Palaeoloxodon Antiquus) and Other Large Mammals from the Middle Pleistocene Butchering Locality Marathousa 1 (Megalopolis Basin, Greece): Preliminary Results

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The skeleton of a straight-tusked elephant (Palaeoloxodon antiquus) and other large mammals from the Middle Pleistocene butchering locality Marathousa 1 (Megalopolis Basin, Greece): preliminary results

* George E. Konidaris a, , Athanassios Athanassiou b, Vangelis Tourloukis a, Nicholas Thompson a, Domenico Giusti a, Eleni Panagopoulou b, Katerina Harvati a a Eberhard Karls University of Tübingen, Palaeoanthropology, Senckenberg Centre for Human Evolution and Palaeoenvironment, Rümelinstr. 23, 72070 Tübingen, Germany b Ministry of Culture, Ephorate of Palaeoanthropology-Speleology, Ardittou 34B, 11636 Athens, Greece article info abstract

Article history: In this article, we present the first results on the large mammal fauna from the new open-air Lower Received 12 July 2017 Palaeolithic locality Marathousa 1 (MAR-1) (Megalopolis Basin, Peloponnesus, Greece). MAR-1 belongs to Received in revised form the Marathousa Member of the Choremi Formation and its large mammal faunal list (collection 2013 14 November 2017 e2016) includes the castorid Castor fiber, the mustelids Mustela sp. and Lutra simplicidens, the felid Felis Accepted 1 December 2017 sp., the canids Vulpes sp. and Canis sp., the elephantid Palaeoloxodon antiquus, the hippopotamid Available online xxx Hippopotamus antiquus, the bovid Bison sp., and the cervids Dama sp. and Cervus elaphus. This faunal association is common in the Galerian (Middle Pleistocene) mammal communities of Europe (ca. 0.9e0.4 Keywords: Fauna Ma). The MAR-1 fauna is consistent with a temperate climate and is indicative of a landscape with Biochronology substantial woodland components with more open areas, close to permanent and large freshwater Palaeoecology bodies. Of particular interest are an elephant cranium and numerous postcranial elements, which were Taphonomy found in close anatomical association and are attributed to a single individual of the straight-tusked Cut marks elephant Palaeoloxodon antiquus. The skeleton belonged to a male individual in its late adulthood Palaeolithic close to or in its sixties, with live skeletal height around 3.7 m at the shoulder and body mass around 9.0 tonnes. The good state of preservation of the MAR-1 bones allows the identification of taphonomic modifications. Cut marks on the elephant skeleton, and on other elephant and mammal bones, indicate human exploitation by means of butchering activities, in accordance with the traits of the lithic assemblage and its spatial association with the bones. Carnivore activity is also recorded on some elephant and cervid bones. Marathousa 1 is among the oldest elephant butchering sites in Europe and the only one known in Southeastern Europe. © 2017 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

1. Introduction lacustrine sediments (Fig. 1a), and is characterized by the presence of economically important lignite seams (Vinken, 1965). Since 1969 The Megalopolis area is an intramontane basin situated in the lignite deposits are mined extensively in open-air quarries and western Arcadia, in the centre of the Peloponnesus (Fig. 1a). used for electricity production. The exposed sections during the Geologically it is a graben, formed in the MesozoicePalaeogene mining operations have offered a unique opportunity to study the basement of the Peloponnesus, and extending along a NWeSE di- stratigraphy and the environment of the Pleistocene palaeolake, rection. The basin is ﬁlled with Neogene and Pleistocene ﬂuvial and which once covered the basin. The lacustrine layers preserve most notably remains of vertebrates and plants, mainly mammalian fossils, wood and fruits.

Abbreviations: DAP, antero-posterior diameter; D, depth; dp, deciduous lower The mammal fossils of Megalopolis are known since antiquity, premolar; DT, transverse diameter; H, height; L, length; m/M, lower/upper molar; and ‘giant’ bones attributed to mythical beings are reported in p, lower premolar; W, width. ancient Greek writings about the region (see Melentis, 1961; Darlas, * Corresponding author. 2003). The ﬁrst systematic palaeontological excavations in the E-mail address: [email protected] (G.E. Konidaris). https://doi.org/10.1016/j.quaint.2017.12.001 1040-6182/© 2017 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Fig. 1. a, Geographic position of the Megalopolis Basin and the Marathousa 1 site (lignite mines expansion taken from www.openstreetmap.org in 2015), and stratigraphic column of the basin [modified from van Vugt (2000) and Panagopoulou et al. (2015)]. The globe was taken from commons.wikimedia.org and the map of Greece from www.shadedrelief. com; b, simplified stratigraphic columns of Area A and Area B showing the stratigraphic units and correlation between the two areas [modified from Karkanas et al. (this issue)]; absolute elevation in m.a.s.l. (meters above sea level). The elephant skeleton from Area A lies at the contact of UA3/UA4 and is covered by UA3; c, distribution map of the elephant remains in Area A, grey zones indicate the 2013e2016 excavated areas.

Megalopolis Basin took place in several localities in 1902 by Pro- Tourloukis et al., this issue a), micro- and macro-fauna (fishes, fessor T. Skouphos (University of Athens) (Bürchner, 1903; amphibians, reptiles, birds, mammals; see also Doukas et al., this Skouphos, 1905; Melentis, 1961), but unfortunately, as with issue and Michailidis et al., this issue), as well as micro- and several historical collections, the fossils lack precise stratigraphic macro-flora (Field et al., this issue). Area A represents mainly a information. Therefore, this collection, which was studied in detail dense accumulation of elephant bones, belonging to a single indi- by Melentis during the 1960s (Melentis, 1961, 1963a, b, 1965a, b, c, vidual of the straight-tusked elephant Palaeoloxodon antiquus, d, e, 1966), does not represent a single stratified fossil accumulation associated with lithic artefacts and other faunal remains. Area B, and cannot be used for biochronology. Since then, further palae- which lies ca. 60 m to the south of Area A, is generally characterized ontological work in the basin has been conducted, including the by the presence of relatively abundant lithic artefacts in association collection of Pleistocene fossils and a human molar from the Mar- with faunal remains. Lithological and stratigraphic correlations athousa Beds by a team of German geologists in the 1960s between the excavated sequences (Fig. 1b; Karkanas et al., this (Sickenberg, 1975); the study on the magneto- and cyclo- issue), along with spatial patterns (Giusti et al., this issue), stratigraphy of the basin, as well as the micro- and macro- archaeological (Panagopoulou et al., this issue; Tourloukis et al., this vertebrate fauna of the Choremiou section (Marathousa Member), issue a) and palaeontological/zooarchaeological evidence (this conducted by van Vugt and colleagues in the 1990s (van Vugt, study) altogether indicate that the find-bearing units in the two 2000); the palaeontological excavations by the University of Ath- excavation areas are correlated and belong to the same depositional ens (Theodorou, 2014) and the study by Athanassiou and colleagues event. Consequently, notwithstanding the fact that the archaeo- in the Kyparissia mine (Athanassiou et al., this issue). The Pleisto- logical signature of the two excavation Areas may represent cene fossil fauna from Megalopolis is therefore relatively well different episodes and/or human activity areas (Panagopoulou known. However, most of the collections mentioned above do not et al., this issue; Tourloukis et al., this issue a), the palae- represent a single stratified assemblage, which can be used for ontological material is treated here as a single faunal assemblage. secure biochronological and palaeoecological interpretations; The preliminary faunal lists for Area A and Area B are presented in furthermore, none of this material was found in association with Table 1. More information on the locality and the stratigraphy can archaeological remains and therefore it cannot provide information be found in Panagopoulou et al. (this issue), Karkanas et al. (this about early human activities. Here we report on the first such issue) and Tourloukis et al. (this issue b). stratified assemblage associated with palaeolithic cultural remains, The studied material includes all the large mammal specimens recovered from a controlled excavation context at the new open-air collected from Marathousa 1 during the field seasons 2013e2016. Lower Palaeolithic site Marathousa 1. The aim of this article is to All specimens are stored at the Ephorate of Palaeoanthropology- provide preliminary results about the large mammal fauna and the Speleology in Athens (Greece). Measurements were taken with biochronology of Marathousa 1, as well as remarks on palae- digital calipers or, in the case of very large specimens, with a oecology and taphonomy. measuring tape. Cut mark analysis: We inspected every bone with a stereomi- e 2. Locality, materials and methods croscope (Leica MZ95, 10 eyepiece, 0.63 6 zoom objective lens) or in the case of very large specimens with hand lenses under Marathousa 1 was discovered during systematic survey by a strong direct light. Specimens with marks were molded using high- joint team of the Ephorate of Palaeoanthropology-Speleology of the resolution silicone (polyvinyl siloxane; Regular Body Microsystem, Greek Ministry of Culture and the Palaeoanthropology department Coltene President). Each cast was subsequently poured with epoxy of the University of Tübingen in the framework of the ERC project resin (Epo-tek 301 Parts A and B) in order to produce replicas. Data m PaGE ePalaeoanthropology at the Gates of Europe: human evolu- were acquired with the use of a Sensofar Pl Neox confocal imaging fi tion in the southern Balkanse (Harvati and Tourloukis, 2013). The pro ler with 10 objective lens with a vertical resolution of < m survey was conducted in 2012 and 2013, with the goal of locating 50 nm, a lateral sampling interval of 1.66 m, and an aperture of fi sites with archaeological/palaeoanthropological interest (see 0.30. Three dimensional representations and pro les of the stria- Thompson et al., this issue). The new locality Marathousa 1 (MAR- tions were obtained with the SensoMAP 7.0 software using the fi 1) was discovered in 2013, when stratified bones and lithic artefacts default settings without application of lters, apart from removing fi fi were identified in a profile of the Marathousa Member (Choremi outliers, while all data were rst leveled. Pro les between 30% and Formation) (Panagopoulou et al., 2015). The find-bearing layers 70% of the marks' length were studied for the cross-sectional occur between two lignite seams (Fig. 1b) and are composed mainly analysis (Mate Gonzalez et al., 2015). of silty sands. Two excavation areas were defined: Area A and Area B, both yielding lithics (see Panagopoulou et al., this issue and

Table 1 Faunal list of the large mammals from Marathousa 1 (MAR-1). Specimens from the section between Areas and B originate from the upper part of the sedimentary sequence, below UA2 and UB2 (see Fig. 1b).

Order Family Genus Species Area A Area B Section between Areas A and B

Rodentia Castoridae Castor ﬁber UA2, UA3, UA3/UA4 Carnivora Mustelidae Mustela sp. (large-sized) þ Carnivora Mustelidae Lutra simplicidens UA3 Carnivora Felidae Felis sp. UB4 Carnivora Canidae Vulpes sp. UB4 Carnivora Canidae Canis sp. UB4 Proboscidea Elephantidae Palaeoloxodon antiquus UA3/UA4 UB4, UB5 þ Artiodactyla Hippopotamidae Hippopotamus antiquus UA2, UA3 UB4 Artiodactyla Bovidae Bison sp. UB4 Artiodactyla Cervidae Dama sp. UB4, UB5 Artiodactyla Cervidae Cervus elaphus UA3, UA3/UA4 UB4, UB5 þ

3. Palaeontology by Laws (1966). That author used the lower teeth, but a similar wear pattern can be assumed for the upper teeth. The anterior parts 3.1. The Marathousa 1 elephant skeleton of the M3s are not well preserved so the exact number of plates is difficult to reconstruct (at least pending final preparation of the The MAR-1 excavation is in progress, but important skeletal cranium). Nevertheless, it appears that the teeth likely fall within elements of the elephant have been already unearthed in Area A, Laws' group XXVII, which corresponds to the average age of 53 in including the cranium (MAR-1-942/676-6; pending final prepara- African Equivalent Years (AEY). Recent revisions on Laws' age as- tion), which bears the pair of upper third molars and the proximal signments propose that this age group corresponds to an upper part of the right tusk (Fig. 2a). The morphology of the cranium with limit of 60e66 AEY with 70þ longevity for the African elephant widely laterally divergent (flaring) premaxillary tusk alveoli, (Stansfield, 2015; Haynes, 2017). Palaeoloxodon antiquus had a accompanied with dental features such as the relatively narrow significantly larger body size than extant African elephants, and crown, the intensively folded enamel of the lamellae and the due to the general positive scaling of longevity to body size across presence of pointed midline sinuses (Saegusa and Gilbert, 2008; mammals, also probably had a longer total lifespan (Eisenberg, Lister et al., 2012), support the attribution to Palaeoloxodon. The 1990; Maiorana, 1990); therefore, a slightly older true age is ex- combination of the above features excludes an attribution to pected for the MAR-1 elephant. When the formula of Blueweiss Mammuthus, the other candidate elephantid genus of the European et al. (1978) (longevity in days proportional to body mass in Pleistocene [which is characterized by tusk alveoli that are sub- grams0.17) is applied, assuming a longevity of 70 years and a body parallel and situated close to one another, relatively wide molars mass of 6.0 tonnes for males of L. africana, and a body mass of 9.0 with weakly folded enamel and absence of pointed midline si- tonnes for the MAR-1 elephant, a longevity of approximately 75 nuses; Lister and Stuart (2010), Lister et al. (2012)], and are shared years can be hypothesized for the latter. Consequently, the esti- with the European straight-tusked elephant Palaeoloxodon anti- mated upper limit of the ontogenetic age for the MAR-1 elephant is quus. The final preparation and study of the cranium will likely 60 (to 66) 75/70 ¼ 64e71 years. Therefore, it is reasonable to reveal additional relevant morphological features. assume that the MAR-1 elephant was in its late adulthood (but not The proximate spatial accumulation of several elephant bones, senile) close to or in its sixties. This age is in agreement with the the lack of duplication, and the consistency in size and ontogenetic extent of epiphyseal fusion of the skeletal elements. Elephants have age indicate the presence of a single elephant individual (Fig. 1c). a prolonged period of growth and the fusion of their long bones There is no anatomical connection between the bones; despite this, extends into their middle age or even further, and males complete however, the skeleton is not largely dissociated so that several their epiphyseal fusion sequence well after females (Haynes, 1991, originally articulated bones occur in approximate anatomical as- 2017). Considering that the MAR-1 elephant is male (see below), sociation (Fig. 1c). To date, numerous skeletal elements have been and following the fusion sequence in recent male elephants and recovered in addition to the cranium, such as vertebrae (including mammoths provided by Roth (1984), Lister (1999) and Haynes atlas, axis and sacrum), ribs, the left humerus (pending prepara- (2017), the last epiphyses to fuse are the proximal femur, and tion), the right ulna, the pelvis, the right femur, the left tibia and then the distal radius and ulna. Assuming similar fusion series for several carpals, tarsals, metapodials and phalanges, for a total of 77 the MAR-1 elephant, the not completely fused distal ulna and the teeth and bones recorded so far (Fig. 3). fused proximal femur (Fig. 2d, f), support the advanced age for the The available skeletal elements of the MAR-1 elephant provide MAR-1 elephant. metric data for the approximate estimation of its shoulder height Elephants, both recent and extinct, are characterized by pro- and body mass. For the estimation of shoulder height from isolated nounced sexual dimorphism, particularly with regard to body size limb bones, we used the regression equations of Lister and Stuart emales being considerably larger than females (Haynes, 1991; (2010), which have been developed on the basis of mounted skel- Sukumar, 2003). Several observations on the MAR-1 elephant etons of mammoths, and we obtained a live shoulder height indicate that it was male: a) Based on the Neumark-Nord (Ger- (including flesh) of approximately 3.7 m for the MAR-1 elephant many) P. antiquus (Palombo et al., 2010, fig. 22), female individuals (Table 2). The ulna is considered problematic for the estimation of are not expected to significantly surpass a shoulder height of 3.0 m; the shoulder height (Lister and Stuart, 2010) and therefore is not the estimated shoulder height of 3.7 m for the MAR-1 elephant included. The body mass of the elephant can be estimated using the exceeds this limit considerably. b) The dimensions of the post- prediction equations of Christiansen (2004), which are based on cranial elements help to distinguish the sex of extinct elephantids samples of recent Loxodonta africana and Elephas maximus. Using based on the larger size observed in males (e.g., Kroll, 1991; the best predictors proposed by Christiansen for the available MAR- Averianov, 1996; Tsoukala and Lister, 1998; Palombo and Villa, 1 bones (Table 3), body mass estimates range significantly from 2003). The large size of several bones, such as the atlas (W max: ~6.7 to 11.0 tonnes with an average of 8.8 tonnes. Among the 491 mm), axis (H max: 333 mm), ulna (L max: 1035 mm), femur (L possible measurements, ulna length and ulna least circumference max: 1330 mm), tibia (L max: 792 mm) and calcaneus (D max: of the diaphysis are the most reliable estimators, given their lowest 274 mm), fit within the ranges of corresponding bones belonging to standard error of the estimation (SEE) and prediction error (PE) presumed male individuals of P. antiquus (Kroll, 1991). c) The percentages. These provide a mass of ~9.0 tonnes for the MAR-1 developed dorsal and ventral tubercles and the marked muscle elephant. Another method for the body mass estimation uses scars on the cranial articular surfaces of the atlas, which contrast prediction equations of body mass against shoulder height for with the presumed female atlas from Crumstadt, Germany (Kroll, recent elephants (Roth, 1990 and references cited therein). Given 1991, p. 14), as well as the robust odontoid process (dens) of the the estimated live shoulder height of 3.7 m for the MAR-1 elephant, axis (Fig. 2b and c), point to a male (Averianov, 1996; Marano and an average body mass of ~9.1 tonnes is obtained. The results of the Palombo, 2013). d) The morphology and the ratios between mea- best osteological predictor (ulna) and of the mass estimate based on surements of the pelvic girdle are essential to the sex determina- the shoulder height are in very good agreement, and therefore a tion of proboscideans (Lister, 1996a; Palombo and Villa, 2003). The body mass of 9.0 tonnes is proposed for the MAR-1 elephant MAR-1 pelvis is still under preparation and the two innominates (Table 3). were found separated almost at the pubic symphysis. Therefore, the Age at death can be estimated by the dental-wear-based age ratio between the pelvic aperture width and the ilium shaft width, criteria for the extant African elephant Loxodonta africana provided a major criterion for sex determination, is not available at the

Fig. 2. aef, Palaeoloxodon antiquus remains from the Marathousa 1 (Area A) skeleton; a, cranium in situ (MAR-1-942/676-6) and detail of the left and right upper third molars; b, atlas, MAR-1-940/674-39, cranial view; c, axis, MAR-1-940/675-41, lateral view; d, right ulna, MAR-1-940/677-21, medial view; e, right half of the pelvic girdle, MAR-1-938/674-37, caudal view; f, right femur, MAR-1-940/675-50, cranial view; g, ratio between pelvic aperture height and ilium shaft width (measurements are shown in ‘e’); circles: males, squares: females; data from Lister (1996a), Gohlich€ (2000) and Lister et al. (2012).

Fig. 3. Schematic drawing of a Palaeoloxodon antiquus skeleton (by K. Schauer/C. Beauval available at www.archeozoo.org) showing the identiﬁed anatomical parts (in parentheses the side or the number of specimens; R: right, L: left) belonging to the Marathousa 1 (Area A) skeleton (collection 2013e2016).

Table 2 Estimation of the shoulder height of the Marathousa 1 (Area A) elephant from femur and tibia lengths, based on linear regression equations (SH ¼ aX þ b; SH: shoulder height, X: the bone length in mm, a, b: slope and intercept of regression equation) given by Lister and Stuart (2010).

Element/measure MAR-1 measure (mm) a b R-square MAR-1 skeletal SH estimate ± 95% MAR-1 live SH (mm) conﬁdence/prediction interval (mm) þ6% for ﬂesh

femur length ('best') 1330 3.0438 549.104 0.8901 3449 ± 112/349 3656 tibia length ('best') 792 3.8455 462.384 0.7219 3508 ± 225/593 3718 Average SH 3687

Table 3 Estimation of the body mass of the Marathousa 1 (Area A) elephant based on predictive equations of modern elephants, a, from bone lengths (Christiansen, 2004); SEE: standard error of the estimation and PE: prediction error percentages, and b, from shoulder height (Roth, 1990 and references cited therein); BM: body mass, SH: shoulder height.

a. Element Equation MAR-1 measure (mm) a b % SEE % PE Estimated ВМ (kg)

ulna length log (BM) ¼ a þ b (logX) 1035 4.135 2.674 8.41 5.34 8452 ulna least circumference where X is the bone 430 1.349 2.022 5.78 4.42 9460 femur length variable (mm) 1330 5.568 3.036 14.54 6.15 8242 tibia length 792 3.064 2.378 12.47 6.93 6748 tibia least circumference 360 2.724 2.647 10.72 6.57 11,029 Average (all bones) 8786 Average (ulna) 8956

b. Source population Equation (SH in cm) MAR-1 SH (cm) Estimated ВМ (kg)

L. africana (males) BM ¼ 5.07 10 4 SH2.803 370 8011 L. africana (males) BM ¼ 3.06 10 4 SH2.890 8088 L. africana (males) BM ¼ 1.81 10 4 SH2.97 7678 E. maximus (males and females) BM ¼ 4.68 10 5SH3.263 11,228 E. maximus (males, captive) log (BM) ¼3.436 þ 2.906 log (SH) 10,646 Average 9130 Average (both methods) 9043

Please cite this article in press as: Konidaris, G.E., et al., The skeleton of a straight-tusked elephant (Palaeoloxodon antiquus) and other large mammals from the Middle Pleistocene butchering locality Marathousa 1 (Megalopolis Basin, Greece): preliminary results, Quaternary International (2017), https://doi.org/10.1016/j.quaint.2017.12.001 G.E. Konidaris et al. / Quaternary International xxx (2017) 1e20 7 moment. However, the ratio between pelvic aperture height and and are almost exact to PEC 1600 of V. praeglacialis from Petralona ilium shaft width, an alternative sex indicator (Lister, 1996a) can be (Greece; L W: 14.2 5.7 mm; Baryshnikov and Tsoukala, 2010). calculated for the MAR-1 specimen. The value for this ratio (2.19) They are smaller than V. vulpes from L'Escale (L: 15.0e16.9, W: places the MAR-1 elephant among the male individuals of 5.0e6.2 mm; Bonifay, 1971), as well as from Agios Georgios Cave P. antiquus, as well as those of Mammuthus (Fig. 2e, g), again sup- (Greece; L W: 15.5 6.2 mm; Tsoukala, 1992), but within the porting male sex. e) As was mentioned above, the fused proximal range of variation of Late Pleistocene and recent red foxes (e.g., femur and the not completely fused distal ulna, combined with the Ballesio, 1979; Petronio et al., 2006). Although an attribution to advanced ontogenetic age, is consistent with a male sex. V. praeglacialis is possible, taking into account the large size variability in foxes (e.g., Kurten, 1965; Davis, 1977) we attribute the 3.2. Other large mammals from Marathousa 1 MAR-1 fox to Vulpes sp. A carnivore axis (MAR-1-933/594-77) and a fifth metatarsal The presence of a beaver in the MAR-1 fauna is recognized by (MAR-1-932/596-79) resemble in morphology those of canids and incisors, cheek teeth and postcranial elements. The lower incisors correspond to the size of a wolf. The reduced ventral keel and the with straight anterior margin, smooth enamel on the outer side and caudally bent articular facets for the atlas, which are only slightly triangular cross-section, the high-crowned cheek teeth with long dorsoventrally extended (Fig. 4j), are different from Cuon priscus and deep striiae/striids, and the humerus with a wide caput humeri from Hundsheim and resemble Canis (Thenius, 1954). Its di- attached to the tuberculum majus (Fig. 4g, Table 4), are some mensions [LCDe: 50.6, LAPa: 50.2, BFcr: 30.9, H: 39.0; according to morphological features distinct from Trogontherium cuvieri, and von den Driesch (1976); articular facets for atlas: 17.2 11.5; all in similar to Castor fiber (Guenther, 1965; Kretzoi, 1969; Fischer, 1991; mm] are similar to Canis mosbachensis from Untermaßfeld (Ger- Fostowicz-Frelik, 2008). many) and C. lupus lunellensis from Lunel-Viel (France) (Sotnikova, The carnivores are represented by two mustelids, two canids 2001; Boudadi-Maligne, 2010). The morphology and the di- and one felid. The first mustelid is known by a mandibular frag- mensions of the metatarsal also match those of Canis (Fig. 4i, ment, showing typical Lutra-like appearance (MAR-1-939/675-35; Table 4). The metatarsal is much smaller than that of C. lupus lupus Fig. 4k; Willemsen, 1992; Sotnikova and Titov, 2009). Features on (L: 85.7e90.9, DT diaphysis: 7.0e8.2, DT distal: 11.4e12.9; all in the m1, such as the trigonid that is broader than the talonid mm), and is more similar in size to that of C. l. lunellensis (L: (Table 5), the shallow talonid basin, the large metaconid, and the 70.6e77.2, DT diaphysis: 5.5e8.9, DT distal: 8.8e10.1; all in mm); absent hypoconulid, all distinguish the MAR-1 otter from L. lutra however, it fits best with the C. mosbachensis specimens from and resemble L. simplicidens (Willemsen, 1992). The studied spec- L'Escale (France; L: 66.7e75.7, DT diaphysis: 4.7e5.5, DT distal: imen's m1 is very similar in its dimensions to known lower molars 8.4e9.9; all in mm) (Boudadi-Maligne, 2010). Although an attri- of L. simplicidens from various European localities (Willemsen, bution to the latter species seems plausible, we attribute both 1992; Sotnikova and Titov, 2009; Cherin and Rook, 2014). specimens to Canis sp., because of a certain degree of overlap be- Sotnikova and Titov (2009) erected the new subspecies L. s. tam- tween C. mosbachensis and C. l. lunellensis, and especially due to the anensis, based on the morphology of a lower carnassial from lack of more diagnostic specimens. Chumbur Kosa (Sea of Azov, Russia), whose talonid width is equal An isolated upper carnassial (MAR-1-933/596-19) belongs to a to that of the trigonid, the metaconid is slightly enlarged, the small felid. The protocone is broken; the relatively strong parastyle talonid blade bears a complete number of well-developed cusps is slightly labially situated and a lower but distinct ectoparastyle is and the distal occlusal outline is relatively pointed. Apart from the located at the mesiolabial side. The paracone is prominent and the latter feature, the simplified talonid morphology and the overall metastyle becomes rounded at its distal end (Fig. 4aec). The di- proportions of the MAR-1 m1 are distinct from L. s. tamanensis, and mensions of the tooth [L W: (11.0) (5.4) mm] are clearly smaller similar to L. s. simplicidens from Hundsheim (Austria, holotype), than those reported for Lynx (15.9e19.1 6.6e9.1 mm) and they Voigtstedt, Mosbach-2 (Germany), East Runton and West Runton fall comfortably within the range of variation of the extant Euro- (England) (Thenius, 1965; Willemsen, 1992). If this subspecific pean wildcat Felis silvestris silvestris (9.6e12.1 4.5e6.7 mm), close distinction is corroborated in the future by additional samples, then to the Felis specimens from West Runton (11.3 6.1 mm) and Lunel the MAR-1 mandibular specimen could be referred to the nominal Viel (12.2 6.1 mm) (Bonifay, 1971; Ballesio, 1980; Lewis et al., subspecies. 2010; Ghezzo et al., 2015; Boscaini et al., 2016). Candidate Felis The other mustelid is represented by a tibia (MAR-1-939/637-2; species for the Pleistocene of Europe are F. lunensis and F. silvestris. Fig. 4h). Its size (Table 4) is smaller than recent Meles and Martes, Upper dentition that could be referred with certainty to the former but larger than Mustela erminea and M. nivalis, and falls within the species is so far unknown, while additionally its taxonomy is in size variation of M. putorius, M. eversmanni and M. lutreola question, mainly due to the scarcity of findings, with some re- (Archaeozoology collection-University of Tübingen; Galik, 1997). searchers considering F. lunensis as a subspecies of F. silvestris. In the Because taxonomic identification based on a single tibia is very absence of additional relevant comparative material, we attribute difficult (Ziegler, 1996; Galik, 1997), the range of variation of the MAR-1 wildcat to Felis sp. Pleistocene specimens is unknown and cranial or dental remains In addition to the Palaeoloxodon antiquus skeleton in Area A, have not been recovered to date for this taxon from the site, we isolated elephant bones were excavated in Area B, as well as refer to this mustelid as Mustela sp. (large-sized). recovered from the section between the two areas. These include A lower carnassial (MAR-1-932/595-69) belongs to a small several tusk, vertebra, rib and tibia fragments. The lack of diag- canid, and presents morphology and dimensions similar to Vulpes. nostic specimens does not permit a taxonomical attribution; The m1 is characterized by a rather strong and distinct metaconid, however, given that they originate from the same geological layer which is higher than the entoconid and lower than the paraconid; as the skeleton, they are tentatively attributed also to P. antiquus. bicuspid talonid with stronger hypoconid; a weak entoconulid; The artiodactyls are represented by a hippopotamid, a bovid and and, a low hypoconulid (Fig. 4def). The dimensions of the tooth two cervids. The presence of the hippopotamid is documented by a (L W: 14.2 5.8 mm) are larger than V. alopecoides, e.g., from dp2 (MAR-1-939/676-6) and a third metacarpal (MAR-1-1). The Dafnero-1 (Greece; L: 12.8e13.2, W: 5.3e5.6 mm; Koufos and dp2 is long and narrow [L: 27.5, W: (11.2), H: 15.1; all in mm]. It is Kostopoulos, 1997), fall at the upper limit of V. praeglacialis, e.g., formed by one conical main cusp, whose distal wall bears a from L'Escale (France; L: 13.2e14.3, W: 4.9e5.7 mm; Bonifay, 1971), conjunct low cusplet; distolabially there is an additional, relatively

Fig. 4. aec, Felis sp., right P4, MAR-1-933/596-19; a, labial; b, lingual; c, occlusal view; def, Vulpes sp., left m1, MAR-1-932/595-69; d, labial; e, lingual; f, occlusal; g, Castor fiber, left humerus, MAR-1-941/676-54; caudal view; h, Mustela sp., right tibia, MAR-1-939/637-2, dorsal view; i, Canis sp., left fifth metatarsal, MAR-1-932/596-79, dorsal view; j, Canis sp., axis, MAR-1-933/594-77, lateral view; k, Lutra simplicidens, left mandibular fragment with p3em1, MAR-1-939/675-35, lingual view; l, Cervus elaphus, right third and fourth metacarpal, MAR-1-939/637-1, dorsal view; men, Dama sp., right mandibular fragment with p2em3 showing carnivore gnawing, MAR-1-934/594-71 and MAR-1-932/598-51 (refitted specimens); m, labial; n, lingual view; o, Palaeoloxodon antiquus, spinous process of a vertebra showing carnivore gnawing, MAR-1-935/602-83, lateral view.

Table 4 Postcranial measurements (in mm) of the large mammals from Marathousa 1.

Species Castor ﬁber Mustela sp. Canis sp. Hippopotamus antiquus Bison sp. Dama sp. Cervus elaphus

Specimen humerus tibia MtV McIII PhII (hindlimb) radius radius McIIIþIV

Inventory number MAR-1 941/676-54 939/637-2 932/596-79 1 934/609-23 933/601-6 933/596-82 939/637-1

Length 85.1 55.5 68.4 172.2 50.8 e 281.8 270.2 DT proximal 24.5 10.3 11.9 61.7 35.9 e 55.8 45.4 DAP proximal 20.4 8.6 8.7 60.4 41.2 e 30.2 31.1 DT diaphysis 9.5 3.5 5.2 49.5 27.4 e 34.5 26.6 DAP diaphysis 10.6 4.0 6.8 30.0 28.4 e 19.7 25.0 DT distal 30.3 8.0 8.6 60.0 28.7 37.8 47.4 45.9 DAP distal 10.7 6.3 9.8 47.5 35.9 26.8 36.0 30.7

Table 5 Dental measurements (in mm) of the large mammals from Marathousa 1. Measurements in parentheses represent the greatest measurable value of the parameter. Superscripts indicate in which lobe of the tooth the greatest width was measured.

Species Lutra simplicidens Felis sp. Vulpes sp. Hippopotamus antiquus Dama sp.

Inventory number MAR-1 939/675-35 933/596-19 932/595-69 939/676-6 934/594-71, 932/598-51

L premolars 39.0 L molars 62.9 L dp2 27.5 W dp2 (11.2) Lp2 10.9 Wp2 7.5 L p3 6.1 13.3 W p3 3.7 8.1 L p4 7.9 13.5 W p4 4.3 9.6 (2) L m1 13.1 14.2 16.9 L m1 trigonid 7.9 10.1 W m1 7.0 5.8 11.9 (2) W m1 trigonid 6.3 5.3 W m1 talonid 6.0 5.7 Lm2 18.3 Wm2 12.7 (2) Lm3 27.2 Wm3 12.7 (1) L P4 (11.0) W P4 (5.4)

strong and more isolated cusplet (Fig. 5aec). The dp2s of the two The high values of the DAP/DT proximal (1.15) and DAP/DT distal species commonly occurring in Middle Pleistocene European lo- (1.25) indices of MAR-1-934/609-23 are different from those of calities, Hippopotamus antiquus and H. amphibius, show morpho- B. primigenius and B. priscus, being closer to B. schoetensacki (Fig. 6f logical variability. MAR-1-939/676-6 is morphologically similar to and g). Although an allocation to the latter species could be that of H. amphibius figured by Hooijer (1942, pl. 10) and to some considered, a single phalanx does not allow for a secure specific specimens of H. antiquus from Untermaßfeld (Kahlke, 1997). The attribution. We therefore prefer to refer the MAR-1 large bovid to comparative material is rather limited, but MAR-1-939/676-6 is Bison sp. for the time being. longer than those of H. amphibius [L: 21.0e22.0 mm (n ¼ 2); Faure The cervids belong to two size-groups. A small- to medium- (1985)] and falls at the upper limit of H. antiquus from Unter- sized cervid is recognized by a mandibular fragment bearing the maßfeld [L: 20.5e27.5 mm (n ¼ 4); Kahlke, 1997)]. The third p2em3 (MAR-1-934/594-71 refits with MAR-1-932/598/51; metacarpal is more robust than those of H. amphibius and fits Fig. 4men, Table 5). Morphological features of the mandible and comfortably within the range of variation of H. antiquus (Fig. 5def, the teeth, such as the small labial foramen below the distal edge Table 4). The taxonomy of the Middle Pleistocene hippopotamuses of the p2, the shorter mandibular diastema than the molar tooth from Europe is under debate, but following the two species/groups row, the separated paraconid and parastylid on the p3, the stronger concept of Petronio (1995), Palombo and Valli (2003, appendix) and ectostylid on the m1 relative to the m2, the antero-posteriorly Pandolfi and Petronio (2015), both specimens can be referred to the separated protoconid and hypoconid on the molars, and the over- large-sized H. antiquus. lap of the anterior entoconid and the posterior metaconid wings on A large-sized bovid is represented by an intermediate phalanx of the m3, are common Dama characters (Lister, 1996b; Breda and the hindlimb (MAR-1-934/609-23) presenting Bison-like features Lister, 2013; Breda, 2015). The MAR-1 teeth present some (Fig. 6aed, Table 4; Sala, 1986; Balkwill and Cumbaa, 1992). Its morphological similarities with the type series of the newly erected length differentiates it from the longer phalanges of Bison priscus species D. roberti from Pakefield (England) and Soleilhac (France) and the much longer ones of Bison menneri; it falls within the range [p3: paraconid and parastylid are not well-separated; p4: meta- of variation of Bos primigenius and Bison schoetensacki (Fig. 6e). conid and entoconid form a closed lingual wall (“molarised”); m3: According to Sala (1986), the hindlimb intermediate phalanges absence of an additional stylid between hypoconid and talonid, lack among the candidate bovid species can be distinguished by the of a clear step between the lingual walls of second and third lobes] proportions of the proximal epiphysis and by the narrow distal end. (Breda and Lister, 2013). However, they also present some features

Fig. 5. aec, Hippopotamus antiquus, left dp2, MAR-1-939/676-6; a, labial; b, lingual; c, occlusal view; dee, Hippopotamus antiquus, left third metacarpal, MAR-1-1; d, dorsal; e, proximal view; f, logarithmic ratio diagram comparing the third metacarpal measurements of Hippopotamus from Marathousa 1 (MAR-1-939/676-6) to those of H. amphibius and H. antiquus [data from Faure (1985), Mazza (1995), Kahlke (1997), Galobart et al. (2003), and Tsoukala and Chatzopoulou (2005)]. Standard of comparison: H. amphibius, recent (n ¼ 9e11) (Faure, 1985). Measurements: 1, L; 2, DT proximal; 3, DAP proximal; 4, DT diaphysis; 5, DAP diaphysis; 6, DT distal; 7, DAP distal. which are absent from that species but common in D. dama, molars (L: 19.1e20.9, W max: 20.3e21.8; all in mm) with moderate including D. d. clactoniana [p4: presence of an anterior hypoconid to strong entostyle and lingual cingulum, and no fold inside the wing; molars: not very strongly developed ectostylids]. The sig- anterior wing of the protocone (Lister, 1996b) are also attributed to nificant inter- and intraspecific dental morphological variability this species. among Dama species/subspecies, and especially the absence of antlers in MAR-1, do not allow for a definite specific determination. 4. Biochronology Therefore, we attribute the MAR-1 fallow deer to Dama sp. Addi- tionally, a distal radius fragment (MAR-1-933/601-6; Fig. 9f) pre- Palaeoloxodont elephants migrated from Africa to Eurasia at the sents morphological features similar to those of Dama (Lister, end of the Early Pleistocene (~1.0 Ma) via the Levantine corridor 1996b) and its dimensions (Table 4) distinguish it from both the (Saegusa and Gilbert, 2008). Their earliest occurrence in Europe is smaller Capreolus and the larger Cervus. documented with the species Palaeoloxodon antiquus at Slivia (Italy, The second cervid is of medium to large size and its presence Slivia Faunal Unit), dated after the Jaramillo submagnetochron and was confirmed by some dental and postcranial specimens, before the Brunhes/Matuyama boundary, ~0.9e0.8 Ma (Palombo, including a radius (MAR-1-933/596-82), a metacarpal (MAR-1-939/ 2014). During the Middle and Late Pleistocene, the species was 637-1) and a calcaneus (MAR-1-939/635-1). The metacarpal is not widespread in Central and Southern Europe (Palombo et al., 2010). well preserved (Fig. 4l); however, the clear space between the Its last appearances are post-Eemian, dated at ~35 ka (Sousa and facets in the volar part of the proximal epiphysis, the rather strong Figueiredo, 2001; Mol et al., 2007; Athanassiou, 2011). The tuberosity in dorsal view, and the visible split in the distal epiphysis appearance of Cervus elaphus in Europe roughly coincides with the in dorsal and volar views, point to Cervus elaphus (Lister, 1996b). Its arrival of the palaeoloxodonts. The earliest records of the true red dimensions (Table 4) distinguish it from the smaller Dama,aswell deer C. elaphus with the subspecies C. e. acoronatus are reported just as from the giant deer genera Praemegaceros and Megaloceros. below the Brunhes/Matuyama boundary (end of Epivillafranchian, Although crania or antlers are absent to date from MAR-1, the di- ~0.9e0.8 Ma) in Dorn-Dürkheim 3 (Germany), Gran Dolina TD6 and mensions of the metacarpal match with those of the corresponding TDW4 (Atapuerca, Spain), Slivia (Italy) and possibly in Saint-Prest specimens of the red deer C. elaphus from Voigtstedt, Mosbach, (France, ~1.0e0.9 Ma) (Palombo, 2017 and references cited Neumark-Nord (Germany), Boxgrove and Petralona Cave (Greece) therein). With regard to Lutra simplicidens, its oldest records are [apart from the length which is smaller than Voigtsted (294.4 mm, known from Upper Valdarno (Italy), correlated to the beginning of n ¼ 1) and Boxgrove (308.0 mm, n ¼ 2), and larger than Petralona the late Villafranchian (Olivola and Tasso Faunal Units), and from (255.4e262.5 mm, n ¼ 2); however, the number of specimens is Chumbur Kosa (Russia, 1.1e0.8 Ma), correlated to the Epivilla- low] (Kahlke, 1965; Tsoukala, 1989; Di Stefano and Petronio, 1992; franchian (Cherin and Rook, 2014; Sotnikova and Titov, 2009). The Pfeiffer, 1999; Lister et al., 2010). Therefore, an attribution to this more advanced morphological features of the MAR-1 L. simplicidens species is plausible. The morphology and the dimensions of the indicate a younger age. The last appearances of L. simplicidens are radius (Table 4) and the calcaneus (L: 117.6, DT: 35.9, H: 46.7; all in reported from Hundsheim and Mosbach-2, correlated to the MIS 15 mm) match also with those of C. elaphus. Three isolated upper or MIS 13 (~0.6e0.5 Ma), while the modern L. lutra is reported from

Fig. 6. aed, Bison sp., left intermediate phalanx of the hindlimb, MAR-1-934/609-23; a, dorsal; b, adaxial; c, abaxial; d, proximal view; eef, boxplots with selected ratios comparing the intermediate phalanx from Marathousa 1 (MAR-1-934/609-23) with the intermediate phalanges of the hindlimb of Bos primigenius, Bison menneri, Bison schoetensacki and Bison priscus [data from Sala (1986) and Sher (1997)].

Hoxne, England, at MIS 11 (~0.4 Ma) (Stuart et al., 1993; Kahlke Pleistocene age (Doukas et al., this issue). Preliminary post-infrared et al., 2011) [note however, that the Hoxne lutrine is regarded as Infrared Stimulated Luminescence (post-IR IRSL) indicate an age Lutra sp. by Willemsen (1992), thus for the moment the timing of between ~0.5 and 0.4 Ma (Jacobs et al., this issue). The Electron Spin the replacement of the older species by the younger remains un- Resonance (ESR) of a mollusk sample from UA2 provided a mini- certain]. The oldest appearances of Hippopotamus in Europe are mum age at 473 ± 54 ka, while five subsamples of a cervid tooth observed in the middle Villafranchian localities Elis (Greece) and from UB4c gave an age of 502 ± 13 ka (Blackwell et al., this issue). Coste San Giacomo (Italy) (Thenius, 1955; Reimann and Strauch, Finally, the study of the magnetostratigraphy suggests that the 2008; Bellucci et al., 2012). Hippopotamus antiquus (and closely find-bearing stratigraphic units (UB4ceUB5 and UA3ceUA4) have related forms classified as H. tiberinus or H. ex gr. H. antiquus)is an age broadly comprised between ~0.48 Ma and ~0.42 Ma recorded in several localities of the late Villafranchian and until the (Tourloukis et al., this issue b). Therefore, the large mammal Middle Pleistocene (Palombo, 2014; appendix). The last appear- assemblage is consistent and further supports all of these pre- ances of this species and its replacement by H. amphibius are quite liminary age estimates for MAR-1. vague. In Italy H. antiquus is present at ~0.6 Ma at Isernia la Pineta, Maglianella and Via Portuense (Isernia Faunal Unit, MIS 15), while 5. Palaeoecological remarks the first appearances of H. amphibius might be traced back to MIS 13 e or MIS 11, 0.53 0.45 Ma during the Fontana Ranuccio Faunal Unit Palaeoloxodon antiquus had a wide and flexible ecological fi (Palombo, 2014; Pandol and Petronio, 2015). However, according adaptation, as the species inhabited mild humid, warm to warm- Н to Mazza and Bertini (2013) H. ex gr. . antiquus survived until temperate and moderately wooded to wooded environments, but e ~0.13 0.11 Ma. also wooded or even rather arid grasslands (Palombo et al., 2010). Overall, the MAR-1 large mammal faunal association is common In Northern and Central Europe, the species occurred during the in the Galerian (beginning of Galerian sensu Bellucci et al., 2015) interglacial phases and, with few exceptions, was generally absent mammal communities of Europe and the current biochronological from the intervening cold stages of open habitats. During these data indicate that the locality is dated between 0.9 Ma and 0.4 Ma. times, its range was limited to Southern Europe, which acted as a e The study of the micromammals suggests a middle late Middle refugium (Tsoukala and Lister, 1998; Lister, 2004). Isotopic and

Please cite this article in press as: Konidaris, G.E., et al., The skeleton of a straight-tusked elephant (Palaeoloxodon antiquus) and other large mammals from the Middle Pleistocene butchering locality Marathousa 1 (Megalopolis Basin, Greece): preliminary results, Quaternary International (2017), https://doi.org/10.1016/j.quaint.2017.12.001 12 G.E. Konidaris et al. / Quaternary International xxx (2017) 1e20 dental microwear analyses on P. antiquus teeth from La Polledrara substantial woodland components and more open areas. Taking di Cecanibbio (MIS 9), Casal de' Pazzi (MIS 7), Neumark-Nord (MIS into account the good representation of semiaquatic taxa (Castor, 7) and Megalopolis Basin [Middle Pleistocene; specimens from Lutra, Hippopotamus), the locality was close to the shore of large several sites within the basin described by Melentis (1961)], and permanent freshwater body. This is also supported by the showed that the species was a mixed feeder, but with a significant presence of a rich avian fauna mostly adapted to a lake environ- amount of grasses in its diet (Palombo et al., 2005; Grube et al., ment (Michailidis et al., this issue), the water vole Arvicola (Doukas 2010; Rivals et al., 2012). et al., this issue) and the dominantly aquatic and waterside vege- In the MAR-1 fauna, three semiaquatic mammals are present: tation (Field et al., this issue). This lakeshore setting was ideal for Hippopotamus, Castor and Lutra. Morphological and palae- the subsistence of several animals. Herbivores (cervids, bovids, el- oecological studies on Hippopotamus antiquus indicate a rather ephants) would visit this region in order to drink water and feed on aquatic habitat with high dependence on water, as the species the nearby vegetation, while the small carnivores (Mustela, Lutra, foraged mainly on aquatic vegetation, in contrast to the grazing Felis, Vulpes) would prey on small mammals, frogs, birds, fishes and (terrestrial) dietary habits of the extant H. amphibius (Mazza, 1995; invertebrates (all are recorded in the MAR-1 faunal assemblage). Palmqvist et al., 2003; Madurell-Malapeira, 2012). Hippopotamuses Such a location would attract also larger carnivores, such as wolves, are generally considered indicators of temperate climate; however, which could find a wide range of prey for their survival. This it is the presence and amount of water (linked to precipitation), landscape was probably also ideal for early humans, offering the rather than temperature, that are of vital importance to this taxon protection and the plant resources of the nearby woodlands, as well (Mazza and Bertini, 2013). The similar dentition of the otter Lutra as accessible freshwater and a variety of feeding opportunities simplicidens with the recent Eurasian otter L. lutra indicates a probably throughout the year, including ungulate and megafauna comparable diet consisting dominantly of fish (Willemsen, 1992); carcasses, close to the shores of the lake. however, other food sources like frogs, crayfish, crabs, birds, turtles or small mammals would complement its diet (Lanszki et al., 2006; 6. Taphonomical remarks MacDonald, 2009). Although otters forage mainly in water, they spend much of their time resting on land (MacDonald, 2009). For the taphonomic investigation of the faunal remains we Willemsen (1992) suggested that the postcranial skeleton of studied the specimens originating from the upper part of the L. simplicidens shows stronger aquatic adaptations than L. lutra. The sedimentary sequence below the Lignite Seam III of Area A and Area Eurasian beaver Castor fiber is a riparian and semiaquatic species B (see Karkanas et al., this issue; Fig. 1b). We treated each of these as inhabiting ponds, lakes, rivers and streams. Beavers are generalist one assemblage in order to increase the sample, i.e., one assem- herbivores with a diet ranging from relatively non-woody plants blage includes UA2, UA3 and the specimens found at the contact of (e.g., leaves, roots, herbs, ferns, grasses, algae) to shrubs and trees UA3/UA4 from Area A, and UB3, UB4, UB5 from Area B. However, it (e.g., aspen, birch, maple, oak, dogwood, fruit trees) (MacDonald, should be noted that the majority of the fauna (as well as of the 2009). lithics) from Area A and Area B originate from UA3c and UB4c, Cervus elaphus is mainly found today at the edges of woodland, respectively (Fig. 1b). where it can browse the foliage of trees, and also consume grasses The large mammal assemblage examined from Area A includes and low herbaceous vegetation (Lister, 1984). However, the species 136 bones or bone fragments and 28 teeth or tooth fragments. The presents a high ecological flexibility, as it is encountered from taxa representation and composition is presented in Table 6 and woodlands to open or even treeless environments, feeding on Fig. 7a. As expected, the majority of the identifiable specimens shrubs and tree shoots in the former, and grasses, sedges and belong to Palaeoloxodon antiquus (skeleton) or to ?Palaeoloxodon rushes in the latter (Lister, 1984; Parfitt, 1999). During the Middle antiquus (79.0%), while Castor fiber and cervids represent 9.2% and Pleistocene, the species was present during both interglacial and 6.7% respectively of the assemblage. Because the elephant is the cold stages (Lister, 1984). On the other hand, the presence of Dama main faunal element in Area A and is of primary interest, its full sp. in the MAR-1 assemblage provides more information. The taphonomic study will be conducted separately, after all the fallow deer in Northern and Central Europe is generally limited to elephant bones of the skeleton are excavated. interglacial stages, as it is less tolerant to open and cold conditions The analyzed sample from Area B includes a total of 211 bone/ than the red deer and needs temperate climate (Lister, 1984). It bone fragments and 35 teeth/tooth fragments. Palaeoloxodon anti- mostly inhabits open deciduous or mixed woodland with a well- quus is the main faunal component also in Area B, representing developed shrub layer, feeding in forest clearings and woodland 27.3% of the identifiable specimens, followed by fallow deer (25.0%) margins (Parfitt, 1999). Although a specific attribution of the MAR-1 and red deer (9.1%). The combined cervids are the dominant group bison is not secure at the moment, the Middle Pleistocene European (56.8%) of the Area B sample (Table 6, Fig. 7a). Bones from Area B are Bison species are generally considered as open landscape dwellers, characterized by a high degree of fragmentation: a) Their maximal having a significant grass component in their diet (Palombo, 2016; diameter ranges from 14.4 mm to 390 mm (mean value: 59.7 mm, appendix). standard deviation: 53.1 mm) with the great majority measuring The recent European wildcat is associated with open forest <80.0 mm (Fig. 7b). This size range is much smaller than the ex- habitats. Small cats are opportunistic and highly effective hunters, pected length of complete long bones (except phalanges) belonging killing their own prey (small mammals, amphibians, reptiles, birds, to medium-sized herbivores like cervids (apart from a complete fish), although occasionally they also eat carrion (MacDonald, Cervus radius all other values over 200 mm correspond to elephant 2009). Middle Pleistocene European Vulpes and Canis species are bone fragments). b) Bone fragments make up 93.4% of the assem- generally ecologically flexible taxa, able to live in shrubland or blage, whereas the percentage of complete/almost complete bones woodland, as well as in an open landscape, or at the edge of both; is only 6.6%. The above observations clearly demonstrate the highly foxes are considered as flesh-eaters consuming mainly small fragmented nature of the bone assemblage and account for the low mammals, while wolves are bone crushers capable of grinding percentage of identifiable bones. Of the 211 bones, only 46 (21.8%) bones, but also quite efficient in consuming flesh (Palombo, 2016; could be attributed to a specific skeletal element, while the rest appendix). were unidentifiable (78.2%, mostly splinters or shaft fragments). Overall, the large mammal fauna from MAR-1 is consistent with Bones that could be identified at generic level include 17 specimens a temperate climate and is indicative of a landscape with (8.1%). Regarding the dental elements, of the 35 teeth specimens, 16

Table 6 Taxa representation from Marathousa 1 (Areas A and B, and total). NISP: Number of identiﬁed specimens [each distinct anatomical element is tallied as 1; Stiner (1994, p. 69) and Lyman (2008, pp. 34e35)].

Taxon Area A Area B both Areas

Bones Teeth Total Bones Teeth Total Total

NISP % NISP % NISP % NISP % NISP % NISP % NІSP %

Castor ﬁber 3 3.1 8 38.1 11 9.2 11 6.7 Lutra simplicidens 1 1.0 3 14.3 4 3.4 4 2.5 Felis sp. 1 4.3 1 2.3 1 0.6 Vulpes sp. 1 4.3 1 2.3 1 0.6 Canis sp. 2 9.5 2 4.5 2 1.2 Palaeoloxodon antiquus (skeleton) 74 75.5 3 14.3 77 64.7 77 47.2 ?Palaeoloxodon antiquus (?skeleton) 17 17.3 17 14.3 17 10.4 Palaeoloxodon antiquus 8 38.1 4 17.4 12 27.3 12 7.4 Hippopotamus antiquus 2 9.5 2 1.7 2 8.7 2 4.5 4 2.5 Bison sp. 1 4.8 1 2.3 1 0.6 Dama sp. 5 23.8 6 26.1 11 25.0 11 6.7 Cervus elaphus 2 9.5 2 1.7 1 4.8 3 13.0 4 9.1 6 3.7 Cervidae indet. 3 3.1 3 14.3 6 5.0 4 19.0 6 26.1 10 22.7 16 9.8 Total 98 21 119 21 23 44 163 Indeterminate 38 7 45 190 12 202 247 Total 136 28 164 211 35 246 410

are complete (45.7%), while 17 could be identified to genus (48.6%) stony substrate, as well as the minor transportation of the bones (Table 6). and their relatively fast burial/limited time of exposure. Carnivore The bone surfaces from both Area A and B do not show traces of tooth marks have usually a U-shaped cross-section (scores; rounding, which suggests limited, if any, transport over a short Blumenschine, 1995) and carnivore gnawing in general is so far not distance and/or for a short time span. Although the number of recognized in any of the elephant bones in Area A. Moreover, the bones in Area A (excluding the elephant skeleton) is much lower excavation of bones was carried out with wooden tools, so that the than in Area B, similar weathering percentages are obtained possibility these marks (which were originally also filled with fine (Fig. 7c): the majority of the bones in both Areas present rather sediment) to have a recent origin should be excluded. Cut marks on well-preserved surfaces with minor cracking and flaking [weath- the astragalus are possibly related to the cutting of the joints/lig- ering stages 0e2ofBehrensmeyer (1978)]. This indicates that most aments, and the disarticulation of the tarsals, probably to exploit bones were buried relatively quickly and/or were kept moist and the fat content and the soft tissues enclosed in the distal part of the protected by the lake vegetation [corroborated by the diverse fossil hindlimb and the foot cushion [Weissengruber et al., 2006; see also floral assemblage (Field et al., this issue) and the palae- Saunders and Daeschler (1994) for the disarticulation of mam- oenvironmental reconstructions of the wider region during that moths' forelimbs including carpals]. On the other hand, the location time (e.g., Nickel et al., 1996)]. of the tibia cut mark on the mid-shaft in the plantar side of the The good state of the bone preservation allows the identification of bone, where strong muscles are developed (Weissengruber and taphonomic modifications. Human modifications on the MAR-1 Forstenpointner, 2004), could possibly be attributed to filleting. bones indicative of butchering activities include percussion damage Although cut marks on proboscidean bones are rare (see below), a and cut marks. The former is created during the knapping of bones, cut-marked astragalus (on the proximal articular facet for the tibia) most commonly related to marrow processing (and is also associated was recently recorded at the early Homo site HWK EE (Olduvai with the overall high bone fragmentation and anthropic fractures), Gorge, Tanzania; Pante et al., in press). Cut marks on mammoth while the latter indicates carcass disarticulation and meat removal. tibiae have previously been reported at the Late Pleistocene sites Cut marks were identified on elephant bones belonging to the Yudinovo (Russia; Germonpre et al., 2008), Dent and Clovis (U.S.A.; skeleton from Area A, as well on elephant and other mammal bones Saunders and Daeschler, 1994; Saunders, 2007). from Area B. Specifically, so far identified cut-marked elephant Elephant bones from Area B showing cut marked surfaces bones from the skeleton include an astragalus (MAR-1-939/674-46) include a broken rib fragment (MAR-1-933/595-71), which pre- and a tibia (MAR-1-938/673-19). The astragalus shows two almost sents on the ventral side a single V-shaped mark, as well as a cluster parallel striations located on its distal side, on the lateral articular of scrape marks [see e.g., Rodríguez-Hidalgo et al. (2017, Fig. 6b) for facet for the calcaneus (Fig. 8a and b); while the tibia presents a similar marks on a bison bone] (Fig. 9aed). All these traces are single, wider groove, located on the plantar side of the diaphysis, present only locally with the rest of the bone lacking any other and obliquely oriented relative to the long axis of the bone (Fig. 8g). signs of marks. Cut marks (although of a different pattern compared The astragalus and tibia striations are almost straight and V-shaped to the MAR-1 ones) on the ventral side of an elephant rib from in cross-section, they show Hertzian cones and they partially pre- Aridos 2 (Spain) have been attributed to evisceration, which occurs serve internal microstriations (Fig. 8). These characteristics are at the very early stages of carcass consumption (Yravedra et al., observed with high frequency in cut marks (e.g., Domínguez- 2010). The intensive effort applied in the form of the observed Rodrigo et al., 2009; Fernandez-Jalvo and Andrews, 2016), cluster in the MAR-1 rib might be related to the removal of the including those created during exploitation of proboscidean car- periosteum during butchery or to the cleaning of the bone from fat casses (e.g., Yravedra et al., 2010; Sacca, 2012). Trampling, which before breaking it (Voormolen, 2008; Starkovich and Conard, 2015; mostly creates sinuous trajectories with flat- and broad-based Fernandez-Jalvo and Andrews, 2016; Gingerich and Stanford, in grooves in cross-section (Domínguez-Rodrigo et al., 2009, 2010), press and references cited therein). Indeed, on the dorsal side of is a less likely cause for these modifications, due to the low abrasive the rib, the absence of cortical layer (classic peeling) in the frac- nature of the sediment bearing the fossils and the absence of a tured region (Fig. 9e) indicates that the bone was broken when it

Fig. 7. a, composition of the Marathousa 1 large mammal fauna; b, frequency histogram of maximal diameter of the large mammal bones from Area B with ﬁtted normal distribution curve; c, comparison between Area A and Area B of the weathering stages observed in the large mammal bones. Weathering stages according to Behrensmeyer (1978).

was still fresh. Butchering marks on the rib accompanied by peeling deer, showing four short and parallel striations (Fig. 9f and g). Their indicates early access to an elephant carcass by hominins (Pickering location is possibly related to the disarticulation of the joint be- et al., 2013). tween the radius and the carpals (Rabinovich et al., 2008; Soulier Apart from elephants, ungulate exploitation is indicated by a and Costamagno, 2017). Furthermore, a spiral fractured humerus cut-marked distal radius epiphysis (MAR-1-933/601-6) of a fallow diaphysis fragment of a small/medium-sized mammal (MAR-1-

Fig. 8. Modifications on Palaeoloxodon antiquus bones from the Marathousa 1 (Area A) skeleton. aef, right astragalus, MAR-1-933/591-71; a, distal view, in the rectangle the region of the cut marks, shown in ‘b’; b, close view of the cut marks; c, detail of the cut marks, dashed line indicates the location of the profile shown in ‘d’; d, profile of the cut mark at 50% of its length; e, detail of the morphology of the cut mark in 3D; f, 3D model of the cut marks; gek, left tibia, MAR-1-938/673-19; g, plantar view (proximal to the right), in the rectangle the region of the cut mark; h, detail of the cut mark, dashed line indicates the location of the profile shown in ‘i’; i, profile of the cut mark at 30% of its length; j, detail of the morphology of the cut mark in 3D; k, 3D model of the cut mark.

Fig. 9. Modifications on bones from Area B. aee, Palaeoloxodon antiquus, rib fragment, MAR-1-933/591-71; a, ventral view, in rectangle the region of modification shown in ‘b’; b, detail of the rib showing the cut mark (rectangle, originally filled with sediment) and the scrape marks; c, detail of the cut mark, dashed line indicates the location of the profile shown in ‘d’; d, profile of the cut mark at 60% of its length; e, classic peeling, detail of the dorsal side of the rib showing the region of missing cortical layer; feg, Dama sp., distal fragment of left radius with cut marks, MAR-1-933/601-6; f, caudal view; g, detail of the cut marks; hei, spiral fractured humerus diaphysis fragment of small/medium-sized mammal with cut marks and percussion damage, MAR-1-933/600-113; h, medullary view; i, cortical view and detail of the cut marks.

933/600-113), shows oblique cut marks relative to the axis of the from Ambelia and Perdikkas (Western Macedonia; see Tsoukala bone and a percussion notch with its negative scar (Fig. 9h and i). et al., 2011) among the most complete skeletons of this species Finally, the presence in Area B of bone flakes/tools, as well as of a ever found in Greece. Apart from the elephant, other macro- bone shaft fragment (possibly belonging to an elephant) with flake mammals are so far poorly represented in MAR-1; however, Canis and impact scars, is indicative of bone knapping (Tourloukis et al., sp. [reported also in Athanassiou et al. (this issue)], Mustela sp., Felis this issue a). sp. and Lutra simplicidens are recorded for the first time in the Carnivore gnawing is recorded in Area B and includes one Megalopolis Basin, while L. simplicidens is documented for the first spinous process of an elephant vertebra (forrowing, crenellation, time in Greece. salivary rounding; Fig. 4o), one diaphysis of a cervid radius (salivary During recent decades a significant number of localities doc- rounding) and one mandibular (corpus) fragment of fallow deer, umenting proboscidean exploitation by hominins has been which presents single and overlapping notches on its ventral side discovered (Santucci et al., 2016, table 4 and references cited (Fig. 4men). Although overlapping notches may result from dy- therein), considerably enriching our knowledge on the subsistence namic loading (hammerstone percussion), they are produced in strategies of Homo and their interactions with megaherbivores. The higher frequency through static loading (carnivore pressure) due to preliminary dating of Marathousa 1 to ca. 0.5e0.4 Ma places the the close-set carnivore teeth and the simultaneous bite of adjacent locality among the oldest in Europe, where both lithic artefacts and teeth cusps, creating closely spaced impact points (Domínguez- cut marks are associated with a butchered elephant. It is also the Rodrigo and Barba, 2007). Moreover, in contrast to humans who first such locality to be discovered in Southeastern Europe. The usually break mandibles transversely into segments and produce recovery of additional faunal material from MAR-1 in the future will rather clean and straight fractures, carnivores produce irregular be essential for refining the biochronological, palaeoecological and edges (Stiner, 1994, pp. 140e144). taphonomic interpretation of the site, enhancing our understand- Cut marks on the MAR-1 elephant skeleton, as well as in other ing of the Lower Palaeolithic hominin food acquisition strategies elephant and mammal bones, indicate butchering activities by and subsistence behavior, as well as enriching our knowledge about humans. This is in accordance with the traits of the lithic industry human-proboscidean interactions in the Palaeolithic. and its spatial association with the mammal bones (Giusti et al., this issue; Panagopoulou et al., this issue; Tourloukis et al., this issue a). Acknowledgments Cut marks occur in mammals ranging in size from the small/medium fallow deer to elephant (megafauna), indicating a rather wide This research was supported by the ERC StG no. 283503 PaGE spectrum of exploitation by humans. The fact that carnivore and by the ERC CoG no. 724703 CROSSROADS awarded to K. Harvati, gnawing occurs in the same taxa as cut marks could potentially the University of Tübingen (2016 field season) and the “Minister- suggest a certain degree of carnivore-human competition for early ium für Wissenschaft, Forschung und Kunst Baden-Württemberg”. access to the animal carcasses. We thank S. El-Zaatari, Y. Sahle, B. Starkovich (University of Proboscideans, the largest terrestrial animals during the Pleis- Tübingen) and G. Koufos (University of Thessaloniki) for their help tocene, constituted an ideal food package and one of the most during the research. We are grateful to all the field team members attractive targets for early humans (Ben-Dor et al., 2011; Reshef and for their contribution during the excavations, and to the Greek Barkai, 2015; Agam and Barkai, 2016). The human engagement in Ministry of Culture, the Municipality of Megalopolis, the authorities proboscidean assemblages, as well as the degree of their interaction of the Peloponnesus Prefecture and the Public Power Corporation is not always straightforward. For the MAR-1 skeleton in particular, S.A. for their support. We thank E. Tsoukala (University of Thessa- the small number of cut marks observed to date does not neces- loniki) and two anonymous reviewers for making constructive sarily indicate that the human activity was minor, as cut marks are comments and suggestions leading to an improvement of the rarely produced during the exploitation of large carcasses. 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