Elephas Antiquus in Greece: New ﬁnds and a Reappraisal of Older Material (Mammalia, Proboscidea, Elephantidae)

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Quaternary International xxx (2010) 1e11

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Elephas antiquus in Greece: New ﬁnds and a reappraisal of older material (Mammalia, Proboscidea, Elephantidae)

Evangelia Tsoukala a,*, Dick Mol b, Spyridoula Pappa a, Evangelos Vlachos a, Wilrie van Logchem c, Markos Vaxevanopoulos d, Jelle Reumer e a Aristotle University, School of Geology, University campus, 54 124 Thessaloniki, Macedonia, Greece b Natural History Museum Rotterdam and Mammuthus Club International, Gudumholm 41, 2133 HG Hoofddorp, The Netherlands c Mammuthus Club International, Bosuilstraat 12, 4105 WE Culemborg, The Netherlands d Ministry of Culture, Ephorate of Palaeoanthropology-Speleology of Northern Greece, Navarinou 28, 55131, Thessaloniki, Greece e Faculty of Geosciences, Utrecht University and Natural History Museum Rotterdam, PO Box 23452, 3001 KL Rotterdam, The Netherlands article info abstract

Article history: This paper brieﬂy describes some recently discovered remains of the straight-tusked elephant, Elephas Available online xxx antiquus, from Greece. Material of this extinct proboscidean was found in four localities in Northern Greece: Kaloneri and Sotiras in Western Macedonia, Xerias in Eastern Macedonia, and Larissa in Thessaly. In addition, published elephant remains from Ambelia, Petres and Perdikas, also from Northern Greece, are reinterpreted and also attributed to E. antiquus. Of all these, the Kaloneri elephant shows an inter- esting paleopathology: it was disabled by a broken right tusk. Ó 2010 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction of E. namadicus comes from India. Maglio (1973) considered the Asiatic form E. namadicus to be a senior synonym of the European Fossil Proboscidea are known from Neogene and Quaternary form E. antiquus. In that case the taxon E. namadicus/E. antiquus is sediments in Greece, including deposits on the Greek islands. known from Europe and Asia. This paper will be restricted to the Dermitzakis and Theodorou (1980) presented a map of the main European form and applies the commonly used name E. antiquus, proboscidean localities, and Doukas and Athanassiou (2003) pre- pending taxonomic revision of the fossil straight-tusked elephants. sented a revision of the Pliocene and Pleistocene Proboscidea from E. antiquus is very common in the Middle and Late Pleistocene of Greece. During the last few years several new findings came to Europe (e.g., Maglio, 1973) including Greece (Dermitzakis et al., light, and some older ones needed re-interpretation. This paper 1982; Tsoukala, 1992; Doukas and Athanassiou, 2003). Its occur- focuses on the extinct straight-tusked elephant, Elephas antiquus rence has been identified in several fluvial and lacustrine deposits Falconer and Cautley, 1847. in northern Greece. The published material usually consists of E. antiquus is a large elephant characterized by straight tusks. fragmented tusks and molars, while post-cranial skeleton remains Different opinions about its nomenclature are found in the litera- are either rare or unpublished. Therefore, the nearly complete ture. Maglio (1973) attributed the species to the genus Elephas skeleton of the straight-tusked elephant from Perdikas and the Linnaeus, 1758; others (Aguirre, 1968/69) place it in the genus partial skeleton from Ambelia (Tsoukala and Lister, 1998; both sites Palaeoloxodon Matsumoto, 1924. Beden (1983) considered Palae- in Northern Macedonia) are of great interest. oloxodon to be a subgenus of the genus Elephas. As for the taxonomy In Table 1, records are given of the straight-tusked elephants in on the species level, Lister (personal communication, 1992, in Greece, including some references to Mammuthus. Mammuthus Shoshani and Tassy, 1996) considered the large extinct elephant meridionalis is the oldest representative of the Elephantidae in from Europe and Asia to consist of two different species: E. antiquus Greece. It appeared during the Early Pleistocene (MN 17) and it Falconer and Cautley, 1847, and Elephas namadicus Falconer and coexisted for a short time period with Anancus arvernensis (e.g., Cautley,1845, respectively. This division is based on the fact that the Athanassiou and Kostopoulos, 2001). This co-occurrence is found at holotype of E. antiquus originates from Europe, while the holotype Sesklo, although it is not absolutely certain that the remains of the two species originate from exactly the same stratigraphic level. Furthermore, M. meridionalis is also known in Macedonia from e * Corresponding author. Fax: þ30 20 2310 99 8482. Polylakko-Libako-Kapetanios of the broader Grevena Kastoria E-mail address: [email protected] (E. Tsoukala). basin (Steensma,1988), from Sotiras (with Mammuthus trogontherii,

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Table 1 Records of straight-tusked elephants E. antiquus and selected other proboscideans in Greece. The numbers refer to the numbers shown in Fig. 1.

Taxon Locality References A. Continental species (numbers with black dots) 1 E. antiquus M. meridionalis Symvoli (Angitis river) Doukas and Athanassiou, 2003 2 E. antiquus Xerias (Kavala) Present study 3 E. antiquus Epanomi Dermitzakis et al., 1982 4 E. antiquus Trilophos Dermitzakis et al., 1982 5 E. antiquus Allatini Dermitzakis et al., 1982 6 E. antiquus Vathylakkos (Axios) Dermitzakis et al., 1982 7 E. antiquus Amindeo, Sotiras, Petres Velitzelos and Schneider, 1973; Present study 8 (M. meridionalis), E. aff. antiquus, E. antiquus Perdikas Poulianos and Poulianos, 1980; Poulianos, 1984e1986; Present study 9 E. antiquus Tsakoni (Agia Triada, Kastoria) Athanassiou, 2004 10 E. antiquus Kaloneri Present study 11 E. antiquus, M. meridionalis Tsotili Melentis, 1966 12 E. antiquus AmbeliaeGrevena Tsoukala and Lister, 1998 13 E. antiquus, M. meridionalis, M. primigenius Outfall of Stomio (Penios river) Doukas and Athanassiou, 2003 14 E. antiquus Gorge of Kalamaki (Penios river) Athanassiou and Kostopoulos, 2001 15 E. antiquus Larissa, Terpsithea, Nea Lefki Boessneck, 1965; Present study 16 E. antiquus Trichonia (Mesologgi) Athanassiou and Kostopoulos, 2001 17 E. antiquus Patras Athanassiou and Kostopoulos, 2001 18 E. antiquus Lousiká Athanassiou, 2010 19 E. antiquus Nission Doukas and Athanassiou, 2003 20 E. antiquus Roupaki (Ilia) Athanassiou and Kostopoulos, 2001 21 E. antiquus Corinth Channel Athanassiou and Kostopoulos, 2001 22 E. antiquus, M. meridionalis, M. primigenius Megalopolis Melentis, 1961, 1963, 1965; Sickenberg, 1976 23 E. antiquus Kythera island Dermitzakis et al., 1982

B. Insular species derived from E. antiquus (Dermitzakis and Theodorou, 1980; italic numbers in white rectangles) 1e22 Various localities on Crete Dermitzakis et al., 1982; Poulakakis et al., 2002; Doukas and Athanassiou, 2003 23 Rhodos Doukas and Athanassiou, 2003 24 Tilos Doukas and Athanassiou, 2003 25 Dilos Doukas and Athanassiou, 2003 26 Seriphos Doukas and Athanassiou, 2003 27 Milos Doukas and Athanassiou, 2003 28 Naxos Doukas and Athanassiou, 2003 29 Paros Doukas and Athanassiou, 2003 30 Kythnos Doukas and Athanassiou, 2003

Velitzelos and Schneider, 1973), from Philippi (also with M. trogontherii, Psarianos, 1958; Dermitzakis et al., 1982), from Nea Komi (Mitzopoulos, 1967), as well as from the late Early Pleistocene of Apollonia (Koufos and Kostopoulos, 1997) and Kalamotó in the Mygdonia basin (Tsoukala and Chatzopoulou, 2005). At the latest Pliocene and during the Early Pleistocene, M. meridionalis appears to be the only proboscidean. At the early Middle Pleistocene the species is gradually replaced by new, more advanced forms of the same lineage, such as Mammuthus primigenius, which occurs in very few localities in Macedonia (Drama; Koufos, 1981), while more or less at the same time E. antiquus arrived in Europe. To date, more than 20 localities on the Greek mainland and one on Kythera island with straight-tusked elephant remains have been recorded, together with w30 insular localities with remnants of dwarfed elephantids. These endemic taxa derived from E. antiquus after their isolation on several Greek islands (Fig. 1, Table 1). Various researchers (Mol et al., 1996 and references therein) have suggested that the insular proboscideans from Crete include some full-sized elephants, which would perhaps represent founder populations of the mainland E. antiquus. However, Theodorou (1986) refuted these identiﬁcations. Dwarfed elephantids are referred to several endemic species (Symeonidis and Theodorou, 1982; Dermitzakis et al., 1982; Theodorou, 1983; Mol et al., 1996; Poulakakis et al., 2002; Doukas and Athanassiou, 2003). Continental E. antiquus has been divided by some authors into 3 subspecies: the nominal E. antiquus antiquus, E. antiquus italicus Osborn, 1931 and E. antiquus germanicus Stefanescu, 1924. In the past, Greek E. antiquus has sometimes been attributed to both Fig. 1. Map of Greece with the localities of continental Pleistocene elephants. The subspecies E. a. germanicus and E. a. italicus (e.g. from Megalopolis; localities are listed in Table 1. The numbers 7e12 correspond to localities in Northern Melentis, 1961, 1963, 1965; Dermitzakis et al., 1982). Macedonia, and the numbers in squares are referred to the insular faunas.

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2. New material from sites of Macedonia and Thessaly These consist of up to 4 km thick sediments of Middle Eocene to middle Upper Miocene. The sediments of Aliakmon River and its 2.1. Kaloneri elephant tributaries are found above the molasse sequence; they constitute the upper part of the sedimentary sequence. 2.1.1. Introduction and historical overview The fossil remains were found at the south-eastern shore of the The elephant was discovered in the ‘Pera Raches’ area (altitude Mirichos, one of the tributaries of the Aliakmon, in fluvial deposits 581 m a.s.l.), 2 km SW of Kaloneri municipal center of the Munic- of predominating yellow sand with some inclusions of clay and ipality of Askio (Kozani Prefecture), 180 km SSW from Thessaloniki. small gravels. The sand deposits are often calcareous and locally It is situated in a mountainous area of Western Macedonia (Greece), cemented and belong to a Plio-Pleistocene fluvialelacustrine on the foothills of Askio Mountain (Siniatsiko, 2111 m), (x: 284667, sequence (Brunn, 1956). y: 4462000, Greek Grid), (40 170 0.225900 N, 21 280 7.521500 E, WGS84) (Fig. 2). 2.1.3. Material The broader GrevenaeKastoria basin is situated close to the Distal part of the maxillar alveoli (AKL 1) including the left tusk Pindos Mountain Chain and to the river Aliakmon, where many (AKL 2 and AKL 4) and several small fragments of the left tusk, the mammalian fossils have been found (Brunn, 1956; Psarianos, 1958; right tusk (AKL 3); cranial fragments: frontal (AKL 5) and parietal Melentis, 1966; Paraskevaidis, 1977; Steensma, 1988; Tsoukala and (AKL 7a and 7b); right humerus diaphysis fragment AKL 8. Lister, 1998; Tsoukala, 2000; Athanassiou, 2004). The site was discovered in May 2006 during road construction. 2.1.4. Description The works were immediately discontinued and excavations took The remains of the Kaloneri Elephant are well preserved. place in the same month, by a team of the Aiani Archaeological Unfortunately, it became clear at the time of discovery that the skull Museum and the Northern Greece Ephorate of Paleoanthropology e was completely destroyed by a bulldozer. Only the distal end of the Speleology (Ministry of Culture), in co-operation with the School of tusk alveoli and the left and right tusk, including both tips, Geology (Thessaloniki Aristotle University). The material (Figs. 3 and remained (Fig. 5, 6). 4) was in a friable condition, for which reason plaster jackets were The measurements (Fig. 7; Tables 2e4) and especially the used for extracting it. maximum width of the specimen of 602 mm and the distance between the tusk roots of 130 mm, indicate a large, most probably 2.1.2. Geological setting male individual. In general, a large sexual dimorphism exists in The study area is located in the eastern margin of the Meso- proboscideans. The measurements of the alveoli show that the Hellenic trough, which is the largest and most important molassic distance between the tusk-germs of 130 mm is much shorter than trough in Greece. With a length of 130 km and a width of the alveolar diastema of 250 mm. This indicates an extreme 30e40 km, it extends southward from the GreekeAlbanian border divergence of the tusks which is typical for the straight-tusked to the plain of Thessaly, and was formed on the NWeSE oriented elephant, E. antiquus. suture between the Apulian microplate and the Pelagonian conti- The cross-section of both tusks shows that they are of undula- nental block, and filled with molasse sediments (Mountrakis, 1985). tory shape, and in original anatomical position. The frontal and

Fig. 2. Geological map of the Kaloneri area with the excavation site depicted (based on Savoyat et al., 1971)

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Fig. 3. Excavation of the Kaloneri elephant (AKL) in 2006, showing the poorly preserved tusks. lateral sides of the tusks are strongly ﬂattened. After mounting the specimens in their original position, it became clear that the right tusk is pointed downwards and outwards, and slightly curved in frontal position. The left tusk is pointed downwards and strongly outwards, and weakly in a forward direction; the tip is curved towards the axis. The original anatomical positions of the tusks in the alveoli indicate that the carcass of the animal was deposited shortly after death. Crania of extant proboscideans that were exposed on the surface for a long time, are often found without tusks as they become quickly separated due to climatic inﬂuences (especially drought). The undulatory shapes of the cross-section of the tusks and of Fig. 5. Elephas antiquus; The Kaloneri Elephant (Kozani, Western Macedonia); distal the morphology of the tusk sockets of the Kaloneri elephant show part of the maxilla alveoli (AKL 1) including left (I2, AKL 2 and AKL 4) and right tusk that the extreme divergence is authentic. (AKL 3), on display in the Town Hall of Kaloneri. Top: dorsal view, bottom left: right The tip of the left tusk has a narrow and pointed shape, and the lateral view, bottom right: left lateral view. Scale bar is 50 cm (Photo: Hans Wildschut). tooth has its natural length. The right tusk, which is much shorter than the left one, was broken during the lifetime of the animal. It Amindeo (Florina Prefecture, W. Macedonia; Lilibaki-Akamati and was used by the animal after it became damaged, which is shown Trochidis, 1999e2001). The wider region is well known for the by the polished tip giving it a very bold morphology. The remains abundance of fossils found either in sandpits or in lignite deposits, are on display in the Kaloneri new Town Hall. most of them now destroyed (Velitzelos and Schneider, 1973; Doukas and Athanassiou, 2003: Table 1). 2.2. Sotiras elephant 2.2.1. Material Three isolated remains of elephants are exhibited in the An almost complete mandible with m1, m2 sin þ dex (SAF 1; Archaeological Museum of the city of Florina. These specimens Figs. 8 and 9), a proximal fragment of left ulna (SAF 2), a distal were found, years ago, in a sandpit of Sotiras, close to the town of fragment of right femur (SAF 3; Fig. 10).

2.2.2. Description The mandible is relatively short with left and right m1s and m2s, the m1s being in full use, whereas the m2s are partly in use. Part of the ramus with the processus coronoideus and the mandibular condyle is missing. As a result of this, the (empty) alveolus for the left m3 is visible in the ramus behind the m2, thereby assuring the identiﬁcation of the two functioning molars as m1 and m2. The morphology and the measurements (Table 5) unequivocally attribute this specimen to E. antiquus. This conclusion is based on two characteristic features, viz. (a) the intensively folded enamel of the lamellae which is visible on the occlusal surface, and (b) the slightly loxodont conﬁguration (i.e., presence of a loxodont sinus) of the lamellae of left and right m1s (see Fig. 8). These characters are typical for E. antiquus and absent in the genus Mammuthus. The molars indicate an individual age for this specimen of Fig. 4. Excavation of the Kaloneri elephant (AKL) in 2006, showing part of the left tusk. approximately of 22 2 AEY (Laws Age Group XIV) at the time of

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Fig. 7. Parameters measured in the Kaloneri Elephant. The results are in Table 2.

(1973: Figs. 8 and 9), appears to be a fragment of a heavily worn m3. The morphology of the fragment, especially of the roots, makes Fig. 6. Elephas antiquus; The Kaloneri Elephant (Kozani, Western Macedonia); the it clear that this fragmentary m3 belonged to a very old individual. tusks without the alveoli, left tusk (AKL 2 and AKL 4) and right tusk (AKL 3). Scale bar is The occlusal view again shows the characteristic, strongly folded 10 cm (Photo: Hans Wildschut). enamel and the rhombic shape of the lamellae. Hence, this specimen is also attributed to E. antiquus. The third specimen, a fragmented m3 dex, from the same death. Since the m2s are not fully exposed, the total number of gravel-pit was attributed by Velitzelos and Schneider (1973: Figs. 9 lamellae cannot be given (Figs. 8 and 9). and 10) to “A. meridionalis”. Their identification appears a classical The proximal end of a left ulna, damaged at the olecranon and misidentification. Again, this is a heavily worn remnant of an m3 of with a maximum width of 280 mm, indicates a very large (male?) which the occlusal view shows the concave grinding surface. The individual. posterior part of the molar shows that the plates are becoming The third specimen from this locality is the distal end of a right narrower, a typical feature of elephantid last molars. As noted in the femur with the condyles completely preserved. The maximum original description, the lamellae are heavily curved in posterior width measured over the condyles is 290 mm, whereas the direction and worn in an oblique angle, giving the impression that maximum distal width of this femur fragment is 330 mm. Again, the enamel is extremely thick, which would be typical for the this specimen may be attributed to a very large (male?) individual (Fig. 10). Velitzelos and Schneider (1973) described and figured three Table 2 Measurements of the Kaloneri elephant; parameters refer to the numbers in Fig. 7. molars from the same area, one of which is more or less complete. Two of these molars were attributed to “Mammontheus (Elephas) Parameter Description Size in mm trogontherii”, the steppe mammoth, and the other one to “Archi- 1 Maximum width of tusks 2060 diskodon meridionalis”, the southern or ancestral mammoth. The 2 Maximum distance between tips of 2400 first one is a complete m1 sin. This lower molar counts 10 lamellae tusks 3 Transverse distance between tip of 1750 and a talonid, which is typical for an m1 of E. antiquus. The provided right tusk and left tusk figures (Velitzelos and Schneider, 1973: Figs. 6 and 7) show this 4 Perpendicular distance between 1600 molar in lingual and occlusal view. The molar is narrow, a character rostrum and line of parameter 3 for the straight-tusked elephant, and relatively high-crowned. 5 Distance between alveoli (diastema) 250 6 Maximum preserved length of 200 Moreover, the occlusal view of this molar shows many character- left alveolus istics of the cross-section of the lamellae of E. antiquus: rhombic 7 Maximum preserved length of 230 shape of the plates and intensively folded enamel. There are no right alveolus arguments to attribute this molar to M. trogontherii. 8 Width over tusks at place of eruption from alveoli 570 The second specimen, a fragment of an m2 or m3 dex, also 9 Distance between alveoli in the fragmentary rostrum 130 10 Maximum width of rostrum 602 attributed to the steppe mammoth by Velitzelos and Schneider

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Table 3 Measurements in mm of the left tusk of the Kaloneri elephant.

Anterior/posterior diameter 170 Maximum width 160 Maximum length at outer curvature at eruption of alveole 2230 earliest mammoths. However, if the enamel thickness is measured under the right angle, it will become clear that the enamel is considerably thinner. This third specimen is also attributed to E. antiquus.

2.3. Xerias elephant

The fossils from the village of Xerias, close to the Municipality of Chrissoupoli (Kavala Prefecture, Eastern Macedonia), 190 km from Thessaloniki, were found in 2006 during the construction of the Egnatia Highway. The fossils were collected by Mr. G. Adamidis and subsequently donated to the Museum of Geology and Paleontology (Aristotle University, Thessaloniki). They were found at a distance of about 600 m from the village of Xerias, preserved in red clay that is hardly consolidated in some parts.

2.3.1. Material Elephant: Fragmented tusk parts (I2), nrs. XEK 4, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15; an m3 dex (XEK 3); juvenile right femur diaphysis (XEK 5); see Fig. 11. Associated fauna: Megaloceros sp.: skull fragment (XEK 2); Testudo sp.: partial carapace and plastron (XEK 1; Fig. 12).

2.3.2. Description Eleven tusk fragments were found. Every specimen shows some degree of deformation, which is probably due to pressure of the overlying sediment. After detailed study, at least three different individuals were identified, whereas three specimens (XEK 10, 13, Fig. 8. Elephas antiquus; The Sotiras Elephant (Amindeo, Florina, Western Macedonia); almost complete mandible with m1, m2 left and right, SAF 1. Top: occlusal and bottom: 14) were indeterminable. In general, every part belongs to the posterior view. Scale bar is 10 cm (Photo: Evangelia Tsoukala). straight-tusked elephant. XEK 8 and 11 are parts of a right tusk with a small diameter XEK 6 and 12 belong to a left tusk. XEK 6 has a large diameter (Fig. 11). The two specimens share the same morphology and are similar to that of the right tusk of the male individual. XEK 12 is nearly equal in size; hence, it is assumed that they belonged to the a tusk tip and the most deformed specimen. Based on similarities in same individual. XEK 11 is a fragment not far from the tip of the morphology and diameter, XEK 6 and 12 belonged to the same tusk, while on the rear part of XEK 8 the pulp cavity is recognizable. (male?) individual. This shows that the two specimens together represent almost the XEK 3 is the posterior part of an m3 dex of E. antiquus. The full size of the tusk. identification of this molar rest is based on the rhombic (loxodont) XEK 15 is part of a left tusk; it is morphologically close to XEK 8 shape in the cross-sections of the lamellae and on the strongly and 11, but it is significantly larger and thus belonged to another folded enamel (Fig. 11). The concave occlusal surface in this molar individual. These specimens are attributed to two different female rest indicates it being a lower molar. This is confirmed by the individuals, in which the former is more slender in size and of posteriorly directed narrow keel-shaped root. An anterior part of a younger individual. the tooth is missing and is attached to XEK 4, as mentioned above. XEK 7, 4 and 9 are parts of a left tusk with a relatively large Six lamellae are preserved. The molar is extremely worn; all diameter (Table 6) Because of their similar morphology and preserved plates are used, indicating a very old individual, probably diameter, it can be assumed they belonged to the same individual. more than 50 AEY. The thickness of enamel varies from 5.0 to XEK 7 is the longest part and the one closer to the skull. On the 6.4 mm. The maximum width including the cementum is inferior side of the XEK 4 are the remains of a molar. This shows 102.49 mm and the length of the preserved part is 170 mm. that the fossils became mixed after deposition. Because of the large Of the post-cranial skeleton a right femur diaphysis (XEK 5) is size and the robust morphology, these specimens are attributed to preserved. Both epiphyses are missing, but the sutures between an adult male individual. them and the diaphysis are clearly visible in the distal part. This indicates a young individual. The length of the fragment is 640 mm, Table 4 the transversal diameter is 124 mm and the antero-posterior Measurements in mm of the right tusk of the Kaloneri elephant. diameter is 289 mm. Anterior/posterior diameter 165 One artiodactyle remnant (Cervidae Gray, 1821) and the remains Maximum width 155 of a tortoise (Testudinidae Batsch, 1788) were found in association Maximum length at outer curvature at eruption of alveoli 1320 with the elephant remains. Maximum length at the inner curvature at eruption of alveoli 1170 The posterior part of the megaceroid skull (XEK 2) is poorly Circumference at eruption of alveoli 570 preserved. The right occipital condyle is broken, while most of the

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Fig. 10. Elephas antiquus; The Sotiras Elephant (Amindeo, Florina, Western Fig. 9. Elephas antiquus; The Sotiras Elephant (Amindeo, Florina, Western Macedonia); Macedonia); Distal end of a right femur. Top: caudal view and bottom: posterior view almost complete mandible with m1, m2 left and right, SAF 1. Top: left and anterior of the condyles. Scale bar is 10 cm (Photo: Evangelia Tsoukala). view and bottom: right lateral view. Scale bar is 10 cm (Photo: Evangelia Tsoukala). the Testudo marginata group. The maximum preserved length is 260 mm and width is 170 mm, while the estimated total length of basioccipital is well preserved. Of the temporal bone, only the right the turtle is about 330 mm. bulla tympanica is preserved, and both petrosals are large and well distinguished. It is a rather robust skull which can be attributed to 2.4. Larissa elephants Megaloceros sp., and compared with skulls of Megaloceros from Italy, Germany and Great Britain (Caloi, 1972). The measurements of Elephant remains were found in two new sites in the Prefecture of the XEK 2 cranium are: occipital width >190 mm; occipital height Larissa: Terpsithea and Nea Lefki, both veryclose to the city of Larissa, (basioneacrocranion) 122 mm; width of condyles ca. 114 mm; Thessaly. Here, the Penios river is found, in deposits of which many height of foramen magnum (basioneopisthion) 36.5 mm; width of fossils have been uncovered through time (Boessneck,1965, Table 1). foramen magnum 42 mm; width of basioccipital e maximum A partial left mandible with a nearly complete m3 (TEL 1; Fig. 13) under condyles 70.5 mm. is presented in the collection of the Archaeological Museum of The tortoise material consists of a heavily compressed and Larissa. It originates from Penios river sediments, close to the village deformed partial carapace and plastron (XEK 1). A number of plates of Terpsithea, 3 km W of the city of Larissa (personal communica- are preserved, but most of them are incomplete, forming the tion, Mr. G. Toufeksis of the Archaeological Museum of Larissa). The dextral part of a land tortoise. Six neurals, six pleurals and eight specimen consists of the horizontal ramus, the ascending ramus is peripheral bony plates are counted in dorsal view, while the sutures missing, and it is broken at the symphysis. The anterior part of the between the horny plates are very deep. The medially alternative long and short pleurals, versus laterally short and long, which coincide with alternative quadrate and octagonal neurals is typical Table 5 Measurements in mm of the Sotiras elephant mandible with m1 and m2. for a testudinid shell (Lapparent de Broin et al., 2006). In ventral view, parts of the right hyoplastron, hypoplastron and xiphiplas- Maximum width of the mandible 590 tron are poorly preserved (Fig. 12). In the right hyoplastron the Maximum height of the mandible 190 Maximum length anterior/posterior of the mandible 410 pectoro-abdominal sulcus can be observed, while the posterior part Maximum width of m1 sin 82 is marked by the presence of the hypo-xiphiplastral hinge. The Maximum width of m2 sin 83 presence of this hinge indicates that XEK 1 is a Testudo s.s. species Maximum width of m1 dex 86 (Lapparent de Broin et al., 2006). Further identiﬁcation is, however, Maximum width of m2 dex 85 Maximum length of grinding surface of m1/m2 sin 202 impossible as both characteristic posterior and anterior parts are Maximum length of grinding surface of m1/m2 dex 218 missing. However, in the right posterior part the shell widens, as in

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Table 6 Measurements in mm on tusk parts of the Xerias elephant. White lines separate the different individuals.

XEK no. Gender Max. length Diameter antero-posterior (preserved) (preserved) 8 \ 325 96 11 \ 263 82 15 \ 431 103 4 _ 209 158 9 _ 155 137 7 _ 402 163 6 _ 347 144 12 _ 205 89

From the site of Nea Lefki, which is 13 km south of Larissa, 120 m of altitude, there are two large tusk fragments (NLL 1 and 2) and a group of over 10 smaller fragments (NLL 3). They were found in coarse conglomerates by Mr. K. Floras. The morphology of the fragment, the straight shape without curvature and its large size, all indicate an old and large male individual of E. antiquus.

3. Older material reinterpreted

3.1. Ambelia skeleton

The Ambelia skeleton, a large male, adds signiﬁcantly to the limited knowledge of the range of variation of this species as a whole (Figs. 14 and 15). The ESR dating of lamellae from Ambelia

Fig. 11. Elephas antiquus; The Xerias Elephant; Top: fragment of a right tusk of a female individual (XEK 8). Bottom: posterior fragment of an extremely worn right m3 (XEK 3; top: lingual view, middle: occlusal, bottom: buccal view). Scale bar is 10 cm (Photo: Evangelia Tsoukala). molar is damaged. The ﬁrst plate is only half preserved and from here 14 plates are counted up to and including the talon (N15). Up to 12 plates are in function, which indicates an adult individual. The morphology of the cross-section of the enamel shows all the characteristics of E. antiquus: the rhombic shape of the cross-sections of the plates and the strongly folded enamel.

Fig. 12. Testudo sp.; Drawing of the deformed shell of a tortoise from Xerias locality (XEK 1). Left: dorsal view of carapace and right: ventral view of the plastron. White Fig. 13. Elephas antiquus; The Larissa Elephant (TEL 1); Top: occlusal view of a left areas are bony plates, stripped areas eroded bony plates, dark areas are missing, lines mandible with incomplete m3 and bottom: lingual view. Scale bar is 10 cm (Photo: indicating sutures between plates. Scale bar is 5 cm (Drawing: Evangelos Vlachos). George Toufeksis).

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Fig. 14. Elephas antiquus: The AmbeliaeGrevena straight-tusked elephant, thoracal vertebra (GRE 141), caudal (left) and lateral (right) view. Scale bar is 10 cm (Photo: Hans Wildschut). indicate the presence of E. antiquus in Greece during Oxygen Isotope Stage 6 (OIS 6; Tsoukala and Lister, 1998). In northern and central Europe, E. antiquus is generally regarded as an indicator of interglacial climate, being present during temperate, wooded episodes corresponding to odd-numbered MIS stages (Stuart, 1982). It is supposed to have become extinct at the end of the Eemian interglacial. Mol et al. (2007) have discussed 14C dates showing that there is a strong possibility that this species became extinct in northwestern Europe much later than generally accepted, during the Weichselian glacial. During intervening cold episodes, the species was generally absent from these regions during the Late Pleistocene. For the majority of these long, cold episodes, the range of E. antiquus is presumed to have contracted to the south. The dating of the Ambelia elephant remains to MIS 6 corroborates this suggestion, indicating Northern Greece as part of the glacial refu- gium of the species (Tsoukala and Lister, 1998). Fig. 15. Elephas antiquus: The AmbeliaeGrevena straight-tusked elephant; the right The Ambelia skeleton has been described in detail by Tsoukala anterior foot in anatomical connection. Top: dorsal view, middle: axial view and and Lister (1998). These authors have shown that this skeleton bottom: lateral view. Scale bar is 10 cm (Photo: Hans Wildschut). has all characteristics of the straight-tusked elephants. However, some characteristics that have not been mentioned in the past might be useful to distinguish skeletal remains of straight-tusked Macedonia. From one of these, a locality near Petres, they described elephants from other European Pleistocene proboscideans. Firstly, an M2 dex. and an M3 dex., found in a badly preserved maxilla with the length of the corpus vertebra in all vertebrae is much longer a damaged right tusk. The tusk was in a very bad state of preser- antero-posteriorly and the vertebrae are more heavily built than in vation and heavily fragmented, and was preserved. Pictures were the genus Mammuthus. The dorsal spines of the vertebral column taken of the specimen in situ and included as Fig. 2 in their paper fl are much longer antero-posteriorly and extremely attened (Velitzelos and Schneider, 1973). The remains were attributed to the (Fig. 14). This character is absent in other proboscideans. This straight-tusked elephant, indicated by them as Palaeoloxodon fl morphology of the vertebrae will have an extreme in uence on the (Elephas) antiquus and dated to the Early to Middle Pleistocene. body shape of the straight-tusked elephant. Velitzelos and Schneider (1973) provided measurements and Another feature noticed by analyzing the more or less complete descriptions of the molars, as well clear black and white images right front foot (Fig. 15) is that the Ambelia straight-tusked (their Figs. 2e5). It is not clear from their paper where these fossils fi elephant was thumbless. The rst metacarpal (Mc I) is well from Petres are stored. Study of these descriptions and images led preserved and the distal epiphysis is not showing an articular to different results concerning the nature of the molars. surface for the attachment of the first phalanx, which means that there were only four digits in this specimen. This character, among (1) Fig. 2 in Velitzelos and Schneider (1973) shows that the partial others, is often seen in M. primigenius (Mol et al., 2004, 2006). cranium fragment, still in situ, possessed a heavily fragmented right (straight) tusk, an M1 and an M2 dex, in anatomical order. 3.2. Petres material (2) Fig. 2 in Velitzelos and Schneider (1973) shows the composed fragmented maxilla with M1 dex., M2 dex. and the M2 sin. Both Velitzelos and Schneider (1973) described and figured five M2s show, in lateral view, the s-shaped morphology of the Pleistocene elephantid molars from two localities in Western lamellae in the blunt posterior part.

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(3) Fig. 4 in Velitzelos and Schneider (1973) shows the relatively 4. Conclusion high-crowned left M2 seen from the lingual side, and Fig. 5 does so in occlusal view. The morphology of the molar makes The straight-tusked elephant E. antiquus is widespread in it clear that this is an M2. However, Velitzelos and Schneider central and northern Greece. New material described here from (1973) identified this molar as an M3. Kaloneri, Sotiras, Xerias, Terpsithea and Nea Lefki testifies to this (4) Velitzelos and Schneider (1973) provided measurements, importance. Two nearly complete skeletons found in Ambelia and lamellar formulae, pictures and descriptions of the dental Perdikas add to this knowledge. The easternmost distribution of E. elements, that make clear that these should be identified as M1 antiquus in continental Greece is Xerias in Kavala, E. Macedonia, dex and M2 dex (complete, counting 10 lamellae and a talon while a specimen is also known from Lousiká on the Peloponnese and positioned in anatomical order), and the M2 sin. peninsula (Athanassiou, 2010). The original anatomical positions of the tusks in the alveoli of These remains belong to E. antiquus. The stage of wear of the the Kaloneri elephant indicate that the carcass of the animal has three molars corresponds with the Laws (1966) age groups been deposited shortly after death. The specimen indicates an XIVeXV, which is more or less equivalent to 20e26 African extreme divergence of the tusks which is typical for E. antiquus and Elephant Years (AEY) at the time of death of the animal. helps understand the behaviour of the animal. The measurements and descriptions provided here show clearly a large male individual fi 3.3. Perdikas elephant of old age. The large diameters of 170/165 mm underline identi - cation of the specimen as a male individual; in female specimens of The elephant of Perdikas is a straight-tusked elephant, E. anti- E. antiquus such sizes have never been recorded. The broken right quus. The nearly complete skeleton was discovered in 1978 by Mr. tusk and the slender and pointed shape of the left tip indicate that Isaac Passalidis and was excavated thereafter. The remains of this the Kaloneri Elephant used its tusks intensively. The cause of the skeleton are still in anatomical order and preserved in situ in a small breakage is unknown and can only be speculated upon. building that was erected on the site in order to protect the pale- Except for the Ambelia E. antiquus, there are no data available fi ontological heritage. The skeletal material itself is unfortunately about the stratigraphical position ( rst and last occurrences) of this poorly preserved (Fig. 16). Poulianos and Poulianos (1980) were the species in the Pleistocene of Greece. Based on the MIS 6 for the e first to study the Perdikas elephant material. They attributed it to Ambelia Grevena specimen it is clear that E. antiquus was present “A. meridionalis” and subsequently (Poulianos, 1984e1986)toEle- in Greece around the end of the late Middle Pleistocene. It is phas aff. antiquus. The loxodont form in the cross-sections of the assumed, looking at the few faunal elements that were discovered lamellae, the strongly folded enamel, and the narrow crown leave in association with E. antiquus, that the species is restricted to the no doubt about the identity of this specimen as E. antiquus Middle and/or Late Pleistocene. As an example, the associated fauna (Tsoukala and Lister, 1998). A Pliocene age for the Perdikas spec- of Ptolemaida consists of Hippopotamus antiquus and Bos imen, as suggested by Poulianos and Poulianos (1980), therefore primigenius. fi seems unlikely, because E. antiquus arose from the African Elephas The material described in this paper con rms that E. antiquus is recki (Maglio, 1973), and first entered Europe in the early Middle a heavily built and large representative of the Elephantidae in the Pleistocene (Lister, 1996). In addition, the stratigraphical and Pleistocene of Greece. The fact that E. antiquus is now known from geological setting of the region indicates a Middle to Late Pleisto- so many localities, should stimulate further research in order to cene age (Pavlides, 1985). help understand the paleobiology, biostratigraphy and paleo- Based on the morphology of the left and right molars (m3) biogeography of this animal. present in the mandible, it is clear that the skeleton belongs to the straight-tusked elephant. The sizes and the morphology of the Acknowledgements bones indicate that the skeleton belonged to an old male individual of approximately 50 AEY. A rough estimate of its shoulder height is The authors would like to thank the team members who between 380 and 410 cm. This skeleton is unique, because of the participated in the time-pressured excavations of the Kaloneri rarity of complete findings in anatomical order. It requires detailed elephant in May 2006: V. Makridis, C. Tsagouli, M. Miteletsis, study, preservation and protection. C. Lokana, L. Zdoupas, I. Nenos and G. Natas. For taking part in the 2008 restoration and the display in the Kaloneri Town Hall: V. Makridis, G. Lazarides, O. Koukousioura, M. Spyridaki, H. Wild- schut, K. Tsekoura and D. Labretsa. We are indebted to the mayors of Herakleotes, Mr. A. Karagiannis, and Askion, Mr. V. Patras, for their interest. H. Wildschut provided most of the photographs used in this paper. Two anonymous reviewers provided helpful sugges- tions for improving the text.

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