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Homotherium latidens (Owen, 1846) Machairodus latidens Owen, 1846: 179. Homotherium latidens ��� Galobart et al. 2003: 99-141. HOLOTYPE. ��� The upper canine described and figured as Machairodus latidens by Owen (1846: fig. 69). In total there are five upper canines and two I3 from Kent���s Cave; the holotype, numbered 443/ 103 in the Museum of the Royal College of Surgeons, is that described by Owen, but it was destroyed during bombing in 1941 (Barnett 2014). TYPE LOCALITY. ��� Kent���s Hole, Torquay, England. LOCALITY. ��� Dafnero-3, DFN3, Grevena Basin, Western Macedonia, Greece. DESCRIPTION Cranium The cranium DFN3-152 is dorsoventrally compressed, deformed and damaged in the occipital region (Fig. 1). The dentition preserves both I3, the left canine and part of the left carnassial. The worn incisors and carnassial indicate an old individual. The premaxillae are well developed and projected rostrally to protrude the large incisors. The incisor row is semicircular as in the Sen��ze (France) cranium FSL- 210 991 (Ballesio 1963: fig. 11d) and the Incarcal (Spain) crania IN-I-929 and IN-II-37 (Galobart et al. 2003: figs 3, 4). The incisor row of the Perrier cranium of Homotherium (MNHN.F.PET2000) is less convex than that of DFN3-152, while that of IN-I- 825 from Incarcal is straight (Bonis 1976; fig. 3; Galobart et al. 2003: fig. 1). Between the canine and I3 of DFN3-152, there is a short diastema (4.3 mm) as in the Sen��ze (FSL-210 991; 9.0 mm), (Ballesio 1963) and Perrier crania (7.7mm; MNHN.F.PET2001). In the Incarcal crania the diastema varies greatly; it is long in IN-I929 and absent in IN-I-825 and IN-II-37 (Galobart et al. 2003). A large diastema (16.0 mm) also exists between the canine and the P 3 in DFN3-152, as in the Sen��ze cranium and the Incarcal crania IN- I-1929 and IN-I-825 (Table 1). The maxillary bones are relatively small and have a swelling that corresponds to the strong root of the upper canine. The nasals are rectangular, like in all machairodontines; their posterior margin is almost straight and their suture with the frontals is a straight line, which coincides with the line connecting the anterior margins of the orbits. This feature is clear in the Sen��ze and Perrier crania (Ballesio 1963: fig. 11d; Bonis 1976: fig. 9a). The nasal cavity is large, sub- rectangular and higher than wide; the index height/breadth of the nasal cavity is 89, versus 91 for the Sen��ze and 81-95 for the Incarcal crania (Galobart et al. 2003). The right orbit is better preserved and not deformed; it is rounded, with its anterior margin above the distal end of the carnassial. The infraorbital foramen is very large (Fig. 1A; Table 1). Its anterior margin is situated above the mesial part of the P3, while the posterior margin is above the parastyle of the carnassial. The zygomatic arches are small, relatively low and close to the braincase, starting just behind the distal end of the carnassial and directed slightly externally to connect with the glenoid fossa. The frontals are compressed and deformed; however, the groove between them and its continuity with the nasals is clearly distinguished. The sagittal crest is poorly preserved but based on its remnants seems to be thick and high. Although the development of the sagittal crest varies in the known material of Homotherium as a result of sexual dimorphism (Galobart et al. 2003), it is impossible to determine the sex of DFN3-152. The palate is elongated, wide and bears two strong crests running mesio-distally (Fig. 1D). The choanae are poorly preserved, deep and their anterior margin is situated behind the distal end of the carnassials. The mandibular fossa is long, with strong retroarticular process. The retroarticular foramen is situated in a large and deep fossa between the mandibular fossa and the tympanic bulla. The right part of the occipital is well preserved, having a rounded shape with strong nuchal and external occipital crests; the distal margin of the occipital bone does not exceed the level of the occipital condyles. Dentition The third incisors are worn and partially broken (Fig. 1D). They are large, canine-like, slightly inclined distally, with almost rounded occlusal outline, and bear a small lingual cusp. The canine is well preserved, long, narrow, sharp, inclined distally, like a double-edged knife. Although the wear is advanced, clear serration can be distinguished on its mesial and distal borders, like in Homotherium (Fig. 1E, F). The canine is strongly compressed laterally; the breadth/ length index is 42. The root is very strong and elongated, running parallel to the margin of the nasal cavity to the end of the nasal bone. There is no trace of the P2, while there is a clear alveolus for the P3; its dimensions are 10.5 �� 5.1 mm. The P4 is damaged and only the paracone and the worn metacone are preserved. The carnassial is elongated and relatively narrow. Although it is damaged there is evidence of a pre-parastyle, and the paracone is strong and high. There is no apparent protocone but as the tooth is very worn the presence of a small one as in all Homotherium, but totally worn down, cannot be ruled out. Humerus The humerus DFN3-153 is complete but the epiphyses are badly preserved (Fig. 2). The articular surface of the humerus caput is flattened, as in Homotherium from Incarcal (Anton et al. 2005). The major tubercle is well developed, and the articular head is wide and projected, like in the Incarcal Homotherium material. The intertubercular groove is wide. The diaphysis is long and straight; the deltoid ridge is strong and is not projected laterally. The distal epiphysis is wide and bears a large trochlea, and the olecranon fossa is deep., Published as part of Koufos, George D., Kostopoulos, Dimitris S. & Merceron, Gildas, 2020, The saber-toothed cat Homotherium latidens (Owen, 1846) from the lower Pleistocene locality Dafnero, Western Macedonia, Greece, pp. 139-149 in Geodiversitas 42 (10) on pages 142-144, DOI: 10.5252/geodiversitas2020v42a10, http://zenodo.org/record/3827749, {"references":["mental and paleoecological implications. Quaternary Geochro- nology 21: 2 - 15. https: // doi. org / 10.1016 / j. quageo. 2012.12.006 OWEN R. 1846. - A History of British Mammals and Birds. John van Voorst, London, 560 p. https: // doi. org / 10.5962 / bhl. title. 13248","GALOBART A., PONS- MOYA J., ANTON M. & MAROTO J. 2003. - mental and paleoecological implications. Quaternary Geochro- Descripcion del material de Homotherium latidens (Owen) de los nology 21: 2 - 15. https: // doi. org / 10.1016 / j. quageo. 2012.12.006 yacimentos del Pleistoceno inferior de Incarcal (Girona, NE de OWEN R. 1846. - A History of British Mammals and Birds. John van la Penisula Iberica). Paleontologia i Evolucio 34: 99 - 141. Voorst, London, 560 p. https: // doi. org / 10.5962 / bhl. title. 13248","BARNETT R. 2014. - An inventory of British remains of Homo- KOUFOS G. D. 2018. - New material and revision of the Carnivtherium (Mammalia, Carnivora, Felidae), with special reference ora, Mammalia from the Lower Pleistocene Apolonia 1, Greece, to the material from Kent's Cavern. Geobios 47: 19 - 29. https: // in PALOMBO M. - R. (ed.), Advances in Quaternary studies: The doi. org / 10.1016 / j. geobios. 2013.12.004 contribution of mammalian fossil record. Quaternary 1 (6): 1 - 38.","BALLESIO R. 1963. - Monographie d'un Machairodus du gisement basin (Macedonia, Greece). Annales de Paleontologie 78: 205 - 257. villafranchien de Seneze: Homotherium crenatidens Fabrini, 1890. KOUFOS G. D. 2001. - The Villafranchian mammalian faunas and Travaux du Laboratoire de Geologie de la Faculte des Sciences de biochronology of Greece. Bollettino della Societa Paleontologia Lyon, nouvelle serie, 9: 1 - 127. Italiana 40 (2): 217 - 223.","BONIS L. DE 1976. - Un Felide a longues canines de la colline de KOUFOS G. D. & KOSTOPOULOS D. S. 2016. - The Plio-Pleistocene Perrier (Puy-de-Dome). Annales de Paleontologie 62: 159 - 198. large mammal record of Greece: Implications for early Human","FABRINI E. 1890. - Machairodus (Megantereon) del Val d'Arno MOL D., LOGCHEM W. M. S. VAN, HOOIJDONK K. VAN & BAKKER superiore. Bolletino Reale Committee Geologica Italiana 1: 121 - R. 2008. - The Saber-Toothed Cat of the North Sea. DrukWare, 144 (https: // www. biodiversitylibrary. org / page / 53556172) and Norg, 160 p.","New Data from classic and Innovative Approaches, in 15 th International Congress of the Geological Society of Greece, 2 p. KOUFOS G. D. 1992. - The Pleistocene carnivores of the Mygdonia","PALMQVIST P., TORREGROSSA V., PEREZ- CLAROS J. A., MARTINEZ- NA- VARRO B. & TURNER A. 2007. - A re-evaluation of the diversity of Megantereon (Mammalia, Carnivora, Machairodontinae) and the problem of species identification in extinct carnivores. Journal of Vertebrate Paleontology 27 (1): 160 - 175. https: // doi. org / cfc 688","SARDELLA R. & IURINO D. A. 2012. - The latest early Pleistocene saber-toothed cat Homotherium (Felidae, Mammalia) from Monte Peglia (Umbria, central Italy). Bollettino della Societa Paleontologica Italiana 51 (1): 15 - 22.","ANTON M., GALOBART A. & TURNER A. 2005. - Co-existence of of Greece. Quaternaire, hors serie, 2: 179 - 190."]}

Genus Homotherium Fabrini, 1890 Homotherium Fabrini, 1890: 161. AGE. ��� Early Pleistocene (middle Villafranchian); 2.5-2.0 Ma. MATERIAL. ��� Cranium, DFN3-152; left humerus, DFN3-153. MEASUREMENTS (in mm). ��� The measurements of the cranium and humerus are given in Tables 1 and 2. The dental measurements are: I3 dex = 12.1 �� 11.3; I3 sin = 12.5 �� ���; C = 24.8 �� 11.9; P4 = 36.2 �� ���., Published as part of Koufos, George D., Kostopoulos, Dimitris S. & Merceron, Gildas, 2020, The saber-toothed cat Homotherium latidens (Owen, 1846) from the lower Pleistocene locality Dafnero, Western Macedonia, Greece, pp. 139-149 in Geodiversitas 42 (10) on pages 140-142, DOI: 10.5252/geodiversitas2020v42a10, http://zenodo.org/record/3827749, {"references":["FABRINI E. 1890. - Machairodus (Megantereon) del Val d'Arno MOL D., LOGCHEM W. M. S. VAN, HOOIJDONK K. VAN & BAKKER superiore. Bolletino Reale Committee Geologica Italiana 1: 121 - R. 2008. - The Saber-Toothed Cat of the North Sea. DrukWare, 144 (https: // www. biodiversitylibrary. org / page / 53556172) and Norg, 160 p.","mental and paleoecological implications. Quaternary Geochro- nology 21: 2 - 15. https: // doi. org / 10.1016 / j. quageo. 2012.12.006 OWEN R. 1846. - A History of British Mammals and Birds. John van Voorst, London, 560 p. https: // doi. org / 10.5962 / bhl. title. 13248","TEILHARD DE CHARDIN P. 1936. - Fossil mammals from Locality 9 of Choukoutien. Palaeontologica Sinica, ser. C, 7: 1 - 61.","BALLESIO R. 1963. - Monographie d'un Machairodus du gisement basin (Macedonia, Greece). Annales de Paleontologie 78: 205 - 257. villafranchien de Seneze: Homotherium crenatidens Fabrini, 1890. KOUFOS G. D. 2001. - The Villafranchian mammalian faunas and Travaux du Laboratoire de Geologie de la Faculte des Sciences de biochronology of Greece. Bollettino della Societa Paleontologia Lyon, nouvelle serie, 9: 1 - 127. Italiana 40 (2): 217 - 223.","BONIS L. DE 1976. - Un Felide a longues canines de la colline de KOUFOS G. D. & KOSTOPOULOS D. S. 2016. - The Plio-Pleistocene Perrier (Puy-de-Dome). Annales de Paleontologie 62: 159 - 198. large mammal record of Greece: Implications for early Human","GALOBART A., PONS- MOYA J., ANTON M. & MAROTO J. 2003. - mental and paleoecological implications. Quaternary Geochro- Descripcion del material de Homotherium latidens (Owen) de los nology 21: 2 - 15. https: // doi. org / 10.1016 / j. quageo. 2012.12.006 yacimentos del Pleistoceno inferior de Incarcal (Girona, NE de OWEN R. 1846. - A History of British Mammals and Birds. John van la Penisula Iberica). Paleontologia i Evolucio 34: 99 - 141. Voorst, London, 560 p. https: // doi. org / 10.5962 / bhl. title. 13248"]}

Genus Mammuthus Brookes, 1828 TYPE SPECIES. ��� Mammuthus primigenius (Blumenbach, 1799), fixed as type species by Garutt et al. 1990 (see also Reich et al. 2007 and Lister 2017)., Published as part of Konidaris, George E., Kostopoulos, Dimitris S. & Koufos, George D., 2020, Mammuthus meridionalis (Nesti, 1825) from Apollonia- 1 (Mygdonia Basin, Northern Greece) and its importance within the Early Pleistocene mammoth evolution in Europe, pp. 69-91 in Geodiversitas 42 (6) on page 72, DOI: 10.5252/geodiversitas2020v42a6, http://zenodo.org/record/3708332, {"references":["GARUTT V. E., GENTRY A. & LISTER A. M. 1990. - Mammuthus Brookes, 1828 (Mammalia, Proboscidea): proposed conservation, and Elephas primigenius Blumenbach, 1799 (currently Mammuthus primigenius): proposed designation as the type species of Mammuthus, and designation of a neotype. Bulletin of Zoological Nomenclature 47 (1): 38 - 44. https: // doi. org / 10.5962 / bhl. part. 2651","REICH M., GEHLER A., MOL D. & LISTER A. 2007. - The rediscovery of type material of Mammuthus primigenius (Mammalia: Proboscidea). Abstracts of the IVth International Mammoth Conference, Yakutsk: 190 - 191.","LISTER A. M. 2017. - On the type material and evolution of North American mammoths. Quaternary International 443: 14 - 31."]}

Mammuthus meridionalis (Nesti, 1825) LECTOTYPE. ��� Cranium with M3, IGF-1054 (Natural History Museum, Geology and Paleontology Section, University of Florence, Italy), designated by Dep��ret & Mayet (1923: 126). TYPE LOCALITY. ��� Upper Valdarno (Italy)., Published as part of Konidaris, George E., Kostopoulos, Dimitris S. & Koufos, George D., 2020, Mammuthus meridionalis (Nesti, 1825) from Apollonia- 1 (Mygdonia Basin, Northern Greece) and its importance within the Early Pleistocene mammoth evolution in Europe, pp. 69-91 in Geodiversitas 42 (6) on page 72, DOI: 10.5252/geodiversitas2020v42a6, http://zenodo.org/record/3708332, {"references":["NESTI F. 1825. - Lettere sopra alcune ossa fossili del Valdarno non per anco descritte sulla nuiva specie di elefante fossile del Valdarno. Presso Sebastiano Nistri, Pisa, 24 p.","DEPERET C. & MAYET L. 1923. - Les elephants pliocenes. Deuxieme partie: Monographie des elephants pliocenes d'Europe et de l'Afrique du Nord. Annales de l'Universite de Lyon. Nouvelle serie, I, Sciences, Medecine 42: 89 - 224."]}

Mammuthus meridionalis vestinus (Azzaroli in Ambrosetti, Azzaroli, Bonadonna & Follieri, 1972) (Figs 2-4) HOLOTYPE. ��� Skeleton including the skull with the m3/M3 (Spanish Fortress, L���Aquila, Italy). TYPE LOCALITY. ��� Madonna della Strada (Scoppito, L���Aquila, Italy). LOCALITY. ��� Apollonia-1 (Mygdonia Basin, Macedonia, Greece). AGE. ��� Late Villafranchian (Early Pleistocene). MATERIAL EXAMINED. ��� Right maxilla fragment with DP2-DP3, APL-225; right M3 fragment, APL-686A; left M3 fragment, APL- 686B; right hemi-mandible with m3, APL-716; left m3, APL-687. DESCRIPTION The right maxilla fragment APL-225 preserves the DP2 and DP3 (Fig. 2). The DP2 has rectangular shape and is formed of four, only slightly worn plates, with the posterior one attached to the third one. Cementodonty is present, but weak. The still not completely erupted DP3 consists of eight mostly unworn plates, which are parallelsided and separated. The tooth presents a weak lingual curvature. In lingual view, the first two plates are slightly inclined relative to the occlusal surface, whereas the rest are almost vertical. APL-686B (Fig. 3 A-C) is identified as upper molar based on the obtuse angle that is formed between the occlusal surface to the vertical plane of the lamellae, and on the rootward tilt for the last 6-7 lamellae (Lister et al. 2013). It represents the third (last) molar due to the tapering form at the posterior end of the crown (Lister et al. 2012). The plates were found loose and isolated in the sediment, and they were subsequently glued together. Preserved are thirteen plates, although at least one more should have been anteriorly, increasing thus the PN to possibly 14. The tooth is curved at the lingual side. In labial and lingual views, the plates are slightly inclined relative to the horizontal occlusal surface, and they are rather straight. The only complete enamel loop is the anterior-most preserved one. The enamel loops at the anterior part of the tooth are parallelsided, separated, perpendicular to the long axis of the tooth, and their lateral sides are rounded and evenly directed (not curving anteriorly or posteriorly). Apical digitations (appearing on the occlusal surface as rounded loops) are present at the third and fourth preserved lamellae. The fourth lamella, which is at an early wear stage, shows four subequal rings. The enamel folding is weak. The last eight plates are ovoid of wear.APL-686A corresponds to a fragmented upper molar and preserves only four plates. It was found close to APL-686B and represents possibly its right counterpart. The hemi-mandible APL-716 preserving the m3 (see below) lacks the anterior part of the ventral border of the corpus, the condyle and the coronoid process (Fig. 4). The mandibular symphysis is not well-preserved, but its preserved dorsal surface is straight and steeply directed downwards. In lateral and medial views, the hemi-mandible is relatively short and deep. The corpus is moderately swollen and more evident in the medial side. In lateral view, the depth of the corpus is higher at the level of the anterior part of the m3. The lateral side of the ramus is flat. Corpus and ramus form obtuse angle due to the posterior inclination of the ramus. The rostral margin of the ramus is positioned at a minor obtuse angle, but almost vertical, to the alveolar planum. Two anterior mental foramina (AMF) are located at the posterior border of the symphysis, at the level of the anterior end of the m3, and a large medial mental foramen (MMF) is present approximately at the same level as the AMF. Measurements [in mm; according to Beden (1983)], are A = (220); B = (120); C = 260; F = 150 and J = (225). The molar on the hemi-mandible APL-716 (Fig. 4) is considered as the third one due the long antero-posterior dimension (length) and the narrowing of the crown towards the posterior end. The anterior part is worn down to a dentine platform and the original plate number can only be estimated, due to the wear of the enamel loops. However, the rostral part of the anterior root is visible, indicating that no substantial loss should had happened due to wear, which resulted possibly in the loss of only the anterior talon (Lister & Sher 2015). Therefore, the number of plates is estimated as 12, plus a platelet/talonid at the posterior end. The shape of the molar is elliptic, being wider in the middle, and with a minor curvature at the labial side. In lingual view, the plates are slightly inclined relative to the occlusal surface. In labial view, the tooth shows a slight depression at the middle. Cement deposition is strong on the occlusal surface, as well as on the labial, lingual and posterior sides; however, the enamel is high above the cement. The enamel loops (five complete at the anterior part) are parallel-sided and separated. Anteriorly, the loops are oblique relative to the long axis of the tooth and become perpendicular to that at the posterior end. Apical digitations are few and present posteriorly. The enamel folding is undulating and present at the entire length of the enamel figure. The lateral sides of the enamel figures are rounded and directed anteriorly, stronger on the lingual side. The identification of the APL-687 (Fig. 3 D-F) as a third lower molar is based on the significant preserved length despite the advanced dental wear (a condition possible only in the last molars when the forward progression of them has been terminated), and on the presence of a single root; the anterior first and second roots had been lost, and therefore a substantial part of the original anterior end of the tooth is missing (Lister et al. 2012; Lister & Sher 2015). The tooth preserves seven visible plates, whereas at least one more should be added in the anterior part, where the dentine is confluent. Due to its strong wear, the original plate number, as well as the crown height, cannot be estimated. The lamellar frequency, measured at the base of the crown, results in a low value (Table 1), because of the divergence of the plates towards the base in elephantids (Lister & Sher 2015). The shape of the molar is almost parallel-sided. In labial and lingual views, the tooth shows a slight depression at the middle. Cement covers the regions between the enamel loops as well as the labial, lingual and posterior sides. At the anterior-medial part of the tooth, complete enamel loops are present, which are parallel-sided with anterior and posterior median swellings that are slightly offset from the long axis of the molar (asymmetry). The enamel loops are separated, albeit in some lamellae they are almost in contact, and are oblique relative to the long axis of the tooth. The lateral sides of the enamel figures are rounded. Apical digitations are present on the two distal plates; the last plate, which is at an early wear stage shows four subequal rings. The enamel folding is regular, more intense than APL-716, due to the advanced stage of wear (Van Essen 2011); it is present at the entire length of the enamel figure. Age at death The ontogenetic age is estimated using the dental-wear-based age criteria for the extant African savannah elephant Loxodonta africana provided by Laws (1966). The m3 APL-716 (Fig. 4) falls within Laws��� group XXV, which corresponds to the average age of 49 in African Equivalent Years (AEY). Recent revisions on Laws��� age assignments propose that this age group corresponds to an upper limit of 50 AEY with 70+ longevity for the African elephant (Stansfield 2015; Haynes 2017). Mammuthus meridionalis had a significantly larger body size than extant African elephants, and due to the general positive scaling of longevity to body size across mammals, also probably had a longer total lifespan (Eisenberg 1990; Maiorana 1990); therefore, a slightly older true age is expected for APL-716. When the formula of Blueweiss et al. (1978) (longevity in days proportional to body mass in grams 0.17) is applied, assuming a longevity of 70 years and a body mass of 6.0 tonnes for males of L. africana, and an average body mass of 11.0 tons for M. meridionalis (Larramendi 2016), a longevity of approximately 77.5 years can be hypothesized for the latter. Consequently, the estimated upper limit of the ontogenetic age for APL-716 is 47 �� 77.5/70 = 52 years. Accordingly, the estimated upper limit of the ontogenetic age for the m3 APL- 687 (Laws��� group XXVII; Fig. 3 D-F) is 66-73 years, for the M3 APL-686B (Laws��� group XX; Fig. 3 A-C) is 37 years and for the maxilla APL-225 (Laws��� group II; Fig. 2) is c. 1 year., Published as part of Konidaris, George E., Kostopoulos, Dimitris S. & Koufos, George D., 2020, Mammuthus meridionalis (Nesti, 1825) from Apollonia- 1 (Mygdonia Basin, Northern Greece) and its importance within the Early Pleistocene mammoth evolution in Europe, pp. 69-91 in Geodiversitas 42 (6) on pages 72-74, DOI: 10.5252/geodiversitas2020v42a6, http://zenodo.org/record/3708332, {"references":["AMBROSETTI P., AZZAROLI A., BONADONNA F. P. & FOLLIERI M. 1972. - A scheme of Pleistocene chronology for the Tyrrhenian side of Central Italy. Bollettino della Societa Geologica Italiana 91: 169 - 184.","LISTER A. M., DIRKS W., ASSAF A., CHAZAN M., GOLDBERG P., APPLBAUM Y. H., GREENBAUM N. & HORWITZ L. K. 2013. - New fossil remains of Elephas from the southern Levant: implications for the evolutionary history of the Asian elephant. Palaeogeography, Palaeoclimatology, Palaeoecology 386: 119 - 130. https: // doi. org / 10.1016 / j. palaeo. 2013.05.013","LISTER A. M., DIMITRIJEVIC V., MARKOVIC Z., KNEZEVIC S. & MOL D. 2012. - A skeleton of \" steppe \" mammoth (Mammuthus trogontherii (Pohlig )) from Drmno, near Kostolac, Serbia. Quaternary International 276 - 277: 129 - 144. https: // doi. org / 10.1016 / j. quaint. 2012.03.021","BEDEN M. 1983. - Family Elephantidae, in HARRIS J. M. (ed.), Koobi Fora Research Project. Volume 2: The Fossil Ungulates: Proboscidea, Perissodactyla, and Suidae. Clarendon Press, Oxford: 40 - 129.","LISTER A. M. & SHER A. V. 2015. - Evolution and dispersal of mammoths across the Northern Hemisphere. Science 350: 805 - 809. https: // doi. org / 10.1126 / science. aac 5660","VAN ESSEN H. 2011. - Tracing Transitions. An Overview of the Evolution and Migrations of the Genus Mammuthus Brookes, 1828 (Mammalia, Proboscidea). PhD dissertation, Leiden University, Leiden, 270 p.","LAWS R. M. 1966. - Age criteria for the African elephant, Loxodonta a. africana. East African Wildlife Journal 4: 1 - 37. https: // doi. org / 10.1111 / j. 1365 - 2028.1966. tb 00878. x","STANSFIELD F. J. 2015. - A novel objective method of estimating the age of mandibles from African elephants (Loxodonta africana africana). PLoS ONE 10: e 0124980. https: // doi. org / 10.1371 / journal. pone. 0124980","HAYNES G. 2017. - Finding meaning in mammoth age profiles. Quaternary International 443 (A): 65 - 78. https: // doi. org / 10.1016 / j. quaint. 2016.04.012","EISENBERG J. F. 1990. - The behavioural / ecological significance of body size in the Mammalia, in DAMUTH J. & MACFADDEN B. J. (eds), Body Size in Mammalian Paleobiology. Cambridge University Press, Cambridge: 25 - 37.","MAIORANA V. C. 1990. - Evolutionary strategies and body size in a guild of mammals, in DAMUTH J. & MACFADDEN B. J. (eds), Body Size in Mammalian Paleobiology. Cambridge University Press, Cambridge: 69 - 102.","BLUEWEISS L., FOX H., KUDZMA V., NAKASHIMA D., PETERS R. & SAMS S. 1978. - Relationships between body size and some life history parameters. Oecologia 37: 257 - 272. https: // doi. org / 10.1007 / BF 00344996","LARRAMENDI A. 2016. - Shoulder height, body mass, and shape of proboscideans. Acta Palaeontologica Polonica 61: 537 - 574. https: // doi. org / 10.4202 / app. 00136.2014"]}

Konidaris, George E., Kostopoulos, Dimitris S., Koufos, George D. (2020): Mammuthus meridionalis (Nesti, 1825) from Apollonia- 1 (Mygdonia Basin, Northern Greece) and its importance within the Early Pleistocene mammoth evolution in Europe. Geodiversitas 42 (6): 69-91, DOI: 10.5252/geodiversitas2020v42a6

International audience; We here report on fossil remains of the earliest known crown-Testudo, an extant clade of Mediterranean testudinid tortoises from the late Miocene (Vallesian, MN 10) from the hominoid locality Ravin de la Pluie (RPl) in Greece. The material studied is a small, nearly complete carapace with a clearly distinct hypo-xiphiplastral hinge. This supports the sensu stricto generic assignment. This new terrestrial testudinid specimen is characterized by a possible tectiform, narrow, elongated shell with a pentagonal pygal and a long, posteriorly elevated, lenticular and rounded dorsal epiplastral lip. These unique features differ from those of other known Mediterranean hinged forms and allow the erection of the new species Testudo hellenica sp. nov. This taxon is phylogenetically close to two Greek species, the extant T. marginata and the fossil T. marmorum (Turolian, around 7.3 Ma). This record provides evidence for the first appearance of the genus Testudo sensu stricto at a minimum age of 9 Ma.

Mammuthus meridionalis from Apollonia-1 (Mygdonia Basin, Northern Greece) and its importance within the Early Pleistocene Mammoth Evolution in Europe

We here describe lizards and snakes from the late Miocene (MN 10) of Ravin de la Pluie, near Thessaloniki, Greece, a locality widely known for its hominoid primate Ouranopithecus macedoniensis. The new finds comprise two large-sized lizards (a probable anguine and a varanid) and two snakes (an elapid and a small-sized “colubrine”). Even if the material is represented by few specimens, this is the first record of squamates from the late Miocene MN 10 biozone of southeastern Europe and the third only for the whole continent. The importance of the varanid vertebrae for systematic attributions is discussed. The new varanid limb elements described herein rank among the few such specimens in the fossil record of monitor lizards. Judging from the new and previously published varanid appendicular material, we suggest that Neogene monitor lizards from Europe possessed comparatively short and robustly built limbs. Distinctive scars on one of the limb elements are interpreted as bite marks of a predator or scavenger, offering insights on the palaeoecology of the herpetofauna of the locality.

International audience; The upper incisor lingual morphology of the late Miocene Greek hominoid Ouranopithecus macedoniensis was almost unknown, as the described earlier maxillary remains preserve only worn incisors. During the most recent excavations in the type locality of Ouranopithecus, Ravin de la Pluie (RPl) of Axios Valley (Macedonia, Greece), four little-worn upper central incisors were recovered. This material and a few additional worn upper incisors, discovered recently, are described and compared in this article. Even though a morphological comparison with the old RPl material, lacking unworn or little worn incisors, is impossible, the metrical comparison and the monospecific character of the RPl hominoid sample suggest that the described incisors can be assigned to Ouranopithecus macedoniensis. The described upper central incisors are separated in two size-groups which in general have similar morphology except for some minor differences such as the presence of a pronounced mesial lingual pillar in the small-sized specimens. The observed significant size difference among the studied incisors is attributed to the strong sexual dimorphism of Ouranopithecus, which is also well expressed in the other teeth. The lingual morphology of the upper incisors of Ouranopithecus are not identical to those of extant great apes, though they have some similarities with those of the African great apes (Gorilla and Pan), while they are clearly different from those of the Asian great ape (Pongo). Even though they have some morphological similarities, the O. macedoniensis central incisors are probably not identical to those of the Eurasian Miocene hominoids; the most similar central incisor is that of Ouranopithecus turkae. Among the known African Miocene hominoids, Nakalipithecus upper central incisor is quite similar in morphology and size to that of Ouranopithecus.

Koufos, George D., Sen, Sevket (2016): Equidae. Geodiversitas 38 (2): 225-243, DOI: 10.5252/g2016n2a5, URL: http://dx.doi.org/10.5252/g2016n2a5

Hipparion aff. giganteum Gromova, 1952 LOCALITY. — Küçükçekmece, Turkey. AGE. — Late Vallesian, MN 10; Late Miocene. MATERIAL. — See Table 1. MEASUREMENTS. — See Table 1. DESCRIPTION Upper cheek teeth The upper cheek teeth are large with very rich enamel plication in the fossette's borders and with thin and very deep plis (Fig. 5); the plication number ranges between 14-31 in the premolars and between 23-24 in the molars. The protocone is large, elliptical-oval with flattened lingual wall in the less worn teeth; in MNHN.F.TRQ110 it bears a small mesial spur. The pli caballin is simple to triple. The distal hypoconal groove is well developed and deep while a large lingual hypoconal groove is present in the premolars and a small one in the molars. The P2 TRQ106 has a small and pointed anterostyle, closed fossettes, very rich enamel plication in the fossette's borders (mean plication number = 21), isolated and oval protocone with a small mesial spur, triple pli caballin, deep and narrow distal hypoconal groove, and a shallow and clear lingual hypoconal groove (Fig. 5). The presence of the lingual hypoconal groove, the very rich enamel plication, the multiple pli caballin, and the size of the P2TRQ106 fit well with the Küçükçekmece form-B morphology. Lower cheek teeth The lower premolars are unworn or less worn preventing the description of their occlusal morphology (Fig.5). However, the lower molars (MNHN.F.TRQ46, TRQ111) are more worn and their morphology is clear. The parastylid is well developed and high in TRQ46. The ectoflexid is narrow and deep in the molars with a weak pli caballinid, and the linguaflexid is deep and V-shaped. The pli caballinid is weak and it is reduced by the wear. The metaconid and metastylid are rounded (in the less worn teeth they are elliptical) and the entoconid squarish. The preflexid and postflexid borders are plicated and (or) crenulated. There is not any trace of the ectostylid. Metapodials There is only a fragment of the McIII and an almost complete MtIII (Fig. 6). The slenderness index is 17 for the MtIII and the keel index is 117 for the McIII and 118 for the MtIII, indicating short and robust metapodials with weak keel., Published as part of Koufos, George D. & Sen, Sevket, 2016, Equidae, pp. 225-243 in Geodiversitas 38 (2) on page 232, DOI: 10.5252/g2016n2a5, http://zenodo.org/record/4535533, {"references":["GROMOVA V. 1952. - Le genre Hipparion. Bureau de Recherches geologiques et Minieres CEDP 12: 1 - 288."]}

Koufos, George D., Kostopoulos, Dimitris S. (2013): First report of Brachypotherium Roger, 1904 (Rhinocerotidae, Mammalia) in the Middle Miocene of Greece. Geodiversitas 35 (3): 629-641, DOI: 10.5252/g2013n3a6, URL: http://dx.doi.org/10.5252/g2013n3a6

Genus Brachypotherium Roger, 1904 TYPE SPECIES. — Rhinoceros goldfussi Kaup, 1834., Published as part of Koufos, George D. & Kostopoulos, Dimitris S., 2013, First report of Brachypotherium Roger, 1904 (Rhinocerotidae, Mammalia) in the Middle Miocene of Greece, pp. 629-641 in Geodiversitas 35 (3) on page 634, DOI: 10.5252/g2013n3a6, http://zenodo.org/record/5371747

Mesopithecus pentelicus Wagner, 1839 HOLOTYPE. — Maxillary fragment with M1-M3 described by Wagner (1839) and figured by Wagner (1840). The specimen is housed in BSPM numbered as BSPM ASII. 11. LOCALITIES. — Classical Pikermi ravine (PIK), Attica, Greece (near Athens); Chomateres or Kisdari (CHO), Attica, Greece (near Athens). AGE. — Middle Turolian, MN 12 (late Miocene); more precisely it is referred to the uppermost MN 12 with an age of ~7.0 Ma. Th e fauna of Chomateres is considered slightly older than the Pikermi one. DIAGNOSIS. — Medium-sized colobine monkey; short, upright face; sexual dimorphism in the skull, canines and postcranials; absent or very small sagittal crest in the males; enlarged mandibular angle; shallow mandibular corpus with constant height between p4 and m3; convex anterior symphysis without symphyseal constriction; small and deeply inclined planum alveolare; absent or weak fossa genioglossa; small lingual cusp in the P3, 4; small honing facet in the p3; small hypoconulid in the m3., Published as part of Koufos, George D., 2009, The Neogene cercopithecids (Mammalia, Primates) of Greece, pp. 817-850 in Geodiversitas 31 (4) on page 839, DOI: 10.5252/g2009n4a817, http://zenodo.org/record/5381502, {"references":["WAGNER A. 1839. - Fossile Uberreste von einem Affenschadel und andern Saugethierreste aus Griechenland. Gelehrte Anzeigen Bayerische Akademie der Wissenschaften 38: 301 - 312.","WAGNER A. 1840. - Fossile Uberreste von einem Affenschadel und andern Saugethierreste aus Griechenland. Abhandlungen der Mathematisch-Physikalischen classe der Bayerischen Academie der Wissenschaften 3: 153 - 170."]}

Merceron, Gildas, Koufos, George D., Valentin, Xavier (2009): Feeding habits of the first European colobine, (Mammalia, Primates): evidence from a comparative dental microwear analysis with modern cercopithecids. Geodiversitas 31 (4): 865-878, DOI: 10.5252/g2009n4a865, URL: http://www.bioone.org/doi/abs/10.5252/g2009n4a865

Paradolichopithecus arvernensis (Depéret, 1929) LOCALITY. — Vatera (VTR), Lesvos Island, Greece. AGE. — Middle Villafranchian s.l., MN 17 (late Pliocene). DIAGNOSIS. — Large-sized; clear sexual dimorphism in the canine’s complex; large anterior lower teeth; lingual surface of the lower incisors without enamel; p4 with larger labial than lingual cuspid; m3 with elongated trigonid, low relief and a large sixth cuspid; eruption pattern of lower dentition typically papionin., Published as part of Koufos, George D., 2009, The Neogene cercopithecids (Mammalia, Primates) of Greece, pp. 817-850 in Geodiversitas 31 (4) on page 840, DOI: 10.5252/g2009n4a817, http://zenodo.org/record/5381502

Mesopithecus delsoni Bonis, Bouvrain, Geraads & Koufos, 1990 HOLOTYPE. — Mandible of male adult individual with both tooth rows, RZO-159. It is housed in the Laboratory of Geology and Palaeontology, University of Thessaloniki. LOCALITY. — Ravin des Zouaves-5 (RZO), Axios Valley, Macedonia, Greece. AGE. — Early Turolian, MN 11 (late Miocene); estimated magnetostratigraphic age ~8.2 Ma. DIAGNOSIS. — Large size; deep mandibular corpus; flattened anterior symphysis; strong symphyseal constriction; slightly inclined alveolar plane; large fossa genioglossa; thick inferior transverse torus; large honing facet in the p3; well-developed and bicuspid talonid in the m3., Published as part of Koufos, George D., 2009, The Neogene cercopithecids (Mammalia, Primates) of Greece, pp. 817-850 in Geodiversitas 31 (4) on page 839, DOI: 10.5252/g2009n4a817, http://zenodo.org/record/5381502, {"references":["BONIS L. DE, BOUVRAIN G., GERAADS D. & KOUFOS G. D. 1990. - New remains of Mesopithecus (Primates, Cercopithecidae) from the late Miocene of Macedonia with the description of a new species. Journal of Vertebrate Paleontology 10: 473 - 483."]}

Merceron, Gildas, Blondel, Cécile, Viriot, Laurent, Koufos, George D., Bonis, Louis de (2007): Dental microwear analysis of bovids from the Vallesian (late Miocene) of Axios Valley in Greece: reconstruction of the habitat of Ouranopithecus macedoniensis (Primates, Hominoidea). Geodiversitas 29 (3): 421-433, DOI: http://doi.org/10.5281/zenodo.4665539, {"references":["AGUSTI J., SANZ DE SIRIA A. & GARCES M. 2003. - Explaining the end of the hominoid experiment in Europe. Journal of Human Evolution 45: 145-153.","AXELROD D. I. 1975. - Evolution and biogeography of the Madrean-Tethyan sclerophyll vegetation. Annals of the Missouri Botanical Garden 62: 280-334.","BELL R. H. V. 1971. - A grazing ecosystem in the Serengeti. Scientific American 225: 86-93.","BERNOR R. L. 1984. - A zoogeographic theater and biochronologic play: the time/biofacies phenomena of Eurasian and African miocene mammal provinces. Paleobiologie continentale 14: 121-142.","BONIS L. DE & MELENTIS J. 1977. - Un nouveau genre de Primates hominoide dans le Vallesien (Miocene superieur) de Macedoine.Comptes Rendus de l'Academie des Sciences, Paris 284: 1393-1396.","BONIS L. DE & KOUFOS G. D. 1999. - The Miocene large mammal succession in Greece, in AGUSTI J., ROOK L. & ANDREWS P. (eds), Hominoid Evolution and Climatic Change in Europe: the Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge University Press, Cambridge: 205-237.","BONIS L. DE, BOUVRAIN G., KERAUDAN B. & MELENTIS J. 1973. - Premiers resultats des fouilles recentes en Grece septentrionale (Macedoine). Comptes Rendus de l'Academie des Sciences, Paris 277: 1431-1434.","BONIS L. DE, BOUVRAIN G. & KOUFOS G. D. 1988. - Late Miocene mammal localities of the lower Axios valley (Macedonia, Greece) and their stratigraphic significance. Modern Geology 13: 141-147.","BONIS L. DE, BOUVRAIN G., GERAADS D. & KOUFOS G. D. 1990. - New hominid skull material from the late Miocene of Macedonia in northern Greece. Nature 345: 712-714.","G. D. 1992. - Diversity and paleoecology of Greek","BONIS L. DE, KOUFOS G. D., GUY F., PEIGNE S. & SYL- VESTROU I. 1998. - Nouveaux restes du primate hominoide Ouranopithecus dans les depots du Miocene superieur de Macedoine (Grece). Comptes Rendus de l'Academie des Sciences, Paris 327: 141-146.","BONIS L.DE, BOUVRAIN G. & KOUFOS G. D. 1999. - Palaeoenvironments of late Miocene primate localities in Macedonia, in AGUSTI J., ROOK L. & ANDREWS P. (eds), Hominoid Evolution and Climatic Change in Europe: the Evolution of Neogene Terrestrial Ecosystems in Europe. Cambridge University Press, Cambridge: 413-435.","BOUVRAIN G. 1975. - Un nouveau bovide du Vallesien de Macedoine (Grece). Comptes Rendus de l'Academie des Sciences, Paris, ser. D, 280: 1357-1359.","BOUVRAIN G. 1982. - Revision du genre Prostrepsiceros Major, 1891 (Mammalia, Bovidae). 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Koufos, George D., Vlachou, Theodora D. (2005): Equidae (Mammalia, Perissodactyla) from the late Miocene of Akkaşdağı, Turkey. Geodiversitas 27 (4): 633-705, DOI: http://doi.org/10.5281/zenodo.5376285, {"references":["ARAMBOURG C. F PIVETEAU J. 1929. - Les vertebre s du Pontien de Salonique. Annales de Paleontologie 18: 59-138.","BAKALOV P. F NIKOLOV I. 1962. - The Fossils of Bulgaria. X. Tertiary Mammals. Bulgarian Academy of Science, Sofia, 162 p.","BERNOR R. L. 1985. - Systematic and evolutionary relationships of the hipparionine horses from Maragha, Iran (late Miocene, Turolian age). Palaeovertebrata 15 (4): 173-269.","BERNOR R. L., TOBIEN H. F WOODBURNE M. O. 1990. - Patterns of Old World hipparionine evolutionary diversification and biogeographic extension, in LINDSAY E., FAHLBUSCH V. F MEIN P. (eds), European Neogene Mammal Chronology. Plenum Press, New York: 263-338.","BERNOR R. L., KOUFOS G. D., WOODBURNE M. 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