Bartolini Lucenti, S.

Rook, L.

English

Florence Research

     Available online http://amq.aiqua.it ISSN (print): 2279-7327, ISSN (online): 2279-7335    Alpine and Mediterranean Quaternary, Vol. 31 (Quaternary: Past, Present, Future - AIQUA Conference, Florence, 13-14/06/2018), 169 - 172 THE FOSSIL RECORD OF THE GENUS CANIS (CANIDAE, CARNIVORA, MAMMALIA) FROM THE UPPER VALDARNO: A CRITICAL REVISION IN THE FRAME OF THE EARLY AND MIDDLE PLEISTOCENE CANIDS OF EURASIA  Saverio Bartolini Lucenti1,2, Lorenzo Rook2  1 Dottorato di Ricerca in Scienze della Terra, Università di Pisa, Pisa, Italy 2 Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Firenze, Italy Corresponding author: S. Bartolini Lucenti <saverio.bartolini@dst.unipi.it>   ABSTRACT: The outstanding fossil record of the Upper Valdarno Basin is renown since the XV century. Its Early Pleistocene sample of canids, characterized by the large sized Lycaon falconeri and the medium-sized Canis arnensis and Canis etruscus, has laid the basis for the study of the evolution of the genus Canis in Western Eurasia. In recent years, several discoveries tend to modify our conception on this subject, e.g. the lineage that leads to modern wolves or the inter/intraspecific variability of fossil species, but the record of UV Basin remains one of the most extensive and important of its age.  KEYWORDS: Upper Valdarno, Pleistocene, Canidae, taxonomy, evolution 1. INTRODUCTION  In the international panorama of continental Quater-nary sites, the Upper Valdarno Basin is historically re-nowned from the Renaissance for its outstanding record of large mammals (Rook et al., 2013), and since the 18th century naturalists and early palaeontologists, above all G. Cuvier (Cuvier, 1812; 1821-1824), have studied the mammals recovered from this basin (Cioppi & Dominici, 2010). For the scholars who study the family Canidae, the significance of this basin emerged by the end of the 19th and the beginning of the 20th century (Forsyth Ma-jor, 1877; Del Campana, 1913) with the description of the medium-sized Canis arnensis Del Campana, 1913 and the large Lycaon falconeri (Forsyth Major, 1877). These two species, together with Canis etruscus For-syth Major, 1877 also recovered from the other Tuscan locality of Olivola, represented the earliest species of the genus to be described in Western Eurasia. The fau-nal assemblage of these localities suggested a correla-tion to the beginning of the Calabrian (i.e., at that time, the Plio-Pleistocene boundary) (Azzaroli, 1977), as con-firmed by some magnetostratigraphic calibrations (e.g. Napoleone et al., 2003). The appearance of Canis etruscus came to be known as the “Wolf event” (Azzaroli, 1983; Azzaroli et al., 1988; Rook & Torre, 1996). Other findings, e.g. those of Coste San Giacomo (2.1 Ma, Bellucci et al., 2012) and Vialette (disputed 3 Ma, Lacombat et al., 2008), have shown the diachronic state of the arrival of Canis taxa in Europe (Sotnikova & Rook, 2010 for further discussion). Never-theless, the record of Canis the Upper Valdarno still has to be regarded with great interest by international re-searchers for the implication its fossil have. For in-stance, C. etruscus is generally considered as the an-cestor of the lineage that leads to the modern wolf (Sotnikova, 2001; Brugal & Boudadi-Maligne, 2011). Moreover, several sites across Eurasia indicate the oc-currence of C. etruscus-like canids (see Cherin et al., 2014 and reference therein). On the contrary, C. arnen-sis seems limited to sites of “Tasso FU” faunal compo-nents (Torre et al., 1992). Interestingly, the former ar-gued that this species could be the European represen-tative of a holoarctic coyote-like population, with Canis lepophagus as American correspondent. During the Early Pleistocene the diversity of species of Canis is not limited to these two as Canis senezensis Martin, 1973 was described from Senèze (Massif Central, France; ca. 2 Ma) whereas in Spain, Canis accitanus Garrido & Ar-ribas, 2008 is reported from Fonelas P-1 (Guadix Basin, Granada ca. 1.9-1.7 Ma). Around 1.5-1.4 the first repre-sentatives of Canis mosbachensis Soergel, 1925 ap-pears in Spain and in Italy. Lastly, from the site of Apol-lonia comes the disputed Canis apollonensis Koufos & Kostopoulos, 1997. This variability of species decreases around the Epivillafranchian (ca 1 Ma) as only C. mos-bachensis remained in Western Europe (Sotnikova & Rook, 2010). In Asia, the pattern of decrease in diversity is similar. During the Early Pleistocene several medium- to large-sized species were present [Canis chihliensis Zdansky, 1924 and Canis palmidens (Teilhard de Char-din & Piveteau, 1930) from Nihewan Basin; Canis teil-hardi Qiu et al., 2004, Canis longdanensis Qiu et al., 2004 and Canis brevicephalus Qiu et al., 2004 from Longdan] but in the Middle Pleistocene the medium-sized and mesocarnivorous niche of canids was occu-pied by C. mosbachensis variabilis Pei, 1934 (Jiangzuo et al., 2018). This short review aims to resume past knowledge in the light of recent discoveries with unpublished material      that has significant implications for the phylogeny of this group of Carnivora.  2. MATERIAL AND METHODS  The material considered for the present research is housed at the Museum of Natural History of the Univer-sity of Florence. Comparative fossil and extant sample studied comes from various European sites (Fig. 1) and it is held in different European and North American institution as the Montevarchi Paleontological Museum; La Specola Zoology section of the Museum of Natural History of the University of Florence; the American Mu-seum of Natural History (New York); the Georgian Na-tional Museum (Tbilisi); the Aristotle University of Thes-saloniki; the Hungarian Museum of Natural History and the Geological Society of Hungary (Budapest); Earth Science department of the Université Claude Bernard Lyon-1. Furthermore all the relevant literature on fossil Canis has been reviewed. The morphometric analyses used in this study include log ratio diagrams (Simpson, 1941), box plots, and biplots. Those analyses were performed with PAST ver. 3.14 (Hammer, 2016). The measurements where taken to nearest of 0.1 mm fol-lowing Driesch (1976).  3. RESULTS  The morphometric and morphological comparison of the Tuscan material of C. etruscus and that of C. etruscus from Dmanisi showed important difference both in dental proportions and, more importantly, in cranial features e.g., the elongation of the skull in ros-trocaudal sense; the length of the nasal (Fig. 2); the height of the skull; the anteroposterior divergence of the medial wall of the tympanic bullae. The morphology of these characteristics in C. “etruscus” from Dmanisi are more similar to the other Early Pleistocene canid C. mosbachensis as well as to extant and more derived species e.g. C. lupus rather to the coeval C. etruscus or C. arnensis. Similar characteristics have been recog-nized in the sample of Gerakarou (Greece). Although the size is intermediate between C. arnensis and C. etruscus, the greek specimens show the same mor-phologies that characterize the material of Canis sp. from Dmanisi.  The analyses on several samples taxa from Early and Middle Pleistocene sites across Europe such as Venta Micena, Pirro Nord, Apollonia, Petralona, Vértesszȍlȍs II (Fig. 1) have revealed a wide range of variability in size but a general morphology shared by all these samples, all referable to C. mosbachensis-like canids.  4. DISCUSSION AND CONCLUSIONS  The fossils of C. arnensis and C. etruscus of the Olivola and Tasso FU represent one of the most exten-sive record of the genus Canis of the early Quaternary and among the earliest record in Western Europe. These samples remain pivotal basis for comparison in the attempt to resolve the intricate scenario of Early Pleistocene canids across Eurasia. The morphometric and morphological characteristics found in the Georgian Canis “etruscus” greatly contrast with the peculiarities of C. etruscus. Moreover, the Canis from Dmanisi presents   170   Bartolini Lucenti S., Rook L. Fig. 1 - Geographic localization of the sites considered in this study. 1, Venta Micena, Spain; 2, Cueva Victoria, Spain; 3, Vallparadís Estació-Cal Guardiola, Spain; 4, Olivola, Italy; 5, Upper Valdarno, Italy; 6, Pirro Nord, Italy; 7, Gerakarou, Apollonia-1, Petralona, Greece; 8, Dmanisi, Georgia; 9, Vértesszȍlȍs II, Hungary.      several cranial affinities to Eurasian C. mosbachensis and to other modern wolf-related species, for instance the antero-posteriorly divergent medial walls of the tym-panic bullae, as also noticed by Sotnikova & Rook (2010). Relevance of this feature stands in the fact that only some extant species but no fossil taxa present it, with the exception of C. mosbachensis (Bartolini Lu-centi et al., 2017; Jianzuo et al., 2018). Altogether this evidence leads us to consider this taxon as Canis sp. nov. The new information on the Dmanisi Canis sp. has strong implications on the evolutionary history of the Late Villafranchian canids, because it changes the widely accepted C. etruscus-C. mosbachensis-C. lupus lineage.  Although the status of C. mosbachensis is ques-tioned by some authors (for an updated resume see Mecozzi et al., 2017), the slight morphological differ-ences found in samples of Early to Middle Pleistocene sites attributed to C. mosbachensis-like canids should definitely be regarded as results of the intraspecific variability. Surely, such little variations are not sufficient to erect new species. Likewise, it would be misleading to interpret this diversity as part of the variability of the modern C. lupus. In this sense, the ascription of the Chinese taxon C. variabilis as a subspecies of C. mos-bachensis in Jiangzou et al. (2018) offers an interesting solution to the issue.  ACKNOWLEDGEMENTS The authors are thankful to all the curator of the institutions visited for granting us access to the collec-tion in order to study the variability of Early-Middle Pleis-tocene: Cioppi E.(IGF), Rustioni M. (MPM), Agnelli P. (MZUF), Galkin J., Meng J. (AMNH), Lordkipanidze D., Bukhsianidze M. (GNM), Koufos G. (AUT), Gasparik M. (HMNH), Robert E. (UCBL), Berthet D. (MdC).  REFERENCES  Azzaroli A. (1977) - The Villafranchian stage in Italy and the Plio-Pleistocene boundary. Giornale di Geolo-gia, 41, 61-79. Azzaroli A. (1983) - Quaternary Mammals and the End-Villafranchian Dispersal event - a turning point in the history of Eurasia. Palaeogeography, Pa-laeoecology, Palaeoclimatology, 44, 117-139. Azzaroli A., De Giuli C., Ficcarelli G., Torre D. (1988) - Late Pliocene to early mid-Pleistocene mammals in Eurasia: faunal succession and dispersal events. 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The fossil record of the genus Canis (Canidae, Carnivora, Mammalia) from the Upper Valdarno: A critical revision in the frame of the early and Middle Pleistocene canids of Eurasia

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The fossil record of the genus Canis (Canidae, Carnivora, Mammalia) from the Upper Valdarno: A critical revision in the frame of the early and Middle Pleistocene canids of Eurasia

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