators is interpreted as due to the supposed MNHN Muséum national d’Historie naturelle, Paris, “trapping affect” of the CO2-rich exhalations of the hot France mineral springs. The abundance of Amphicyonidae teeth is MNCN Museo Nacional de Ciencias Naturales, Madrid, remarkable, and there is evidence of at least five species, Spain ranging from the small Amphicyon schlosseri Dehm, 1950 to the biggest Megamphicyon giganteus (Schinz, 1825). NMB Naturhistorisches Museum Basel, Basel, Switzerland The Amphicyonidae association is comparable to that of NMP National Museum, Prague, the Czech Republic Wintershof-West (Dehm 1950), but with some differences; (inv. no. NM-Pv) in Tuchořice, Pseudocyon bohemicus Schlosser, 1901 is TU Tuchořice locality, cast collection (housed in NMP abundant, while this species is absent in Wintershof-West. and MNCN) Amphicyon dietrichi Dehm, 1950 is taxonomically close, UCB Université Claude Bernard Lyon 1, Lyon, France but scarce. The most striking difference is the presence Studied material in Tuchořice of a new Thaumastocyoninae species, a group that seems to be absent in Wintershof-West. The The holotype, NM-Pv 11600 right mandible with c–p4– Tuchořice sample is the most complete fossil record of this m1, alveoli for m2–m3, of Peignecyon felinoides n. gen. et elusive group. New collections and comparisons of similar n. sp. from Tuchořice (Pl. 1) is stored in the collections of mammalian taxa have shown the more advanced character NMP. The fossil associated described in the present work – of some species, indicating a somewhat higher level in TU 73917 left mandible with p4–m2, alveoli p3 y m3 (Pl. the Miocene mammalian zone MN 3 (Fejfar et al. 2003, 2, Fig.1); TU 739137 left mandible fragment with p4 (Pl. Heissig and Fejfar 2007, Fejfar and Heizmann 2016). 2, Fig. 2); TU 739118 left mandible with p4–m3 (Pl. 3, Fig. The Tuchořice fossil localities are now by law Protected 1); TU 739115 right m1–m2 (Pl. 3, Fig. 2); TU 7391147 Palaeontological Site. right p4–m2 (Pl. 3, Fig. 3); TU 739111 right mandible with 398 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC m1–m3 (Pl. 3, Fig. 4 ); TU 739151 left m2 (Pl. 3 Fig. 5); Table 1. Character list and character states. TU 739110 right mandible with p4–m1; TU 739130 right P4 (Text-fig. 2a); TU 739129 right M1 (Text-fig. 2b); TU 1 Ratio LM2/LM1; >0.72 (0), between 0.72–0.62 (1), <0.62 (2) 739130 right M2 (Text-fig. 2c); TU 73916 right M1; 739136 2 Ratio LM1/LP4; <0.76 (0), between 0.76–0.88 (1), >0.88 (2) left M2; TU 739151 left M2 (Text-fig. 2d) – are currently 3 P4 parastyle; small (0), strong (1), very developed (2) lost. The cast of these fossils are available in the collections 4 P4 protocone position; mesial (0), more distal (1) of NMP and MNCN. 5 P4 protocone size; strong (0), reduced (1) For systematic comparison and cladistics analysis, P4 ratio W/L P4; >0.70 (1), between 0.70–0.58 (1), we have directly studied the following taxa: Ysengrinia 6 <0.58 (2) gerandiana (Viret, 1929a) and Cynelos lemanensis (Pomel, 7 M1 shape; subquadrangular (0), subtriangular (1) 1846) from Saint Gérand le Puy (France), early Miocene 8 M1 paraconule; present (0), divided (1), reduced (2) (MN 2) housed in MNHN and UCB; Thaumastocyon dirus M1 metaconule; present (0), divided (1), enlarged (2), 9 Ginsburg et al., 1981 from Los Valles de Fuentidueña reduced or lost (3) (Spain), late Miocene (MN 9) housed in MNCN; the 10 M1 paracone-metacone; low (0), high (1) undescribed dentition of Thaumastocyon n. sp. from 11 M1 distal wall middle inflexion; present (0), absent (1) Batallones-3 (Spain), late Miocene (MN 10) housed in 12 M1 protocone size; low/weak (0), high/strong (1) MNCN; Tomocyon grivensis Viret, 1929b from La Grive 13 M1 protocone position; mesial (0), central (1) Saint Alban (France), middle Miocene (MN 7/8) housed M1 lingual cingulum position; distal /asymmetric (0), 14 in UCB and MNHL; Ysengrinia valentiana Belinchón et distomesial/more symmetric (1) Morales, 1989 from Buñol (Spain), early Miocene (MN 4) 15 M1 lingual area; wide (0), narrowed (1) by casts housed in MNCN; Thaumastocyon bourgeoisi 16 M1 lingual platform; large (0), reduced (1) Stehlin et Helbing, 1925 from Pont-Levoy (France), 17 M1 lingual cingulum; well developed (0), reduced (1) middle Miocene (MN 5) by casts housed in the MNHN; 18 M1 ratio W/L; <0.72 (0), between 0.72–0.84 (1), >0.84 (2) Daphoenodon superbus (Peterson, 1907) from Agate Fossil 19 M2 shape; short and large (0), oval (1) beds National Monument (USA) and Ysengrinia americana 20 M2 ratio W/L; <0.62 (0), between 0.62–0.75 (1), >0.75 (2) (Wortman, 1901) from Harrison Formation (USA), early 21 M2 trigone cusp; differenciated (0), fused (1), reduced (2) Miocene (late Arikarean) by casts housed in MNCN. 22 Premolars (p1–p3); not reduced (0), reduced (1) 23 Ratio Lm2/Lm1; >0.62 (0), <0.62 (1) Cladistic analysis 24 m1 metaconid size; strong (0), reduced or lost (1) We performed a cladistic analysis to better assess the 25 m1 hypoconid; lateral (0), central (1) relationships between Peignecyon felinoides n. gen. et n. sp., m1 talonid – basal distobuccal expansion; absent (0), and other amphicyonid taxa, such as the Oligocene Pseudo- 26 present (1) cyonopsis spp. (Pseudocyonopsis antiquus Ginsburg, 27 m1 ratio W/L; >0.60 (0), <0.60 (1) 1966, Pseudocyonopsis ambiguus (Filhol, 1876) and m2 ratio L talonid / L trigonid; >0.85 (0), between 28 Pseudocyonopsis quercensis (Ginsburg, 1965)), the early 0.85–0.68 (1), <0.68 (2) Miocene Daphoenodon superbus, Cynelos lemanensis, 29 m2 ratio W/L; >0.60 (0), <0.60 (1) Ysengrinia americana, Ysengrinia gerandiana, Ysengrinia m2 height of protoconid; protoconid > metaconid (0), 30 valentiana and Crassidia intermedia (von Meyer, 1849), the protoconid >> metaconid (1) middle Miocene Tomocyon grivensis and Thaumastocyon 31 m2 protoconid mesial cristid; oblique (0), rectilinear (1) bourgeosi, and the late Miocene Thaumastocyon sp. from 32 m2 paraconid; present (0), absent (1) Batallones 3 (Morales et al. 2017), together with the 33 m2 metaconid; present (0), reduced or absent (1) primitive carnivoran Paramiacis exilis (Filhol, 1876). 34 m2 talonid; large (0), narrow (1) We excluded from the analysis some taxa related to the 35 m2 hypoconid; small/buccal (0), enlarged/central (1) Thaumastocyoninae, such as Ysengrinia tolosana (Noulet, 36 m2 buccal wall; almost vertical (0), basally expanded (1) 1876), Ysengrinia depereti (Mayet, 1908), Thaumastocyon dirus and Agnotherium antiquum Kaup, 1833 because they are poorly represented, or there is no information on their upper dentition. The cladistics analysis includes 12 taxa and Systematic palaeontology 36 equally weighted and unordered dental characters (21 Order Carnivora Bowdich, 1821 binary characters and 15 multistate characters; see Tab. 1). Suborder Caniformia Kretzoi, 1943 Some of them were used by Viranta (1996) (3, 5, 7, 12, 13, Infraorder Arctoidea Flower, 1869 19, 23, 24 and 34) and by Peigné et al. (2008) (1, 3, 4, 5, 10, Family Amphicyonidae Trouessart, 1885 22, 23, 24 and 33). The choice of characters was obtained Subfamily Thaumasthocyoninae Hürzeler, 1940 from the molars (M1–M2 and m2) and the carnassial teeth (P4/m1). As information on the premolars is more limited, D i a g n o s i s . Amphicyonidae presenting hyper- they play (except p4) a secondary role in the evolution carnivorous dentition, with reduced M2/m2–m3 and highly of hypercarnivorous amphycyonids. The matrix can be developed and sectorial carnassial teeth. Specialized genera observed in Table 3. We selected P. exilis as the outgroup. with a reduction of the premolars and a large mandibular The analysis was performed with PAUP*4.0b10 (Swofford diastema, well-developed P4 parastyle and reduced M1 2002). lingual area. 399 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC I n c l u d e d g e n e r a . Thaumastocyon Stehlin et Hürzeler (1940) within the Thaumastocyoninae, should Helbing, 1925, Crassidia Heizmann et Kordikova, 2000, be distinguished from Pseudocyon and attributed to Ysengrinia Ginsburg, 1966, Tomocyon Viret, 1929b and Amphicyonopsis? serus (Kuss, 1965). Peignecyon n. gen. Mein and Ginsburg (2002), partially in accordance with Kuss (1965), transferred the fossils of La Grive to R e m a r k s . The taxonomy and systematics of the Pseudocyon sansaniensis and therefore excluded them from Thaumastocyoninae amphicyonid have been considered the Thaumastocyoninae. to be quite complex since Hürzeler (1940), who described Identification of Agnotherium and Ysengrinia in Africa is them as aberrant canids with feline tendencies. The latter problematic, as has recently been discussed by Morales et al. author correctly characterized the subfamily based on the (2016). For these authors, the African amphicyonids previously complete suppression of m1 metaconid, reduction of the identified in these genera acquired their hypercarnivorous premolars, except the p4, which is reinforced, and the adaptation independent from the European taxa. This could oblique abrasion of the teeth. He included in the group be the case of Ysengrinia americana, as will subsequently be Thaumastocyon bourgeoisi, Agnotherium antiquum Kaup, discussed. The Tuchořice Thaumastocyoninae are important, 1833, Tomocyon grivensis Viret, 1929b together with the since they can help to identify the beginning of the more amphicyonid remains from Frohnstteten, figured by Jäger specialized Thaumastocyoninae, and because they are a (1835), unpublished materials housed in the Basel Museum significant link between these genera and the more primitive (Switzerland) from Baigneaux-en-Beauce (France) and species included in the Ysengrini tribe by Heizmann and Chairmoille (Switzeland), and eventually, a mandible Kordikova (2000). fragment from Steinheim (Germany), which was determined Thus, our paper accepts the presence of two tribes by Helbing (1929) as Pseudocyon sansaniensis Lartet, of Thaumastocyoninae in the late Oligocene/Miocene 1851, and subsequently chosen as the holotype of a new of Europe – Thaumastocyonini Hürzeler, 1940 which subspecies, Pseudocyon sansaniensis serus Kuss, 1965 included the genera Thaumastocyon Stehlin et Helbing, (Kuss 1965: 133). 1925, Tomocyon Viret, 1929b and Peignecyon n. gen., and Kuss (1962) performed the first in-depth review of the Ysengrini Heizmannn et Kordikova, 2000 which included group, in which he proposed the inclusion of Tomocyon the genera Ysengrinia Ginsburg, 1966 and Crassidia grivensis in Agnotherium. He considers that A. antiquum Heizmannn et Kordikova, 2000. likely had upper molars with a non-reduced lingual area, an We did not consider some taxa, such as Amphicyonopsis argument which favoured the determination of Frohnstetten serus (Kuss, 1965), which we believe to be a species fossils as A. antiquus, reconstructing the molar morphology of dubious validity, or Agnotherium antiquum Kaup, with a strong development of the lingual area, using as a 1833, defined in Eppelsheim (MN 9), Germany, and only model the molars of T. grivensis (Kuss 1962: fig. 6). For represented by one sectorial m1. This tooth displays an some fossils from Baigneaux-en-Beauce, Kuss (1962: fig. 7) advanced degree of dental wear, which has partially erased proposes their classification as Agnotherium grivensis (right its morphology. The paraconid mesial cristid is displaced m1; NMB S.O. 4459), aff. Agnotherium grivensis (for three lingually and is well marked, connecting to the basal P4; NMB S.O. 1274, 874 and 4460) and ?aff. Agnotherium lingual cingulum; the talonid is short and narrow with a grivensis (left M2; NMB S.O. 1559). high hypoconid and a reduced entoconid. These characters Ginsburg (1999) seconds this proposal, pointing out are associated with the hypercarnivorous tendency, and the presence of Agnotherium grivensis in Simorre (France) might likely have been acquired convergently in different and Agnotherium antiquum in Pedregueras (Spain), and the Thaumastocyoninae taxa. The genera Thaumastocyon and presence of a second Thaumastocyon species, Th. dirus from Tomocyon present an m1 close to A. antiquus, but differ in Los Valles de Fuentidueña (Spain) (Ginsburg et al. 1981). their associated upper and lower teeth. We do not avail of Heizmann and Kordikova (2000) erected the tribe any criteria to relate the Eppelsheim genus more closely to Ysengrini to contain Crassidia, a new genus described in the either one of these genera. Given our current knowledge, same research, together with Ysengrinia and Amphicyonopsis this m1 is insufficient with regard to defining a genus, or Viret, 1929b. For these authors, the Ysengrini occupies even a species. Most of the Agnotherium antiquus published an intermediate position between Amphicyoninae and determinations have been indirect, based upon tooth size Thaumastocyoninae, but since the defining features of and sectorial morphology of specimens. This has given rise Thaumastocyoninae, loss of the m1 metaconid and shortening to multiple errors, as Kurtén (1976) initially pointed out. of the entire facial region, are not present in the Ysengrini, The determination of Agnotherium antiquus in Pedregueras they consider them to constitute a specialized branch of the 2A and Charmoille exemplify this circumstance; in both Amphicyoninae. But they did not take into account that both localities the material comprises a lower canine, p4 and m1; groups share other derived characters, which indicate a close the m1s are morphologically close to the tooth of Eppelsheim, relationship between them. although the mesial paraconid-cristid is less marked. The Amphicyonopsis was described by Viret (1929b) on the canine and p4 present a similar morphology and robustness basis of m1 (MHNL LGR 1136) from La Grive (France), to those of La Grive homologous teeth (Viret 1929a: pl. but no species was included in the new genus. Kuss (1965) 19, figs 3, 7). The conclusion is evident; there is a suite of included it, together with other teeth from La Grive as features such as the advanced degree of m1 sectorialitation Pseudocyon sansaniensis aff. serus n. subsp. Heizmann (loss of metaconid, talonid reduction, sharp cristid, etc.), the (1973) considers that the holotype of this subspecies, robustness of the lower canine and the p4, and others, which the mandible fragment from Steinheim included by enable the Thaumastocyonini clade to be defined, but which 400 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC by themselves do not permit an unequivocal determination within this taxon. Genus Peignecyon n. gen. T y p e s p e c i e s . Peignecyon felinoides n. sp. E t y m o l o g y . In honour of the late Stéphane Peigné. T y p e l o c a l i t y . Tuchořice, the Czech Republic. A g e . Early Miocene (MN 3). D i a g n o s i s . Thaumastocyoninae presenting long mandible diastema, reduced mesial premolars (p2–p3), p4 with inclined distally high main cusp; sectorial carnassials, talonid and trigonid of m1 with similar width, relict m1 metaconid. Reduced M2 and m2. Peignecyon felinoides n. sp. Text-figs 1, 2, Pls 1–3 2003 Tomocyon nov. sp.; Fejfar et al., p. 167. 2016 Tomocyon (? nov. gen.), nov. sp.; Fejfar and Heizmann, p. 325. H o l o t y p e . NM-Pv 11600 right mandible with p4–m1, alveoli for m2–m3 (Pl. 1), deposited in NMP. P a r a t y p e . TU 73916 right M1 (Text-fig. 2b). E t y m o l o g y . “felinoides” from feline, denomination used by Hürzeler (1940) for this group of Amphicyonidae. T y p e l o c a l i t y . Tuchořice, the Czech Republic. A g e . Early Miocene (MN 3). D i a g n o s i s . The same as for the genus. D e s c r i p t i o n . Mandible and lower dentition. NM- Pv 11600 right mandible with c, p1 alveolus, p4–m1, and Text-fig. 2. Peignecyon felinoides n. gen. et n. sp., from Tu- alveoli for m2–m3 (holotype, Pl. 1), the mandible is included chořice, the Czech Republic. a) TU 739130 right P4; a1) buccal in the sediment with the lingual wall exposed. The mandible view, a2) occlusal view, a3) lingual view. b) TU 73916 right M1; ramus was partially broken, but the fragments remained close b1) buccal view, b2) occlusal view, b3) lingual view, b4) mesial to the corpus mandibulae, and this part has thus been restored view, b5) distal view. c) TU 739130 right M2; c1) buccal view, c2) occlusal view, c3) lingual view, c4) mesial view, c5) distal in the image (Pl. 1, Fig. 1a). The overall morphology is close view. d) TU 739151 left M2; occlusal view. to that of the wolf (Canis lupus), but is somehow shorter and more robust. There is a long diastema between the p4 and the alveolus for the p1 without the alveoli for p2–p3. The margo interalveolaris is sharp, and in lingual view it presents the hypoconid is aligned with those of the protoconid and a concave profile (Pl. 1, Fig. 1a, b), common in species paraconid, forming the carnassial blade. The entoconid is with premolar reduction and presence of a long diastema, relatively large and located parallel to the hypoconid, and as occurs in Thaumastocyon bourgeoisi (see Stehlin and both cuspids are separated by a very narrow valley. A small Helbing 1925) or Pseudocyon? astrei Kuss, 1962. The canine cristid in the hypoconulid position connects the entoconid associated with the mandible is well developed, and presents with the hypoconid. The m2 is missing, but the alveoli are a sharp and irregularly crenulated distal cristid (Pl. 1, seen to present a moderate size. The alveolus for the m3 is Fig. 2). The p4 is small in relation to the m1, presenting a very small, with one single root. smooth paraconid. The protoconid is distally inclined, high TU 73917 left mandible with p4–m2 and alveoli for and with a smooth mesial cristid. The distal cuspid is well p3 and m3 (Pl. 2, Fig. 1). The presence of a possible p3 is individualized and the talonid is small. The lingual cingulum marked by a one-rooted alveolus, separated from the p4 by a is somewhat more developed in the distal part of the tooth, small diastema. Both the p4 and the m1 are morphologically where it is slightly expanded. The m1 is a sectorial tooth with close to the holotype. The m1 metaconid is reduced to a moderately laterally compressed paraconid and protoconid. small notch in the distolingual cristid of the protoconid. The paraconid mesial cristid is not totally vertical. The m1 The m1 talonid is also very short, with a small entoconid metaconid is a small bulge in the middle of the distolingual situated in its distolingual corner. The distal end of the protoconid cristid. The talonid is very short, and is mostly talonid is rectilinear. The cingula are very smooth. There is occupied by a tall and long hypoconid. The mesial cristid of a deep vertical occlusion facet in the labial wall, between 401 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC the protoconid and the hypoconid, as well as vertical wear in size. The protocone appears to be reduced, and its position facets on the labial wall of both the paraconid and the is somewhat distal. The paracone is coniform, with a very protoconid. The m2 exhibits a quadrangular shape, with a marked mesial crista presenting a clear distal inclination. complete trigonid, although the paraconid is low and very The metastyle is quite robust. reduced. Its labial cristid connects the mesial cristid of TU 73916 right M1 (Text-fig. 2b). The occlusal outline the protoconid, and appears to be labially attached to the is an isosceles triangle with the labial wall rectilinear, with a metaconid, in such a way that a well-defined mesiolingual smooth ectoflexus. The parastyle is quite well developed. The valley is formed among them. The protoconid is quite large, paracone is rather larger and higher than the metacone. The with a pyramidal morphology; it has three cristids, one metastyle is small. The labial cingulum is high and strong mesial, another distal (parallel to the molar axis) and the at the base of both the paracone and metacone, but weaker third one lingual, contacting the metaconid. Its labial wall in the ectoflexus area. The paraconule and the metaconule is basally widened. The metaconid is smaller and lower than are subdivided and well developed. The paraconule comes the protoconid, and is slightly displaced distally. The talonid into contact with a high cingulum, developed from the is short, with a low and wide hypoconid. The entoconid is parastyle, whereas the metaconule connects with a small also low, and is located in the distolingual corner. Between cusp associated with the metastyle cingulum, which is these two cusps, a narrow and relatively deep valley can be very weak lingually. The protocone is situated in a central observed. The cingula are smooth. The presence of a small position relative to the labial wall; it is strong, with a m3 is attested by a small distal alveolus, partially broken. duniform shape and completely encircled by a strong labial The m3 is represented in some specimens TU 739118 (Pl. cingulum, which reaches the bases of the paraconule and 3, Fig. 1), TU739111 (Pl. 3, Fig. 4); it is morphologically metaconule. It is thickened at its distolingual corner. The close to the m2, but with less defined cuspids and an overall lingual part of the molar is clearly reduced compared to smaller size. the labial one. In the distal wall there is a strong inflexion Some morphological variations can be recognized in at the position of the metaconule, while the mesial wall is other specimens; for example, TU 7391147, a right mandible more rectilinear. The central valley of the trigone is deep, (Pl. 3, Fig. 3), lacks the parastylid in the p4 and its distal cusp revealing a circular wear pattern in the centre, resulting from is somewhat stronger; the m1 retains a small metaconid, the occlusion with the m1 hypoconid. An additional right and the mesial cristid of the paraconid is slightly displaced M1, TU 739129, is poorly preserved; it is somewhat smaller lingually and highly marked. Furthermore, the labial wall but morphologically similar to the other molar. of the hypoconid is somewhat basally expanded, developing TU 739130 right M2 (Text-fig. 2c). It presents an oval an incipient cingulum. The m2 represents quite a different shape and small size in relation to the M1. Its basal cingulum morphotype, in particular due to the greater reduction of the surrounds the complete molar, thickening at the distolingual talonid, which is almost exclusively built by the hypoconid; corner. Both the paracone and the metacone are low and this morphology is also observed in the left m2 TU 939151 somewhat lingually displaced. Both parastyle and metastyle (Pl. 3, Fig. 5). The m1 of the specimen TU 739111 (Pl. 3, are small. The protocone is poorly developed; it is located in a Fig. 4) has a small metaconid, but better developed than that central position and joined by a long crista to the paraconule. of the previously mentioned specimens, as occurs with the The metaconule is included in the distal cingulum, and does mesial cristid of the paraconid, which is well-marked and not make contact with the protocone. There is a small cusp tends to be in a more lingual position. lingual to the metacone, associated with the metastyle. The Upper dentition. TU 739130 right P4, elongated upper trigone valley is broad and flat. Specimen TU 739136 is also carnassial (Text-fig. 2a). The paracone and protocone are an M2, which is similar in shape to TU 739130, but smaller partially broken. There is a well-developed lingual cingulum. in size (Text-fig. 2d). The parastyle is present and individualized, but still moderate D i s c u s s i o n . The hypercarnivorous character of Peignecyon felinoides is unquestionable; it is manifested in Table 2. Measurements (in mm) of the lower teeth and upper the strong development of the sharp mandibular diastema, teeth of Peignecyon felinoides n. gen. et n. sp., from Tuchořice, associated with the reduction of the anterior premolars, the Czech Republic. Abbreviations: L – antero-posterior p4 high, with a distally inclined mesial cristid, carnassials diameter, W – width, *estimated measure, **alveolus measure. with sectorial morphology, characterized by the reduction of the m1 metaconid and the strong development of the Lower teeth Lp4 Wp4 Lm1 Wm1 Lm2 Wm2 Lm3 hypoconid, and in the elongation of P4 with an incipient TU 739115 14.2 7.3* 27.9 12.5* 12** 8** parastyle. The M1 presents a triangular occlusal shape and TU 73917 14.9 7.9 27 12 15.4 11.2 a very reduced lingual area. The second molars, m2 and TU 739118 30 13.6 M2, respectively, are smaller in relation to the m1 and M1. TU 7391147 16.2 8.2 29.3 13.2 14.4 10.8 Some of these characteristics are also found in Ysengrinia TU 739115 27.3 12.6 15.5 11.2 gerandiana (Viret 1929a, Ginsburg 1965) from Saint Upper teeth LP4 WP4 LM1 WM1 LM2 WM2 Gérand le Puy (France), early Miocene (MN 2). However, in the latter species, these hypercarnivorous characters TU 739130 29.6 14.8* 12.9 17.7 are less developed. Among other traits, it possesses a less TU 73916 20.4 24.8 reduced lower premolar row with shorter diastemas; the p4 TU 739129 19.1* 23.6 has the principal cusp smaller and more vertical, with no TU 739136 11.2 15.9 distal inclination of the mesial cristid; the m1 is more robust 402 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC and retains a strong metaconid. Additionally, the talonid is A second genus Crassidia was included close to wider than the trigonid, and its hypoconid and entoconid are Ysengrinia in the Ysengrini tribe by Heizmann and Kordikova comparatively less developed. These differences between (2000). Peignecyon felinoides differs from other species both amphicyonids are noteworthy, and suggest a clear grouped in the genus Crassidia (Heizmann and Kordikova taxonomic separation between them. 2000) in the general gracility of its dentition. Significant Peignecyon felinoides differs from other species included differences are also seen in the morphology of the M1, which in the genus Ysengrinia, such as Ysengrinia tolosana in the holotype of Crassidia intermedia from Ulm has a very (Noulet, 1876) and Ysengrinia depereti (Mayet, 1908), in developed lingual area, as occurs in the maxilla fragment of its smaller dentition, a more gracile m1 with a more reduced Selles-sur-Cher (Heizmann and Kordikova 2000: pl. 6); this metaconid, and the m2, which has lower and smaller cuspids. specimen shows the presence of an M2, which despite the The absence of upper dentition for these taxa severely limits buccal wall being broken, is large and shows a subquadrated the comparisons. However, the differences in relation to occlusal shape, in which the major axis is buccolingual, P. felinoides are more pronounced than those indicated for as is characteristic of Y. americana. Crassidia intermedia Y. gerandiana. appears in our phylogenetic analysis (Text-fig. 3) as the The species Ysengrinia americana (Wortman, 1901) sister group of the rest of the Thaumastocyoninae, close is much better represented; it was originally described as to Y. gerandiana. Peignecyon felinoides differs from these Amphicyon, and included by Hunt (2002) in this European two genera in a suite of advanced features, such as the genus. The relationship between European and American loss of the m1 metaconid and the strong reduction of the species has been exhaustively discussed by this author, M1 lingual area, which it shares with the most specialized who finds obvious affinities between Y. americana and Thaumastocyoninae genera, Tomocyon and Thaumastocyon. Y. tolosana, and between the latter two and Y. gerandiana. Tomocyon grivensis from the late middle Miocene of La However, there are notable differences in the upper Grive (France) differs from P. felinoides in the following dentition, which obscure these relationships. The M1 of characters: a p4 exhibiting a higher protoconid and a much Ysengrinia americana shows a strong reduction of the stronger distal cingulum; m1 long, with a sharp mesial lingual area, similar to that observed in Y. gerandiana and paraconidcristid, and buccolingually compressed, like P. felinoides, but the lingual cingulum does not completely the protoconid and talonid. The distolingual cristid of the encircle the protocone as it does in these two European protoconid is very prominent and has no metaconid, while species. On the contrary, this cingulum is restricted to the the hypoconid has a strong basal cingulum extending distally distolingual margin, so that the mesial wall of the protocone beyond the distal edge of the tooth; in the m2 the metaconid is forms part of the wall itself, and its cingulum is only a small located distally to the very large and high protoconid, so that ridge. Additionally, the M1 of P. felinoides has buccal cusps the trigonid valley is reduced and mesio-lingually located, that are much higher than in the American species, and the the talonid is narrow and similar to what it is observed in the paraconule and metaconule are coniform and subdivided, m1, the base of the hypoconid is distally expanded; the P4 is whereas in Y. americana they form crests and are attached longer, with a strong parastyle and a more distal protocone; to the protocone. In the M2, the differences are even more the M1 exhibits a wider lingual area; consequently, pronounced; in Y. americana the molar is morphologically its occlusal shape is subquadrangular, the paracone is close to the M1, but the protocone is completely surrounded much higher and more developed than the metacone, the by the lingual cingulum, which is comparatively better metaconule is poorly differentiated and the lingual cingulum developed. In Y. gerandiana and P. felinoides, the M2 is very is stronger, especially in the distolingual corner; in the m2 the reduced lingually, and the molar is therefore oval, and the protocone connects both the metaconule and the paraconule, protocone is very reduced. Thus, in both European species, forming a continuous semicircular crista, which joins the the M2 has a similar overall shape of the M3 to Y. americana buccal cingulum. Many of these differences indicate that and other amphicyonid species. This fact can be related to the T. grivensis possesses a better developed sectorial dentition strong tendency of Y. gerandiana and P. felinoides to reduce compared to that of P. felinoides. However, in the M1, it the crushing molar, whilst in Y. americana, this feature is presents a larger lingual area in contrast to the reduction less pronounced. The lower dentition of P. felinoides also observed in the Thaumastocyon species; this is also observed differs from that of Y. americana in the greater gracility in the m2, which retains a relatively large size. of the p4 and the m1. Considering the current data, we The other genus is Thaumastocyon, with two species, conclude that Y. americana cannot be included within the Th. bourgeoisi (type species) from the middle Miocene genus Ysengrinia. Consequently, Y. tolosana and Y. depereti, (MN 5) and Th. dirus from the late Miocene (MN 9) of only represented by their lower dentition, also remain in an Spain (Ginsburg et al. 1981). As we have already pointed uncertain systematic position, awaiting documentation on out, Thaumastocyon is characterized by extreme reduction their upper dentition. of the M1 lingual cingulum feature, which is absent in all Although in our analysis, Y. americana could be the other members of this subfamily. The holotype of Th. considered as the sister group of the Thaumastocyoninae bourgeoisi Sthelin et Helbing, 1925 is a mandible (Pont- (Text-fig. 3), alternatively, it could be related to other Levoy 69) preserving the canine and the m1–m2. The m1 hypercarnivorous Amphicyoninae that have retained a has more compressed buccolingual cuspids than those of well-developed M2 and a reduced p4 (Peigné et al. 2008, P. felinoides; moreover, the reduction of the metaconid Morales et al. 2016). These Amphicyoninae require an in- is complete, although a tiny notch in the distolingual depth taxonomic and systematic review, but this is not the cristid marks its ancestral position; the distobuccal base objective of the present paper. of the hypoconid is more swollen and projected distally. 403 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC Text-fig. 3. Phylogenetic relationship of Peignecyon felinoides n. gen. et n. sp., within some selected Amphicyonidae, and some extinct caniform carnivorans. Paramiacis exilis is the outgroup. Searches were performed by means of the Branch and Bound and a Bootstrap analysis through 1,000 replicates. One tree is obtained (length 73 steps, consistency index (CI) = 0.6301, retention index (RI) = 0.7000). The numbers below nodes are Bremer indices, and the numbers above nodes are Bootstrap support percent- ages (only shown ≥ 50). The m2 is morphologically similar to the morphotype, of Th. bourgeoisi, although we have very little material to Tuchořice, which presents a reduced talonid. Furthermore, ascertain its systematic position. Thaumastocyon dirus is somewhat larger, and differs from the type species of the genus in the almost total loss of the M1 lingual cingulum. Phylogenetic approach Ysengrinia valentiana from the locality of Buñol (Spain), Our cladistics analysis recovers a major clade: the early Miocene (MN 4) (Belinchón and Morales 1989) is quite Thaumastocyoninae, comprising Crassidia intermedia, interesting, although the species is represented by only two Ysengrinia gerandiana, Peignecyon felinoides, Tomocyon teeth: a left M1 (smaller in size that the M1 of P. felinoides), grivensis, Ysengrinia valentiana and Thaumastocyon spp., which displays a reduced lingual area, with a weak lingual is relatively well supported (Text-fig. 3). The subfamily cingulum distally placed. The protocone, although still Thaumastocyoninae is characterized by the start of an m2/M2 in a central position, presents a mesial crista that forms reduction, still moderate in Crassidia intermedia, but part of the mesial wall, similar to that of Y. americana. remarkable in the other species of the clade. In successive Additionally, the reduction of the labial cingulum, with the dichotomies, gradual progress can be observed towards consequent buccolingual shortening of the molar, clearly the development of hypercarnivorous dentition. First, it is differentiates Y. valentiana from the American species. The manifested in the specialization of the carnassial teeth, with second tooth of Y. valentiana, left m2, is morphologically the reduction of the P4 protocone and the m1 metaconid. close to that of P. felinoides (mandible TU 73917; Pl. 2, Next is the appearance of Peignecyon felinoides, which Fig. 1); it presents a strong wear facet in the mesiobuccal already shows the advanced features defining the group, such corner, which perfectly fits with a wear facet on the lingual as the loss of the m1 metaconid and the strong reduction wall of the M1 metacone. In Y. valentiana, the molar of the M1 lingual area. This species appears as the sister morphology is close to Thaumastocyon bourgeoisi, but the group of Tomocyon and Y. valentiana + Thaumastocyon spp. Spanish species still retains a larger M1 lingual cingulum. Consequently, it can be considered to constitute a link between At least regarding the M1, Y. valentiana certainly appears to these taxa, and Y. gerandiana and C. intermedia. Apparently, occupy an intermediate position between P. felinoides and Tomocyon grivensis has retained this intermediate grade 404 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC Table 3. Character matrix of the taxa analysed (see Table 1 for latter in a more advanced node than that of the French the corresponding list of characters and character states). species. Ysengrinia valentiana could possibly be better classified as Thaumastocyon than as Ysengrinia, but there is a need for additional material to confirm this hypothesis. Ysengrinia americana is also classified separately from the other Ysengrinia species considered in the analysis; its classification to the genus Ysengrinia is problematic, as we have previously discussed. Moreover, our phylogenetic hypothesis supports a sister relationship between Thaumasto- cyoninae and Y. americana, although support for this hypo- thesis is low (Text-fig. 3). Another species included in the analysis, Daphoenodon superbus, is placed close to Y. americana, but presents certain more plesiomorphic characters, retaining a complete premolar row showing no teeth size reduction, a short P4 with strong mesial protocone or a large M1 (Hunt 1998); however, it also presents a notable size reduction of M2–M3/m2–m3, which could indicate a clear divergence between them. Cynelos lemanensis represents a species with no hypercarnivorous characters in the dentition, and shows an opposite trend to Thaumastocyoninae and other taxa included in the analysis; this species tends to increase the crushing surface of the molars. Finally, Pseudocyonopsis spp. appears far from the Thaumastocyoninae species and consequently, the similarities with Ysengrinia gerandiana are only the result of shared plesiomorphic characters. Acknowledgements We would like to thank the curators and collection managers of all the institutions visited. We also wish to thank Dr. Robert M. Hunt Jr. (University of Nebraska State Museum) for providing casts of Ysengrinia americana and Daphonoedon superbus. O. F. and J. W. were granted funding by the Ministry of Culture of the Czech Republic (DKRVO 2019–2023/2.V.a, National Museum, 00023272). The Spanish Research Projects CGL 2015-68333-P and CGL2016-76431-P (MINECO/ FEDER-UE), the Research Groups CSIC 64 1538 and CAM-UCM 910607, and the Generalitat de Catalunya (CERCA Programme, and Beatriu de Pinós contract 2017 BP 00223 from AGAUR to J.A.) also funded aspects of this characterizing P. felinoides, although some of the m1 from research. Additionally, this research received support from La Grive show significant size development, associated with the SYNTHESYS Project http://www.synthesys.info/, which prominent and cutting cristids. Nevertheless, Y. valentiana is financed by European Community Research Infrastructure displays a more advanced reduction of the crushing molars, Action under the FP7 “Capacities” Program under the grant with an m2 talonid and M1 lingual area reduced in size. This agreement (SYNTHESYS; CZ-TAF-3329) of J.A. The support trend is even more evident in Th. bourgeosi, whose M1 barely of the DST-NFR Centre of Excellence in Palaeosciences retains a tiny lingual cingulum, which is completely lost in Th. (CoE-Pal) toward this research for A. V. (COE2018-09POST dirus, so that the molar comprises only the trigone, in which and COE2019-PD07) is hereby acknowledged. Opinions the protocone is quite small. However, the clade comprising expressed and conclusions reached are those of the author and Y. valentiana + Thaumastocyon spp. + Tomocyon is better are not necessarily to be attributed to the CoE. supported by bootstrap results and Bremer indices (Text-fig. 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(1846): Mémoire pour servir à la géologie record of a haplocyonine amphicyonid in early Miocene paléontologique des terrains tertiaires du département de of Nei Mongol fills a long-suspected geographic hiatus. l’Allier. – Bulletin de la Société géologique de France, – Vertebrata PalAsiatica, 54: 21–35. 3: 353–373. Wortman, J. L. (1901): A new American species of Amphi- Schaller, O. (2007): Illustrated Veterinary Anatomical No- cyon. – American Journal of Science, 11: 200–204. menclatures, 2nd ed. – Enke Verlag, Stuttgart, 614 pp. https://doi.org/10.2475/ajs.s4-11.63.200 407 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC Explanations to the plates PLATE 1 Peignecyon felinoides n. gen. et n. sp., from Tuchořice, the Czech Republic 1. Right mandible (holotype: NM-Pv 11600 right mandi- ble with c, p1 alveolus, p4–m1, and alveoli for m2–m3); a) buccal view, b) p4–m1 in lingual view, c) occlusal view. 2. Canine of the holotype mandible; a) buccal view, b) dis- tal view, c) lingual view, d) distal cristid in lingual view. PLATE 2 Peignecyon felinoides n. gen. et n. sp., from Tuchořice, the Czech Republic 1. Left mandible with p3 alveolus, p4–m2, and alveolus for m3 (TU 73917); a) buccal view, b) occlusal view, c) lin- gual view. 2. Left mandible with p4 (TU 739137); a) buccal view, b) occlusal view, c) buccal view. PLATE 3 Peignecyon felinoides n. gen. et n. sp., from Tuchořice, the Czech Republic 1. Right mandible with m1–m2 (TU 739118); a) buccal view, b) occlusal view, c) lingual view. 2. Left mandible with p4–m1 (TU 739115); a) buccal view, b) occlusal view, c) buccal view, d) distal view. 3. Right mandible with p4–m2 (TU 7391147); a) lingual view, b) occlusal view, c) buccal view, d) distal view. 4. Right mandible with m1–m3 (TU 739111); a) buccal view, b) occlusal view, c) lingual view. 5. Left m2 (TU 739151); a) occlusal view. 408 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC PLATE 1 409 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC PLATE 2 410 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC PLATE 3 411 Unauthentifiziert | Heruntergeladen 19.03.20 10:47 UTC