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New Material and Revision of the Carnivora, Mammalia from the Lower Pleistocene Locality Apollonia 1, Greece

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New Material and Revision of the Carnivora, Mammalia from the Lower Pleistocene Locality Apollonia 1, Greece quaternary Article New Material and Revision of the Carnivora, Mammalia from the Lower Pleistocene Locality Apollonia 1, Greece George D. Koufos Aristotle University of Thessaloniki, Department of Geology, Laboratory of Geology and Palaeontology, GR-54124 Thessaloniki, Greece; [email protected]; Tel.: +30-2310-998464 Academic Editor: Maria Rita Palombo Received: 19 March 2018; Accepted: 14 May 2018; Published: 17 May 2018 Abstract: During the last field campaigns in the mammal fossiliferous site Apollonia 1 (Macedonia, Greece), new carnivoran material has been discovered. The new collection added two new carnivoran taxa, Homotherium latidens and Panthera gombaszögensis. The new canid material and the revision of the old one (a) suggest the presence of two Canis species, C. etruscus and C. apolloniensis; (b) confirm the presence of the hypercarnivore Lycaon lycaonoides, and (c) allow for re-classifying the vulpine material to Vulpes praeglacialis. The taxonomic status of the species C. apolloniensis and Meles dimitrius is discussed. The composition and diversity of the Apollonia carnivoran assemblage are estimated and compared to those of various Greek and European Villafranchian ones. The results suggest close similarity to the Venta Micena (Spain) and Dmanisi (Georgia) carnivoran assemblages. The biochronological evidence indicates that Apollonia 1 is younger than Venta Micena and older than Untermassfeld (Germany), suggesting an age of 1.3–1.0 Ma. The study of the carnivoran guild structure of Apollonia 1 in comparison to the modern ones from known environments, as well as their functional morphology, suggest an open habitat, agreeing with previous interpretations. Keywords: carnivora; Early Pleistocene; Greece; systematics; biochronology; palaeoecology 1. Introduction The mammal fossiliferous site Apollonia 1 (APL) has been known since the beginning of the 1990s, when it was discovered in the Mygdonia Basin (Macedonia, Greece) [1]. The site provided a rich and interesting fauna with several taxa. The first excavations in the locality were carried out during the period 1990–1996 and the collected material was published in a series of articles [2–5]. Ten taxa were identified in the APL carnivoran assemblage: Ursus etruscus, Canis etruscus, Canis arnensis, Canis apolloniensis, Xenocyon sp., Pachycrocuta brevirostris, Vulpes alopecoides, Meles dimitrius, Megantereon cultridens, Lynx issiodorensis [2,4]. The fossiliferous site APL is located near the village of Nea Apollonia, about 65 km northwest of Thessaloniki (Figure1). The fossils were found into the Platanochori Formation, which overlies the Gerakarou Fm and crops out in the southeastern part of the Mygdonia Basin; the Platanochori Fm consists of grey-white, grey-green fluvio-lacustrine to lacustrine deposits [1,6,7]. The APL biochronological data suggest an Epivillafranchian age, more precisely ranging from 1.2 to 1.0 Ma. More recently (2011–2016), new field campaigns enriched the APL collection with new material, which includes some interesting carnivoran remains. In this article, the new material is described and compared with the old one and that from Eurasian Villafranchian localities. New taxa are added to the faunal list and the presence of others has been confirmed. The study of the new material and the revision of the old collection improve the knowledge of the systematics of the carnivoran fauna and provide additional evidence on the APL age and palaeoenvironment. Quaternary 2018, 1, 6; doi:10.3390/quat1010006 www.mdpi.com/journal/quaternary Quaternary 2018, 2, x FOR PEER REVIEW 2 of 37 revisionQuaternary of2018 the, 1 ,old 6 collection improve the knowledge of the systematics of the carnivoran fauna2 and of 38 provide additional evidence on the APL age and palaeoenvironment. Figure 1. Map indicating the Neogene–Quaternary deposits and the fossiliferous sites in the Figure 1. Map indicating the Neogene–Quaternary deposits and the fossiliferous sites in the Mygdonia Mygdonia Basin (Macedonia, Greece). APL: Apollonia 1; GER: Gerakarou 1; KAL: Kalamoto; TSR: Basin (Macedonia, Greece). APL: Apollonia 1; GER: Gerakarou 1; KAL: Kalamoto; TSR: Tsiotra Vryssi. Tsiotra Vryssi. Map provided by Dr. D. Mountrakis (Aristotle University of Thessaloniki, Department Map provided by Dr. D. Mountrakis (Aristotle University of Thessaloniki, Department of Geology). of Geology). 2. Materials and Methods 2. Materials and Methods The studied material is stored in the Laboratory of Geology and Palaeontology, Aristotle The studied material is stored in the Laboratory of Geology and Palaeontology, Aristotle University of Thessaloniki (LGPUT), together with the old APL collection. All material (new and old University of Thessaloniki (LGPUT), together with the old APL collection. All material (new and old collection) has been measured using a digital caliper with an accuracy of 0.1 mm. All measurements collection) has been measured using a digital caliper with an accuracy of 0.1 mm. All measurements are given in mm; estimated values into brackets. The upper and lower teeth are symbolized with are given in mm; estimated values into brackets. The upper and lower teeth are symbolized with capital and lowercase letters, respectively. The software Excel 10 (Microsoft, Redmond, WA, USA) capital and lowercase letters, respectively. The software Excel 10 (Microsoft, Redmond, WA, USA) is is used for the scatter diagrams and PAST 3.1 [8] for the box-plots, cluster analysis, and principal used for the scatter diagrams and PAST 3.1 [8] for the box‐plots, cluster analysis, and principal component analysis. component analysis. Abbreviations Abbreviations Localities. APL: Apollonia-1, Mygdonia Basin, Greece; DFN: Dafnero 1, Western Macedonia, Localities. APL: Apollonia‐1, Mygdonia Basin, Greece; DFN: Dafnero 1, Western Macedonia, Greece; DMN: Dmanisi, Georgia; GER: Gerakarou, Mygdonia Basin, Greece; KAL: Kalamoto, Greece; DMN: Dmanisi, Georgia; GER: Gerakarou, Mygdonia Basin, Greece; KAL: Kalamoto, Mygdonia Basin, Greece; PIR: Pirro Nord, Italy; SES: Sesklon, Thessaly, Greece; TSR: Tsiotra Vryssi, Mygdonia Basin, Greece; PIR: Pirro Nord, Italy; SES: Sesklon, Thessaly, Greece; TSR: Tsiotra Vryssi, Mygdonia Basin, Greece; UMF: Untermassfeld, Germany; VMC: Venta Micena, Spain; VOL: Volax, Mygdonia Basin, Greece; UMF: Untermassfeld, Germany; VMC: Venta Micena, Spain; VOL: Volax, Eastern Macedonia, Greece. Eastern Macedonia, Greece Institutes. LGPUT: Laboratory of Geology and Palaeontology, Aristotle University of Thessaloniki. Institutes. LGPUT: Laboratory of Geology and Palaeontology, Aristotle University of Measurements. B: breadth (maximum bucco-lingual diameter of the tooth); DAP: anteroposterior Thessaloniki. diameter (maximum mesio-distal diameter of the tooth); DT: transverse diameter; dist: distal; epiph: Measurements. B: breadth (maximum bucco‐lingual diameter of the tooth); DAP: anteroposterior epiphysis; H: height; mid-shaft: middle of the diaphysis; max: maximal; prox: proximal. diameter (maximum mesio‐distal diameter of the tooth); DT: transverse diameter; dist: distal; epiph: Morphological. a.a.c: anterior accessory cusp(-id); p.a.c: posterior accessory cusp(-id). epiphysis; H: height; mid‐shaft: middle of the diaphysis; max: maximal; prox: proximal. Morphological. a.a.c: anterior accessory cusp(‐id); p.a.c: posterior accessory cusp(‐id). Quaternary 2018, 1, 6 3 of 38 Quaternary 2018, 2, x FOR PEER REVIEW 3 of 37 3. Systematic Palaeontology 3. Systematic Palaeontology 3.1. Order Carnivora Bowdich, 1821; Family Ursidae Gray, 1865; Genus Ursus Linnaeus, 1858 3.1. Order Carnivora Bowdich, 1821; Family Ursidae Gray, 1865; Genus Ursus Linnaeus, 1858 Ursus etruscus Cuvier, 1823 Ursus etruscus Cuvier, 1823 Material.Material.New New collection: collection: Right Right radius, radius, APL-759. APL‐759. Measurements.Measurements. LL = 309.5 = 309.5 mm, DT mm,max. prox. DT =max. 31.2 prox.mm, DAP= 31.2max. prox. mm, = 21.4 DAPmm, DTmax.midshaft. prox. = 27.5= 21.4 mm, mm, DTmidshaft.DAPmidshaft.= 27.5 = 16.0 mm, mm, DAP DTmidshaft.max. dist. = =50.3 16.0 mm, mm, DAP DTmax.max. dist. dist.= 33.0= mm. 50.3 mm, DAPmax. dist. = 33.0 mm. Description.Description.The The radius radius APL-759 APL‐759 (Figure (Figure 22a,b)a,b) isis well preserved, preserved, lacking lacking a small a small part part of the of the proximalproximal diaphysis, diaphysis, which which was was broken broken during during its its discovery; discovery; however, however, plaster was was added added in in the the field field in orderin to order preserve to preserve its original its original height height and and the the connection connection withwith the the proximal proximal epiphysis, epiphysis, which which is well is well preserved.preserved. The The diaphysis diaphysis is slender,is slender, slightly slightly curved curved and flattened flattened in in the the dorso dorso-palmar‐palmar direction. direction. The The proximal articular facet is sub‐elliptical, with the medio‐lateral diameter longer than the dorso‐ proximal articular facet is sub-elliptical, with the medio-lateral diameter longer than the dorso-palmar palmar one. The distal epiphysis is wider than the proximal one, bearing a large concave distal one. The distal epiphysis is wider than the proximal one, bearing a large concave distal articular facet. articular facet. Figure 2. (a,b) Ursus etruscus, right radius, APL‐759; (a) palmar, and (b) dorsal view. (c,e) Vulpes Figure 2. (a,b) Ursus etruscus, right radius, APL-759; (a) palmar, and (b) dorsal view. (c,e) praeglacialis,
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