Pleistocene vertebrates from the Kyparíssia lignite mine, Megalopolis Basin, S. Greece: Testudines, Aves, Suiformes

Accepted Manuscript Pleistocene vertebrates from the Kyparíssia lignite mine, Megalopolis Basin, S. Greece: Testudines, Aves, Suiformes Athanassios Athanassiou, Dimitris Michailidis, Evangelos Vlachos, Vangelis Tourloukis, Nicholas Thompson, Katerina Harvati PII:

S1040-6182(18)30230-1

DOI:

10.1016/j.quaint.2018.06.030

Reference:

JQI 7490

To appear in:

Quaternary International

Received Date: 25 February 2018 Revised Date:

19 June 2018

Accepted Date: 20 June 2018

Please cite this article as: Athanassiou, A., Michailidis, D., Vlachos, E., Tourloukis, V., Thompson, N., Harvati, K., Pleistocene vertebrates from the Kyparíssia lignite mine, Megalopolis Basin, S. Greece: Testudines, Aves, Suiformes, Quaternary International (2018), doi: 10.1016/j.quaint.2018.06.030. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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Pleistocene vertebrates from the Kyparíssia lignite mine, Megalopolis Basin, S. Greece: Testudines, Aves, Suiformes

Athanassios ATHANASSIOU1, Dimitris MICHAILIDIS2, Evangelos VLACHOS3,

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Vangelis TOURLOUKIS4, Nicholas THOMPSON4, Katerina HARVATI4

Ministry of Culture, Ephorate of Palaeoanthropology–Speleology, Ardittou 34B, 11636 Athens, Greece

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American School of Classical Studies at Athens, Wiener Laboratory for

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Archaeological Science, Souidias 54, 10676, Athens, Greece 3

CONICET and Museo Paleontológico Egidio Feruglio, Av. Fontana 140, 9100,

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Trelew, Chubut, Argentina

Eberhard Karls University of Tübingen, Palaeoanthropology, Senckenberg Center for Human Evolution and Palaeoenvironment, Rümelinstr. 23, 72070 Tübingen,

Abstract

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Germany

The mining activities in the Middle Pleistocene lacustrine basin of Megalopolis (Peloponnesus, Greece) have exposed expanded sections of lacustrine sediments. In

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particular, the northernmost mine of Kyparíssia has yielded numerous vertebrate fossils, recovered during field surveys and small-scale rescue excavations. The stratified specimens indicate the presence of at least two fossiliferous horizons, which

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are stratigraphically closely situated. The fossils are generally widely dispersed, but they are much more abundant along the western margin of the palaeolake. Four sites showing denser concentration of fossils were located and partially excavated during the 2004–2008 fieldwork. An additional site was identified and excavated in 2012. The fossils were found in organic-rich sediments (mostly lignites), indicating that they were deposited during a warm and humid (i.e., interglacial) time period, in a richly vegetated environment. The recovered fauna is diverse, dominated by hippopotamuses, deer and elephants. The present study presents the chelonian, avian and suiform finds. The following taxa are recognised: Emys orbicularis, Testudo marginata, Sus scrofa, Hippopotamus

ACCEPTED MANUSCRIPT antiquus, as well as a diverse sample of aquatic birds. This faunal composition indicates a temperate, woodland/forest environment, with continuous presence of a large water body, whilst the presence of the darter (Anhinga sp.) points to relatively warm conditions. Biochronologically it points to an age within the early part of the

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Middle Pleistocene.

1. Introduction

The Megalopolis Basin is a lowland area of extensional tectonic origin, situated in

central Peloponnesus and extending along a NW–SE direction (Fig. 1). Its basement

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consists of Mesozoic limestones and Palaeogene flysch that belong to Píndos and

Tripolis geotectonic zones of the alpine orogen. Today the basin is drained by the hydrographic network of the Alpheiós River. This is, however, a fairly recent

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development, as the basin was lacustrine for a considerable part of its Pleistocene history, witnessed by the extensive sedimentary fill of lacustrine deposits. The good preservation potential of the lacustrine environment has provided abundant data for reconstructing the environment of the area, mainly by means of plant (wood, fruits, seeds and microremains) and animal (mainly mammalian) fossils. The main

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conclusion of the already published studies on the Megalopolis area is that the sedimentation in the basin started possibly during the Pliocene and continued as an alternation of fluvial and lacustrine deposits till the end of the Middle Pleistocene (Vinken, 1965; Tsiftsis, 1987; van Vugt et al., 2000). (Note, however, that the

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definition of Pliocene was different during the 1960s, this epoch being partly equivalent to what is currently the Early Pleistocene; nonetheless, a vague reference

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to Pliocene strata in the basin was retained by later authors). An extensive, more recent, alluvial fan covers the sequence. The Middle Pleistocene series is characterised by an alternation of detritic and biogenic sediments that correspond to cold/arid (glacial) and warm/humid (interglacial) climatic conditions (Nickel et al., 1996; van Vugt et al., 2000; Okuda et al., 2002). The presence of scanty vertebrate fossils in the area of Megalopolis is known to the scientific community at least since 1860 (according to the University of Athens archives; Mitzopoulos et al. 1862), but the fossil fauna became well known only in 1902, when Prof. Theodore Skouphos (University of Athens) conducted palaeontological excavations in two or three sites within the Megalopolis Basin, in the area of the villages Karyés and Ísoma (Bürchner, 1903; Skuphos, 1905; Melentis,

ACCEPTED MANUSCRIPT 1961; Sickenberg, 1976). Subsequent geological prospecting carried out in 1960 and 1962–1963 in the whole basin by a German team, working on the assessment of the mining potential of the lacustrine deposits, produced additional, non-stratified palaeontological finds. Most of the available material derived from these early field expeditions was described in a series of publications by Melentis (1961; 1963; 1966a–

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g) and Sickenberg (1976). The published fauna is diverse and indicates a forest/woodland environment in a temperate climatic context. It is stressed, however, that this fauna lacks stratigraphic context, thus the resulting biochronological and palaeoecological interpretations are questionable.

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The rich large mammal samples, including a human specimen, that have been

excavated or collected in the past, have recently regenerated an intense research interest for the basin’s palaeontological and archaeological potential. Recent large-

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scale fieldwork in the basin (palaeontological / palaeoanthropological prospecting and excavations) was carried out by the University of Athens (Theodorou, 2014), as well as by a joint team of the Ephorate of Palaeoanthropology–Speleology of Greece and the University of Tübingen under the European Research Council project “Palaeoanthropology in the Gates of Europe” (PaGE) (Panagopoulou et al., 2015). In

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the frame of the latter project, an extensive part of the Megalopolis Basin was surveyed for palaeoanthropological sites (Thompson et al., this issue). Since 1969, the exploitation of the economically important biogenic deposits (lignite seams) in open-cast mines for electricity production has considerably facilitated the

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discovery of new sites, as many sections are continuously cut through the sedimentary sequence, making the layers much more accessible in horizontal extent and depth. The

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mining activities have taken place in four fields, the Chorémi, the Marathoúsa, the Thoknía, and the Kyparíssia mine (Fig. 1b). The former two are the main operating mines today. Thoknía is closed and backfilled with quarry waste since many years, whereas Kyparíssia, the northernmost mining field, was in full operation till 2006 and it operates thenceforth only occasionally. It was during the mining operations that the first mammalian fossils were located in the Kyparíssia mine: a section collapse in October 2004 revealed skeletal remains of an elephant. Subsequent fieldwork from 2004 to 2008 by the first author, on behalf of the Ephorate of Palaeoanthropology– Speleology (Hellenic Ministry of Culture), produced a wealth of fossil skeletal elements of mammals, birds and turtles. Four main fossiliferous sites were located within the mine (KYP1–KYP4, see below), which looked quite promising for future

ACCEPTED MANUSCRIPT excavations, but numerous other minor findspots were also tracked down. However, no large scale excavation was carried out, while many of the collected specimens are unstratified surface finds. More recently, during the PaGE Project survey conducted in 2012, a new site was found in the mine (KYPT, see below), very close to the village of Kyparíssia. Although it is not quite rich in fossils, it yielded a nice sample

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of Hippopotamus skeletal elements. The present paper is a comprehensive study of the most complete identifiable

specimens of turtles, birds and suiform mammals, collected or excavated in the

Kyparíssia mine from 2004 to 2008, as well as in 2012. Together with the avian

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material from the Marathoúsa mine (Michailidis et al., this issue) this is the first report on the avian remains of the Megalopolis Basin. The rest of the mammalian material, other than Suiformes, deriving from the same research is described in Athanassiou

and Athanassiou et al. (2016).

2. Material and methods

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(this issue). Preliminary reports on both studies were presented by Athanassiou (2016)

All the osteological and dental material presented in this study is stored in the

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collections of the Ephorate of Palaeoanthropology–Speleology (Ministry of Culture, Athens, Greece). Because of the fieldwork conditions (including collection of poorly stratified specimens within the highly disturbed environment of a lignite quarry) most of the available specimens are highly fragmentary. Consequently only a part of the

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collected sample is described below, that is those specimens that are fairly complete and attributable to a taxon. For the reader’s convenience, the catalogue numbers of the

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in situ specimens mentioned in this study include the site number where available (e.g., KYP3-456), though the actual specimens in the collection are invariably labelled as KYP (e.g., KYP-456). Surface finds are simply referred to as KYP throughout the text.

The description of the mammalian finds is based on standard anatomical terminology (König and Liebich, 2004; I.C.V.G.A.N., 2005). The anatomical terminology of the avian remains follows Baumel et al. (1993) and the systematics is according to del Hoyo (1992). The comparative osteological collections of the Museum of Palaeontology and Geology (National and Kapodistrian University of Athens), the Malcolm H. Wiener Laboratory for Archaeological Science (American School of Classical Studies, Athens) and the Natural History Museum of Vienna, Austria, were

ACCEPTED MANUSCRIPT used for the identification of the specimens. All measurements are in mm with an accuracy of up to one decimal digit, when possible. The use of parentheses in the tables denotes an inaccurate or estimated measurement because of specimen distortion or incomplete preservation. To avoid ambiguities about the cited sources, citations to figures, plates, tables etc. of published papers are given in lowercase (e.g., pl. 1,

letter (e.g., Fig. 1, Pl. 1, Fig. 1, Tab. 1).

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fig.1), while citations to figures etc. of the present paper are given with a capital first

Anatomical and dimensional abbreviations: I/i upper/lower incisor; C/c

upper/lower canine; M/m: upper/lower molar; P/p: upper/lower premolar; DAP:

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craniocaudal (anteroposterior) diameter (p: proximal; d: distal; a: articular); L: length; DT or W: transversal diameter (p: proximal; d: distal; a: articular; lat: lateral; med:

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medial); H: height.

3. Geological setting and Taphonomy

The Kyparíssia mine occupies the NW area of the Megalopolis Basin, extending on a basement consisting of Jurassic – Early Cretaceous cherts and Late Cretaceous limestones of the Píndos geotectonic zone (Papadopoulos et al., 1997). The basin

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sedimentary fill in the area consists of organic-rich lacustrine sediments, mainly of a thick lignite sequence, lacking the intermediate detritic layers, which are found in the central and southern areas of the basin. On top of the lignite there is a clayey cover of colluvial origin, which extends along the western margin of the basin. The central part

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of the basin is covered by terrace deposits of the Alpheiós River. Following the stratigraphic scheme detailed by Löhnert and Nowak (1965) and Vinken (1965), the

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sedimentary fill at Kyparíssia belongs to the Marathousa member of the Chorémi Formation. All sites are located near the top of the preserved palaeolake sedimentary sequence in this part of the basin, at an altitude of 330–340 m. Their exact position with regard to the stratigraphy of the whole basin is, however, difficult to assess, mainly because the characteristic lignite seams, which are distinct in the more southern parts of the basin, are merging to each other towards the North, forming the thick mostly undivided biogenic deposit without any significant detrital intercalations. Löhnert and Nowak (1965), based on extensive borehole records, correlate this lignite deposit with the lignite seams I and II, implying that the stratigraphically upper seam III is missing in the north of the basin and in Kyparíssia mine in particular. Indeed, a calcareous–marly layer situated at the base of the KYPT section (see below; Fig. 2a,

ACCEPTED MANUSCRIPT c) may be identified as the thin limestone layer that according to Löhnert and Nowak (1965) characterises the top of the lignite seam II. However, this layer is not present or exposed in the other sites at Kyparíssia, which would have been closer to the lake shore. The fossils crop out mainly along the western margin of the mine, that is along the

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western margin of the Megalopolis palaeolake, while they are extremely rare in the central and the eastern parts of the mine. They are usually found isolated at numerous findspots. Four areas that were characterised by increased density of finds were

dubbed sites 1 to 4 (KYP1 to KYP4) (Fig. 1b). Their geographic coordinates are the

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following: KYP1: 37.4526°N, 22.0696°E; KYP2: 37.4569°N, 22.0697°E; KYP3:

37.4541°N, 22.0686°E; KYP4: 37.4537°N, 22.0696°E (WGS84 datum). KYP1 and KYP2 were partially exploited in 2004 but were subsequently destroyed or buried as a

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result of the mine development. KYP3 and KYP4 still exist and are promising for future research by means of a systematic excavation.

KYP1 is the initially located section-collapse site. It is a sequence of mainly darkcoloured lignite layers with intercalating grey sandy clays of variable thickness. The main grey layer has a thickness of about 3 m, situated at an altitude of 340 m. An

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underlying lignitic layer yielded numerous Hippopotamus specimens that probably belonged to the same individual (see Section 4.4). Due to the disturbance of the site the stratigraphic relationship and the number of the existing fossiliferous horizons are not clear. The most probable reconstruction based on field data includes two

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fossiliferous horizons at about 340 m and 330 m above sea level. KYP2 is the northernmost one at an altitude of 340 m. It was potentially very rich in

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fossils preserved within lignite, as shown by preliminary fieldwork, but it was destroyed before it could be excavated. This site did not yield fossils attributable to the taxa studied in the present paper. KYP3 includes a rich fossiliferous lignitic layer at 340 m above sea level, which was horizontally exposed over an area of about 300 m2. Part of this exposure was covered later because of the construction of a temporary road. The lignitic layer lays on an at least 1-m-thick green-brownish sandy clayey layer with numerous small (diameter of about 5–20 mm) angular pebbles in certain places. A few meters to the North, the clay becomes chestnut-brown and contains nodules and intercalations of pure green sand. The site also includes an additional fossiliferous horizon in a sandy clay layer, poor in finds but only visible in a section, situated about 2 m higher than the main one.

ACCEPTED MANUSCRIPT KYP4 is situated on top of a limestone outcrop, apparently very close to the palaeoshoreline, at an altitude of 330 m. It consists of lignite layers with intercalations of sandy clays. The fossil finds are not accumulated but were found dispersed. The lacustrine sediment outcrop at this site is limited, less that 2 m thick, as most of it has been already mined.

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The new site tracked down during the 2012 survey –dubbed Kyparíssia-T (KYPT)– is situated at the SW margin of the Kyparíssia mine (coordinates 37.4505°N, 22.0742°E; WGS84 datum). A section collapse close to the archaeological site of the ancient town of Trapezoús, very close to the modern village of Kyparíssia, revealed the presence of

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a small accumulation of fossil bones (Fig. 2b, c), belonging mainly to Hippopotamus. Additional scanty finds are referred to Testudines, Aves and Cervidae. The latter are not referable to an infra-familial taxon. Since most of the fossils were already

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exposed, a brief excavation was carried out, in order to recover the finds and document the site. KYPT exhibits a sequence of rather thin layers, including a whitecoloured calcareous one, which may be identified as the characteristic limestone layer at the top of Lignite Seam II (Löhnert and Nowak, 1965, p. 860).

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3.1. Preservation

Most recovered specimens are very well preserved in terms of surface quality (only minor cracking is observed in certain samples), but may have suffered recent fragmentation and surface wear due to operations of the mining machinery. A couple

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of avian specimens show evidence of weathering due to surface exposure [weathering stage 1, slightly weathered (Bochenski and Tomek, 1997)], or pitting caused by

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digestion. Very few isolated surface finds show signs of eroded surfaces and edge rounding because of rolling wear, but this seems to be quite recent and attributed to specimen exposure and transportation caused by the mining operations. Carnivore damage of the collected specimens is quite uncommon in all sites of the Kyparíssia mine; it is well documented only in one surface find (a rhinoceros astragalus; Athanassiou, this issue). An avian specimen (KYP-516, Fig. 5E) shows signs of digestion through the pitting observed on its surface, indicating that this individual fell victim to a bird of prey. Most recovered remains were isolated and there is no evidence of any anatomical association, except for certain cases mentioned below. As a whole, the find assemblages at all sites appear to have been deposited locally and relatively quickly.

ACCEPTED MANUSCRIPT KYP1 and KYP3 yielded specimens that quite plausibly belong to individual megaherbivore skeletons, as indicated by dimensional data, ontogenetic stage and physical appearance (mineralisation, colour, preservation quality), but no relevant in situ taphonomic evidence is available, because the sites were highly disturbed. The specimens recovered from both horizons at KYP1 apparently were in pristine

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condition when still in situ, but the subsequent section collapse caused severe fragmentation. The fossil preservation at KYP2–KYP4 varies according to local geochemical conditions and the effects of the mining operations. However, no

specimen exhibits conspicuous surface cracking or flaking, indicating quick burial

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without exposure to dry aerial conditions or to any stresses associated with high

temperature and moisture changes (Behrensmeyer, 1978). The avian elements from KYP3 identified to the mute swan were found in very close proximity to each other

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and quite probably represent a single partially preserved skeleton, which was quickly deposited. The finds of KYPT exhibit a higher degree of syndepositional fragmentation (Figs 2B, 3), which could have possibly happened due to trampling by animals moving in the lake close to the shore (usually hippopotamuses).

4.1. Turtles

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4. Systematic Palaeontology

Fossil turtles from the Pleistocene of Megalopolis area are known since the work of Melentis (1966g), which presented material that originated from excavations made by

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the University of Athens in 1902. Recently, Vlachos and Delfino (2016) revised the material of Melentis (1966g) based on his published documentation; the material was

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not able to be located for first-hand observations (Vlachos and Delfino, 2016, p. 160). They also presented new fossils from the area of Megalopolis, based on recent collections made by the Utrecht University (The Netherlands). The material from both collections contains shell fragments of two aquatic turtle species, the emydid Emys orbicularis (Linnaeus, 1758) and the geoemydid Mauremys rivulata Vallenciennes in Bory de Saint-Vincent (1833). The new material herein consists of several plates of the shell, mostly fragmented, but all well-preserved. In most cases, the specimens can be identified in detail, which allows further confirming the presence of Emys orbicularis in the area of Megalopolis. Additionally, the KYP turtle material contains few specimens that can be clearly attributed to the tortoise Testudo marginata Schoepff, 1793. This taxon was

ACCEPTED MANUSCRIPT previously unknown from Megalopolis region, but it has been found in other localities of the Peloponnesus peninsula (Vlachos and Delfino, 2016 and references therein). Collectively, the area of Megalopolis appears to be the most diverse in terms of chelonian diversity in Peloponnesus, with three species of freshwater and terrestrial turtles. As all these species form part of the extant turtle fauna of the peninsula, we

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can demonstrate that all extant continental turtle clades from Peloponnesus were already present in the Middle Pleistocene of Megalopolis. The studied material joins the list of few localities with turtle remains in Peloponnesus and enhances our

Order: Testudines Batch, 1788 Sub-order: Cryptodira Cope, 1868

Genus: Emys Duméril, 1805

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Family: Emydidae Rafinesque, 1815

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knowledge of diversity and distribution of turtles in the Pleistocene of Greece.

Emys orbicularis (Linnaeus, 1758)

Material: KYPT-833: nuchal; KYPT-856: partial neural; KYPT-834: suprapygal;

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KYP3-37, partial costal of a juvenile; KYP-10, KYP-547: partial costal; KYP-360, KYP3-656: peripheral; KYP-512: left epiplastron; KYP3-66: right hyoplastron fragment; KYP1-663: left hypoplastron; KYP-746: ?hypoplastron fragment; KYP715: ?hypoplastron fragment; KYP-361: partial right xiphiplastron; KYP3-39, partial

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fragment.

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right xiphiplastron; KYP-517, KYP-843, KYP-747, KYP-523, KYP3-661: shell

Description: The material consists of several isolated shell plates (Fig. 4A–L), which allow the documentation of the various diagnostic characters of the species (see Vlachos and Delfino, 2016). Based on the preserved elements, the morphology of large parts of the shell could be reconstructed. The nuchal is as long as wide, with a cervical scute that is longer than wide and narrower anteriorly. There is no evidence that the pleural 1 covers the lateral parts of the nuchal. The single preserved neural appears to be hexagonal with short antero-lateral sides and is not crossed by any sulcus. The various costals are all laterally longer, therefore being consistent with the above-mentioned neural morphology. The peripherals are short and are covered both by the pleural and marginal scutes. The second suprapygal is wider than long, with a

ACCEPTED MANUSCRIPT trapezoid shape; it is not crossed by any sulcus, thus the vertebral 5 expands on the pygal. The border of the anterior lobe is rounded and smooth. There is no epiplastral lip, and the gular scutes are narrow, covering the anterior part of the entoplastron. Based on the preserved hyoplastron, the humero-pectoral sulcus is probably crossing the posterior part of the entoplastron. As it is evident both the hyoplastron and the

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hypoplastron, a hinge was present between these two plates: the posterior border of the hyoplastron and the anterior of the hypoplastron are smooth, and there is an

overlap of the pectoro-abdominal sulcus with the hyo-hypoplastral suture; this is the

main diagnostic character that allows the attribution to Emys orbicularis. The border

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of the posterior lobe is rounded and the anal scutes are medially longer than the

Family: Testudinidae Batsch, 1788 Genus: Testudo Linnaeus, 1758

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femoral ones.

Testudo marginata Schoepff, 1793

Material: KYP-744: nuchal; KYP-78: partial costal; KYP-753: partial anterior

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peripheral; KYP1-673: bridge peripheral; KYP3-87: 7th right peripheral; KYP-748: two articulated posterior peripherals, probably the 8th and 9th from the right side; KYP-513: left xiphiplastron.

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Description: The material attributed to this taxon consists of few plates from the shell, mostly from the carapace (Fig. 4M–S). The morphology of the material is consistent

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with a testudinid taxon and Testudo marginata in particular, both in size and morphology (see Vlachos and Delfino, 2016). The nuchal is as long as wide, with a narrow anterior border and shallow nuchal notch. The cervical scute is long and narrow. The costal plate is not crossed by any sulcus, but it is laterally short and medially long, thus consistent with a differentiated neural series consisting of quadrangular and octagonal neurals. Several peripherals are known, one from the anterior border, one from the bridge and three from the posterior border. The bridge peripheral is tall, much taller than long. In the seventh peripheral a remnant of the inguinal scute is noted. All peripherals are covered only by the marginal scutes and there is a clear coincidence between the pleuro-marginal sulcus and the costoperipheral suture. The peripherals of the posterior border are elongated, and the ninth

ACCEPTED MANUSCRIPT one appears to develop a posterior flaring; this is the main diagnostic character that allows the attribution to Testudo marginata. The preserved xiphiplastron shows evidence of medially long anals and an anterior border that is smooth and morphologically consistent with the presence of a hypo-xiphiplastral hinge. Based on the preserved material we can estimate the presence of at least one tortoise individual

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as there are no overlapping parts of the shell.

4.2. Birds

A total of 22 skeletal elements have been retrieved from KYP3, KYP4 and KYPT, as

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well as from disturbed findspots, of which 18 were identified as belonging to 9

species in 3 orders and 4 families. It was not possible to identify the remaining 4 skeletal elements further than class, due to their fragmentary preservation. The family

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Anatidae dominates in the assemblage with 15 skeletal elements belonging to 6 species, ranging in size from the teal (Anas crecca) to the mute swan (Cygnus olor). The family Rallidae is represented by the common coot (Fulica atra), Anhingidae by a darter identified only to genus (Anhinga sp.) and the Phalacrocoracidae by the great

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cormorant (Phalacrocorax carbo).

Order: Anseriformes Wagler, 1831 Family: Anatidae Leach, 1820 Genus: Anas Linnaeus, 1758

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Anas crecca Linnaeus, 1758

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Material: KYP-51: left femur, distal end. Description: In caudal aspect, the corpus femori of KYP-51 (Fig. 5N) appears straight and relatively gracile. The fossa poplitea is deeply excavated, the condylus medialis is flattened and the trochlea fibularis relatively shallow. In lateral aspect, the condylus lateralis is relatively robust and the epicondylus lateralis quite marked. All these features can be observed in extant Anas crecca, with which the Kyparissia specimen is also metrically identical.

Anas platyrhynchos Linnaeus, 1758

ACCEPTED MANUSCRIPT Material: KYP-511: right ulna, distal end; KYP4-338: right carpometacarpus, proximal end. Description: In dorsal aspect, the distal ulna (Fig. 5O) appears robust, the incisura tendinosa is well marked and deeply excavated and the condylus ventralis ulnaris is weak. In ventral aspect, the depression radialis is weak and the sulcus intercondylaris

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relatively wide. In dorsal aspect, the carpometacarpus (Fig. 5K) preserves a strongly excavated fossa supratrochlearis, extending distally, a marked fovea carpalis caudalis, with a distinct depression distally and a strong sulcus tendinous, crossing the os

metacarpale majus medio-laterally. In ventral aspect, the trochlea carpalis is relatively

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shallow, the fossa infratrochlearis well excavated, the processus piciformis is robust

and the processus extensorius extends strongly laterally. All of the above features are expressed in the same way in extant Anas platyrhynchos and metrically both skeletal

Genus: Cygnus Garsault, 1764

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elements are also alike extant A. platyrhynchos.

Cygnus olor (Gmelin, 1789)

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Material: KYP3-660: 5th cervical vertebra; KYP3-621: clavicula; KYP3-621: right humerus, proximal end; KYP3-621: left humerus, distal end; KYP3-621: right ulna, distal end; KYP3-621: left tibiotarsus, distal end; KYP3-621: right tibiotarsus, distal end.

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Description: All remains referred to this species, except for the vertebra KYP3-660, were found in very close proximity (actually piled together) at KYP3 and they are

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metrically comparable; they are considered as belonging to a single individual. The corpus vertebrae and partially a zygapophysis cranialis and a zygapophysis caudalis of the 5th cervical vertebra are preserved. The foramen vertebrale is dorsoventrally depressed, the facies articularis cranialis rounded and the facies articularis caudalis dorsoventrally expanded. The clavicula (Fig. 5C) is heavily pneumatized, with strong scapus claviculae and relatively narrow extremitas sternalis claviculae. The right proximal humerus (Fig. 5J) preserves a strong ventral lip of the crista bicipitalis and a wide pneumatized fossa pneumotricipitalis in caudal aspect and an extensive and elevated intumescentia humeri in cranial aspect. The left distal humerus (Fig. 5A) preserves a strong epicondylus ventralis, while the fossa brachialis is constricted proximo-distally, but well excavated. The right distal ulna (Fig. 5B) preserves a

ACCEPTED MANUSCRIPT relatively slender corpus, a strongly excavated incisura tendinosa and a caudally extended condylus dorsalis ulnaris. The depression radialis is relatively small and circular. The better preserved left distal tibiotarsus (Fig. 5I), in cranial aspect has a robust corpus, a proximo-distally extended pons supratendineus with a deeply excavated and wide sulcus extensorius and a well demarcated tuberculum retinacula

extant Cygnus olor allow for a secure identification.

Genus: Mareca Stephens, 1824

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Mareca strepera (Linnaeus, 1758)

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fibularis. All of the characters listed, as well as the metric similarity of the findings to

Material: KYP-509: right coracoid, distal end; KYP-510: right humerus, proximal

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end.

Description: In dorsal aspect, KYP-509 (Fig. 5M) exhibits very marked intermuscular lines at the sternal end of the coracoid. The preserved portion of the cotyla scapularis shows a strong lip and a deeply concave cotyla and the medial end of the sternal extremity of the coracoid is robust. In cranial aspect, KYP-510 preserves a well

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excavated impressio coracobrachialis, the lip of the crista deltopectoralis is well marked and the corpus humeri is relatively robust. In size and morphology, both skeletal elements are identical to extant Mareca strepera.

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Genus: Spatula Boie, 1822

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Spatula clypeata (Linnaeus, 1758)

Material: KYP-799: right humerus, distal end; KYPT-819: left humerus, distal end. Description: In cranial aspect, KYPT-819 (Fig. 5F) shows a lightly curved and slender corpus humeri, an oval shaped and ventrally located fossa brachialis and distinct, deep muscle markings on the ventral side of the epicondylus ventralis. In KYP-799, the shape and size of the fossa brachialis is as described above, as well as the general character of the corpus humeri. Metrically and morphologically, they are both indistinguishable from Spatula clypeata.

Spatula querquedula (Linnaeus, 1758)

ACCEPTED MANUSCRIPT Material: KYP-750: right coracoid, complete. Description: In dorsal aspect, KYP-750 (Fig. 5L) appears relatively slender, with marked intermuscular lines at the sternal end of the coracoid, but not extremely so. The cotyla scapularis is deep and rounded and the processus procoracoideus relatively slender. The sulcus supracoracoidei is deeply excavated with marked lips and the

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impressio ligamenti acrocoracohumeralis is strong and narrow. The facies articularis sternalis is relatively shallow, but proximo-distally extended. Morphologically and metrically it is identical to extant Spatula querquedula.

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Order: Gruiformes Bonaparte, 1854 Family: Rallidae Rafinesque, 1815 Genus: Fulica Linnaeus, 1758

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Fulica atra Linnaeus, 1758

Material: KYP-516: right humerus, complete.

Description: In caudal aspect, KYP-516 (Fig. 5E) appears slender and elongated with a slight dorsal bent at the distal portion of the corpus humeri. The tuberculum dorsale

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is strongly developed and the crista deltopectoralis well defined and squared. The margo caudalis is fairly well elevated. In cranial aspect, the impressio coracobrachialis is well excavated, the condylus dorsalis rounded and the fossa brachialis relatively shallow. All of the above characters are expressed in the same

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way in extant Fulica atra, and metrically the KYP-516 is also similar to humeri of

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extant Eurasian coots.

Order: Suliformes Sharpe, 1891 Family: Anhingidae Reichenbach, 1849 Genus: Anhinga Brisson, 1760 Anhinga sp.

Material: KYPT-804: left humerus, complete. Description: The corpus humeri appears triangularly shaped corpus in caudal view, due to the pronounced margo caudalis, a feature typical of Suliformes (Fig. 5D). In caudal aspect of the proximal humerus, the tuberculum dorsale is distally placed and the distal end of the caput humeri extends distally. The incisura capitis is not deeply

ACCEPTED MANUSCRIPT excavated and is rounded in its distal border. The tuberculum ventrale is robust and dorsally rotated, the fossa pneumotricipitalis less excavated and the crus ventrale fossa relatively gracile. In cranial aspect, the intumescentia humeri appears very elevated and the impressio musculo brachialis strongly etched. The sulcus transversus is relatively shallow and short dorso-ventrally. In cranial aspect of the distal humerus,

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the distal epiphysis appears flattened and squared-off, as is typically the case in Suliformes the border of the fossa brachialis is relatively gracile and the epicondylus dorsalis extends proximally, higher than the condylus dorsalis. The epicondylus

ventralis is robust and has a strongly ventral rotation. The above features can be

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observed in both Anhinga rufa and A. melanogaster but not in Phalacrocorax carbo or P. aristotelis. The lack of additional skeletal elements preserved does not allow a

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more precise identification and the remains are thus identified as Anhinga sp.

Family: Phalacrocoracidae Reichenbach, 1850 Genus: Phalacrocorax Brisson, 1760

Phalacrocorax carbo (Linnaeus, 1758)

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Material: KYPT-812: left tarsometatarsus, complete.

Description: In dorsal aspect, KYPT-812 (Fig. 5G) appears robust, with very strong and dorsally projecting facies subcutanea lateralis, very deep fossa infracotylaris dorsalis and, relatively weak impressio retinacula extensorii. Distally, the foramen

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vasculare distale is large and mediolaterally extended. In proximal aspect, the area intercotylaris is mediolaterally extended and the cotyla lateralis deep and

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dorsoplantarly extended. In size and morphology it is exactly like the tarsometatarsus of extant Phalacrocorax carbo.

4.3. Suids

Order: Artiodactyla Owen, 1848 Family: Suidae Gray, 1821 Genus: Sus Linnaeus, 1758 Sus scrofa Linnaeus, 1758

Material: KYP-700: right c part; KYP3-81: left dentary fragment with m3; KYP-667: left distal humerus; KYP3-658, right proximal radius and ulna.

ACCEPTED MANUSCRIPT Description: The lower canine KYP-700 is partly preserved, lacking its tip and most of its basal part (Fig. 6D). Its preserved length along its labial margin is 115 mm (111 mm in straight line). Its mesiodistal diameter was more than 24 mm, but it is not measurable due to damage at the mesial edge of the crown. The tooth’s cross section

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forms a triangle, the longer side of which is the lingual one, and the shorter the labial one. The distal side bears two wear surfaces that expose the dentine, a distal and a distolabial one, the latter being confined to the basal part of the crown, tapering

towards the tip. The other two sides are covered by enamel. The lingual is slightly

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convex, while the labial is weakly concave because of the presence of a shallow groove that runs along the whole crown in this side. The enamel is rather thin, exhibiting longitudinal striations and developmental cross lines on its surface.

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Compared to published lower canine specimens, KYP-700 seems to be metrically very similar to those of males, despite its incomplete preservation. However, it appears to be less curved than any other canine specimen of Pleistocene S. scrofa (Hünermann, 1975; 1977; 1978; Guérin and Faure, 1997).

The single available molar, a left m3 (Fig. 6C), belongs to a rather aged animal, as the

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main cuspids are fairly worn, particularly the protoconid and the paraconid. The two transverse valleys are blocked by a large median secondary cuspid and much smaller tubercles situated labially in both valleys, and lingually and labially in the distal one. Its dimensions (L = 43.2 mm and W = 20.0 mm) are close to the maximal values of a

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recent sample of the species (Guérin and Faure, 1997) and within the size-range of published Middle–Late Pleistocene samples from Europe (Hünermann, 1975; 1977;

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1978; Faure and Guérin, 1983; Tsoukala, 1989; Guérin and Faure, 1997). The available long bones are partly preserved. They have the typical morphology of the genus. The humerus (Fig. 6B) has a very deep trochlear groove and a very large supratrochlear foramen that connects the olecranon and the radial fossae. The radius and the ulna (Fig. 6A) are fully fused to each other, except for the periarticular area. The humerus distal dimensions are DAPd = 51.0 and DTd = 53.0 mm. The radius proximal dimensions are DAPp = 24.5 and DTp = 39.7 mm. The proximodistal diameter of the trochlear notch (formed by the proximal articular surfaces of the radius and the ulna) is 24.5 mm. The humerus is only slightly smaller than the one of Sus scrofa from Petrálona Cave, Greece, which measures 53.9 and 54.2 mm, respectively (Tsoukala, 1989, p. 329).

ACCEPTED MANUSCRIPT Scarce material of Sus scrofa from the Megalopolis Basin (dental specimens not including an m3, a vertebra, and a tibia) has been already published by Melentis (1966d). This sample is morphologically and metrically consistent with an attribution to Sus scrofa, but it is not comparable to the Kyparíssia sample, due to the lack of

4.4. Hippopotamids Order: Artiodactyla Owen, 1848 Family: Hippopotamidae Gray, 1821

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Genus: Hippopotamus Linnaeus, 1758

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common elements.

Hippopotamus antiquus Desmarest, 1822

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Material: KYP1-797: left I1; KYP3-657: left I2; KYPT-842: right I2; KYP4-334: left C part; KYP-730: right C apex; KYP-324: left P4; KYP3-102: left i1; KYP3-46, KYP3-101: right c; KYP3-429: left c part; KYP-418: dentary fragment with p3–m1; KYP-685: right m2 part; KYP1-407: axis; KYP3-50, KYP3-616: cervical vertebra; KYP3-4, KYP3-44, KYP3-45, KYP3-614, KYP3-796: thoracic vertebra; KYP1-562,

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KYP1-564, KYP3-5: lumbar vertebra; KYP3-544: sacrum; KYP1-775, KYP1-776, KYP3-49, KYP3-326, KYPT-884: caudal vertebra; KYP-687: right scapula; KYP1767: left humerus; KYP1-470, KYP1-793: right humerus; KYP-636: proximal right radius part; KYP1-478: left proximal radius–ulna part; KYP1-773: right radius–ulna;

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KYP1-777, KYPT-801: right intermedium (lunar); KYPT-826: right ulnar; KYP-723: right accessory carpal (pisiform); KYP1-788, KYPT-813: left carpal III (capitatum);

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KYP1-788, KYPT-818: right carpal III (capitatum); KYP1-784, KYPT-808: left carpal IV (hamatum); KYP4-594: right carpal IV (hamatum); KYP1-779, KYPT-869: right metacarpal II; KYP3-454, KYPT-867: left metacarpal III; KYPT-815: right metacarpal III; KYP-525, KYPT-836: left metacarpal IV; KYP-165, KYP1-789, KYP3-545, KYP3-588: right metacarpal IV; KYP1-780, KYPT-825: right metacarpal V; KYP1-771, KYP3-456, KYP3-457: pelvis part; KYP1-766: left femur; KYP4-768: right femur; KYP1-791: distal part of right femur; KYP1-469, KYP4-644: femoral diaphysis; KYP1-772, KYPT-824: right patella; KYP4-314: right tibia; KYP1-559: left tibia; KYP1-795: proximal part of right tibia; KYP1-794, KYPT-838: right astragalus; KYP1-718, KYP4-332: right calcaneus; KYP1-786: right fourth tarsal (cuboid); KYP-424, KYP1-680: left metatarsal II; KYP-425: right metatarsal II;

ACCEPTED MANUSCRIPT KYPT-865: left metatarsal III; KYP1-790, KYPT-820: right metatarsal III; KYPT866: left metatarsal IV; KYP1-781: right metatarsal IV; KYPT-868: left metatarsal V; KYP1-774: right metatarsal V; KYP1-677, KYP1-778, KYP1-782, KYP1-783 KYP4596, KYPT-803, KYPT-822, KYPT-823, KYPT-837, KYPT-841: proximal phalanx; KYP1-785, KYP1-787, KYPT-806, KYPT-807, KYPT-809, KYPT-814, KYPT-827,

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KYPT-840, KYPT-871: middle phalanx; KYPT-811, KYPT-828, KYPT-839, KYPT864: distal phalanx.

4.4.1. Description

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Dentition — The dentition is mainly known from isolated and usually partly-

preserved elements. The available material consists of three upper incisors, two upper canines, an upper fourth premolar, a dentary fragment, bearing the p3, p4 and m1, as

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well as of a lower first incisor, three isolated lower canines and an m2 part. The upper first incisor KYP1-797 is cylindrical with very subtly curved crown. The specimen is broken both at its base and apically; a part of its mesially situated wear surface is, though, preserved. The surface is marked by longitudinal grooves, as well as by developmental cross lines. Preserved length = 104 mm, linguolabial diameter =

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32 mm, mesiodistal diameter = 29 mm.

An upper second incisor (I2, KYPT-842, Fig. 7A) is the only dental find from KYPT. It is rootless, 134 mm long, and rather robust. Its cross section is elliptical, the maximal diameter (31.7 mm) measured in the labiolingual direction, and the minimal

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(26.5 mm) in the mesiolateral direction. The cross section shape and diameters do not change significantly along the crown, but they are maximal in the middle of it. The

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tooth consists almost entirely of dentine; only the labial side is covered by a 20-mmwide enamel band. The dentine surface forms longitudinal grooves, particularly on the mesial and lateral sides. The enamel surface also follows this morphology, but it is wrinkled. The apex is worn mesiolingually through the occlusion with the corresponding lower incisor. The wear surface is slightly concave and bears numerous striations/scratch lines, which run almost vertically. The upper canine KYP4-334 is only apically preserved, where the crown is bevelled because of the occlusion with the lower canine. Except for the wear surface on the labial side of the tooth, the rest of the preserved crown part is covered with rugose enamel. Mesiolingually there is a deep groove directed from the base to the tip of the tooth. The maximal preserved width is 48 mm. Another upper canine apical part

ACCEPTED MANUSCRIPT (KYP-730) is slightly worn, still having a pointed apex and an incipient elliptical wear surface on its labial side. The lingual side is mesiodistally concave. The P4 is a single-cusped conical tooth, with triangular crown in occlusal view, because of the presence of a lingual accessory cusp. The main cusp is somewhat elongated mesiodistally, and it is only slightly worn. A weak cingulum is present at

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the mesial, lingual and distal sides of the crown-base. The roots are not preserved but it seems that they were three. The tooth’s maximal dimensions are 37.4 × 38.5 mm (length × width).

The lower incisors are cylindrical with grooved crowns and curve slightly lingually.

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One canine (KYP3-101, Fig. 7D) and the incisor KYP3-102 (Fig. 7C) were found

closely together, cropping out of the quarry section in the upper level of KYP3, and they could belong to the same individual. Another canine (KYP3-46, Fig. 7E) was

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recovered at the same findspot, but in the lower level. The incisor KYP3-102 is almost strait, preserved in a length of 303 mm. The enamel surface forms shallow longitudinal grooves, two of which are wider and deeper and run along the sides of the crown. The tip is worn laterally due to its contact to the corresponding upper incisor (I1). Its cross section is elliptical with maximal diameters of 62.5 mm

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(linguolabial) and 52.0 mm (mesiodistal).

The canine KYP3-101 (Fig. 7D) is preserved in its greatest part, lacking the mesial part of its tip and a short basal part. It is rootless, 635 mm long along its labial margin (390 mm long in straight line). In cross section it is triangular, the mesial side forming

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the base of the triangle. A shallow groove run along this side, and an even shallower along the distolingual side. The tip has a 183-mm-long wear facet on its lingual side,

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due to contact with the upper right canine. The maximal mesiodistal diameter is 52.1 mm, measured at about the middle of the crown, while the maximal linguolabial one is 71.0 mm, also at the middle of the tooth. The enamel is thin, with uneven surface, which exhibits developmental cross lines and marked longitudinal furrows (2 to 7 mm wide). The specimen KYP3-46 (Fig. 7E) is of similar morphology, though its longitudinal furrows are more prominent. It is less completely preserved (500 mm long along its labial margin) and its corresponding maximal cross-sectional diameters are 48.0 mm and 75.5 mm respectively. KYP3-429, an isolated find at KYP3, preserves only its basal part (250 mm long along its labial margin). Its cross-sectional diameters are 42.5 mm and 71.0 mm respectively.

ACCEPTED MANUSCRIPT The dentary fragment KYP-418 (Fig. 7B, B΄) belongs to a rather young individual, as indicated by the very slightly worn premolars. The two preserved premolars (p3 and p4) are conical, but rather asymmetrical, as their crown is slightly inclined rostrally. The p4 is also rotated, so that its mesiodistal axis is directed labiolingually. Both premolars exhibit strong stylids mesially, distally and distolingually of the main

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cuspid, as well as cinguli around the base of the crown, except for the mesiolingual side of p4. The cingulum of the latter tooth is extremely strong distally. The m1 has the characteristic double-trefoil occlusal pattern of the worn Hippopotamus molars.

The four conids are developed about equally. The talonid is stronger lingually. There

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is prominent mesial cingulum, as well as two weak ones at either side of the distal part of the crown.

A partially preserved isolated molar (KYP-685) is identified as a lower second molar,

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because of the absence of lingual cingulum, its elongate crown, and the absence of a talonid. The specimen (a surface find) is broken in a sagittal direction at the level of the lingual cuspids. Otherwise it is in pristine condition, as the cuspids are still unworn, except for the protoconid which shows a small mesiolabial wear facet. The enamel is rugose. In labial view the cuspids are conical and almost straight, curving

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slightly mesially. In occlusal view the teeth exhibit the typical trefoil-shaped crown morphology, which characterises the genus. Mesially and distally there are equallysized, wide and high cinguli; no cingulum is present, though, labially. A low stylid is observed labially, between the protoconid and the hypoconid. The cheek teeth

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dimensions are given in Table 1. The dental dimensions do not seem to be diagnostic at the species level, as the ranges of the European Pleistocene hippopotamuses, H.

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antiquus and H. amphibius, highly overlap to each other (Faure, 1985; Mazza, 1995). This is also seen in Fig. 8, where the m1 dimensions of KYP-418 are compared to published samples.

Axial skeleton — There is a great number of vertebrae and costae, which are attributed to Hippopotamus, but many of them are very fragmentary. The best preserved finds come from a fossil accumulation at KYP3 and they possibly belong to a single individual. Another group comes from the collapsed site KYP1, potentially also belonging to a single individual, as strongly indicated by their very similar size and the presence of an erosional lesion in their articular surfaces. The axis is known from a fragment of its cranial extremity that only preserves the robust odontoid process and

ACCEPTED MANUSCRIPT the right articular surface for the atlas. The latter is pear-shaped in lateral aspect, tapering towards the odontoid process; its profile is slightly convex, both in dorsoventral and craniocaudal directions. The other two cervical vertebrae belong to the 3rd to 6th part of the cervical series (Fig. 7H). They preserve their entire bodies and parts of their arcs and transverse processes. The cranial (of convex profile) and

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caudal (of concave profile) articulations are circular to elliptical, their dorsal margin tending to show a small median indentation. The ventral side of the body forms a

median sagittal ridge. The lateral processes all rather mediolaterally extended (almost equal to the body diameter). At their bases a circular transverse foramen is formed in

mediodorsally and lateroventrally respectively.

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each side. The cranial and caudal articular processes bear elliptical facets that face

The five available thoracic vertebrae are characterised by the presence of articular

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facets for the ribs at the laterodorsal margin of the bodies. The bodies are almost circular to heart-shaped in cranial and caudal view and have similar, flat cranial and caudal articulations. Lateroventrally they are concave. KYP3-614 (Fig. 7G), which belongs to the cranialmost part of the thoracic series, has very robust transverse processes that bear ventrally large concave articular facets for the corresponding rib

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head. The spinous process is triangular in cross section, forming a sharp edge cranially. Its caudal side is strongly concave in its basal part, but forms a sharp ridge towards its dorsal end. The more caudally situated thoracic vertebrae have progressively more strongly heart-shaped articulations, weaker transverse processes,

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much shorter and narrower spinous processes, and cranial articular processes bearing cylindrical facets. KYP3-44 is the last vertebra of the thoracic series, as it bears only

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cranial articular facets for the ribs. The cranialmost-situated lumbar vertebra KYP3-5 is very similar to the last thoracic ones, but lacks any articular facets for the ribs. The caudally-situated lumbar vertebrae KYP1-562 and KYP1-564 are characterised by flat and mediolaterally and craniocaudally expanded transverse processes (Fig. 7F). The articulations are much wider, but remain heart-like in shape. The sacrum KYP3-544 is formed by the complete fusion of four vertebrae (Fig. 7I, I΄). It is fairly well preserved, lacking a small part at the left side of its cranial end. The spinous processes of the sacral vertebrae are perfectly fused together, forming a dorsally-blunt crest. The same is true for the transverse processes that form a uniform lateral wing. This is greatly expanded laterally at the first sacral vertebra and forms

ACCEPTED MANUSCRIPT the articular surface for the ilium. At the meeting lines between the dorsal crest and the lateral wings open small circular sacral foramina, which perforate the bone in dorsoventral direction. The vertebral bodies are dorsoventrally compressed and wide, as in the lumbars. The ventral side of the sacrum is concave in both craniocaudal and mediolateral directions. The bodies of the fused vertebrae protrude ventrally,

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particularly towards the caudal end of the bone. The caudal vertebrae are cylindrical with short and narrow processes. The axial skeleton morphology does not differentiate from that of other known fossil samples (Reynolds, 1922; Kahlke, 1997).

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Thoracic limb — The thoracic limb is known from several more or less complete humeri, radiocubiti, and autopodial elements. The scapula is preserved only as

fragmentary specimens. The best preserved, KYP-687, has a craniocaudally directed,

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elliptical in outline glenoid cavity. Craniodorsally to the latter there is a very prominent supraglenoid tubercle. The diameters of the articulation are DAP = 97 mm, DT = 84 mm. The maximal DAP of the distal end, including the tubercle, is 159 mm. The humerus is known from three almost complete specimens (Fig. 7N). It is a massive bone with well-developed greater and lesser tubercles and deltoid tuberosity.

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The humeral head is very large, separated from the greater and lesser tubercles by grooves and from the humeral body (diaphysis) by a distinct neck. The diaphysis has an almost straight medial side. The distal end of the bone has a prominent medial and lateral epicondyles and a deep and sharply defined olecranon fossa. The radial fossa,

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in the bone’s cranial face, is more shallow. The humeral condyle (that is the distal articular level) is inclined laterally with regard to the bone’s shaft. It forms a wide

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trochlea medially, while its lateralmost part accommodates a much smaller, separate articular surface for the radius (capitulum). The Kyparíssia humeri are short compared to the H. antiquus samples described by Faure (1985) and Mazza (1995) (see also Fig. 10B), being similar in size to the smallest specimens of this species. Regarding other metrical parameters they are comparable to the minimum values of H. antiquus and the maximum of H. amphibius (Fig. 10A, B). Measurements are given in Table 2. The radius and ulna are known from several fragmentary finds, as well as from three better preserved, though not complete, specimens. KYP-636 preserves only the proximal part of the radius. The other two available specimens (Fig. 7L, M) come from the site KYP1 and may belong to the same individual, as indicated by their very similar dimensions. In Hippopotamus radius and ulna are fused together, forming a

ACCEPTED MANUSCRIPT single very robust bone with very wide proximal and distal ends. The olecranon is high and has a strong tuber. Caudally it forms a prominent keel. The trochlear notch of the ulna forms a continuous articular surface with the proximal articulation of the radius, the contact between the two bones being barely seen. The articular surface is divided into two unequal parts by a prominent ridge. The shafts of the radius and ulna

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are also fully fused to each other, leaving only a short mediolaterally directed foramen close to the radius proximal end. The distal end bears a wide articulation for the

proximal carpal row, which is formed by the fused distal articular surfaces of the ulna and the radius. There are three craniolaterally–mediocaudally directed concave

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surfaces, delimited by sharp crests. The medial and the lateral styloid processes are very prominent. Dimensionally (see Table 2) the Kyparíssia specimens are also

smaller than the average H. antiquus samples, but less so as in the humerus case. In

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general they are comparable to the minimum values of the H. antiquus samples, or fall in the overlapping ranges between H. antiquus and H. amphibius (Fig. 10C). The carpus is not completely known, as several of its elements lack from the locality fossil record. The intermediate (lunar) has a proximally-situated, craniocaudally elongate saddle-shaped articular surface for the radius, that in the specimen KYP1-

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777 articulates perfectly with the corresponding facet of the radio-cubitus KYP1-773. Distally it bears a large, almost rectangular in distal aspect, craniocaudally concave articular surface for the third and fourth carpals. This surface is undivided, the margin between the carpal II and IV facets being hardly visible. Two small facets for the

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ulnar (triquetrum) are located on the lateral side of the bone. The accessory carpal bone (pisiform) is craniocaudally elongate. In lateral aspect it

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curves slightly dorsally; in dorsal view it exhibits a marked bend medially. Its cranial end bears two adjacent articular faces for the ulnar, that form an obtuse angle between them. The third carpal (capitatum) is a craniocaudally elongate bone with a nonarticular caudally expanded part (Fig. 9A). Proximocranially there are two elongate articular facets for the radial and the intermedium carpals, while distally there is a triangular concave surface for the third metacarpal. Smaller facets on the medial and lateral sides correspond to the second and fourth carpal, respectively. Another small mediodistally-situated facet articulates with the second metacarpal. The fourth carpal (a.k.a. unciform, hamatum) looks similar to the third carpal, but it is much wider cranially (Fig. 9A). Two large, proximocranially-situated, well-separated facets articulate to the intermediate and the ulnar. The distal articulation consists of a large,

ACCEPTED MANUSCRIPT medially-situated facet for the fourth metacarpal and a smaller, craniocaudallyelongated one for the fifth metacarpal. The medial side bears a single or divided facet for the third carpal. Measurements of the carpals are given in Table 2. The metacarpals are robust bones with mediolaterally expanded bodies, which are slightly concave palmarly. They are similar in general shape, differing mainly in their

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proximal ends, which are described separately below. Their dorsal face is almost flat, but becomes rather convex towards the distal end of the bone. Their distal articular surfaces are trochlear in shape and cover the distal ends dorsally to palmarly. The

palmar part possesses sagittally-directed ridges for the articulation with the sesamoid

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bones. A certain degree of variation is observed in the articular surface shape and the robustness of metacarpal specimens among different individuals.

The second metacarpal (Fig. 9B) is slender with regard to the more axially situated

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metacarpals. Its corpus curves slightly medially and it is craniocaudally thicker along its lateral margin. The proximal end bears proximally a large subtriangular, mediolaterally concave, surface for the second carpal (trapezoid), and a dorsopalmarly elongate, convex one for the third carpal (capitatum). At the lateral side of the proximal end there are two adjacent small facets for the third metacarpal. The distal

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articular is asymmetrical, being much larger in diameter laterally. The third metacarpal (Fig. 9C) is rather slender and long, the longest in the series. The proximal articulation surface is subtriangular in proximal view and saddle shaped (mediolaterally concave and dorsopalmarly markedly convex). It articulates to the

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third carpal (capitatum). On the medial and lateral sides of the proximal end of the bone there are small articular facets for the adjacent metacarpals. The distal

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articulation is almost rectangular in distal aspect. The fourth metacarpal (Fig. 9D) is the best represented metapodial element in the Kyparíssia sample, as there are five well-preserved specimens available. Their proximal end accommodates a large, craniocaudally convex articular surface for the fourth carpal (hamatum). At the medial side of the proximal end there are two small facets for the articulation of the third metacarpal, while the lateral side has a small facet for the fifth metacarpal at its cranial end. The corpus is almost straight, flat dorsopalmarly and usually concave palmarly. The fifth metacarpal (Fig. 9E) is comparatively short and its body curves slightly laterally. The proximal articulation surface is subtriangular in proximal aspect and slopes caudally. Its dorsopalmar profile is undulate, while its mediolateral one is

ACCEPTED MANUSCRIPT concave, particularly dorsally. The corpus is robust. The distal articulation is strongly asymmetrical in distal aspect. Metrically (Table 2) the metacarpals are similar to the smallest specimens referred to H. antiquus or to the largest ones of H. amphibius (Faure, 1985; Mazza, 1995). In a graphical comparison (Figs 10D–H, 11A) the Kyparíssia specimens separate in most

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cases from those of the extant species, plotting in the area of H. antiquus. A peculiar metrical character of the studied sample is that the abaxial metacarpals (second and

fifth) are comparatively longer than the adaxial ones, as their length plots closer to the longest specimens referred to H. antiquus (Figs 10E, 11A). On the contrary, the

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adaxial ones are shorter, plotting among the smallest H. antiquus and the largest H. amphibius specimens (Fig. 10F–H).

The phalanges differ in morphology according to their anatomical position: The

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proximal ones are robust, with elliptical, concave, undivided proximal articulation, and a trochlear distal one, formed by a medial and a lateral condyle. The middle phalanges are very short, also differing from the proximal ones in their sagittally divided proximal articulation. The distal phalanges are small, having a shallow proximal articular surface. In general, the phalanges of the adaxial digits bear fairly

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symmetrical articular surfaces, while those of the abaxial digits are increasingly asymmetric towards the lateral and medial sides of the manus. However, in many cases their exact anatomical position may be quite uncertain.

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Pelvic limb — Several innominate bone specimens are attributable to Hippopotamus, all, however, are quite fragmentary and not very informative morphologically. Two of

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them, KYP1-771 and KYP3-456, have maximal acetabulum diameters of 94 mm and about 86 mm, respectively. The femur is known from two more or less complete specimens (KYP1-766, KYP4768) and three fragmentary ones, deriving from sites KYP1 and KYP4. It is a long bone with robust extremities (Fig. 7P). The proximal articulation (femoral head) is placed on a short but distinct neck. The greater trochanter reaches the same height to the head and is separated from it by a moderately deep notch. The trochanteric fossa in the caudal side of the proximal end is very deep, particularly at its lateral part. Distally to the fossa, close to the base of the neck on the caudal face of the bone, the lesser trochanter is found, having the form of a low and wide tubercle. The diaphysis is cylindrical, becoming almost rectangular in cross section towards its distal end. A

ACCEPTED MANUSCRIPT fairly deep supracondylar fossa opens in the caudolateral side of this part of the diaphysis. The distal extremity of the femur is massive. The articular condyles are very robust, particularly the medial one, separated by a very deep intercondylar fossa. Craniodistally there is a large asymmetrical trochlea for the articulation of the patella. Two specimens from KYP1 (KYP1-766 and KYP1-791) are very similar metrically

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and belong quite possibly to the same individual. Moreover, KYP1-766 articulates perfectly with the pelvis part KYP1-771, leading to the same conclusion. The

measurements of the available femoral specimens are given in Table 3 and are very

close to each other, although the two measurable specimens come from different sites.

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Compared to published H. antiquus and H. amphibius samples, the Kyparíssia

femoral specimens usually are metrically similar to the largest specimens of the extant species, falling within the metrical range of H. antiquus only in a few width

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parameters (Faure, 1985; Mazza, 1995). The Kyparíssia femora are particularly short when compared to H. antiquus finds from other European localities (Fig. 11B). Two right patellae (KYP1-772 and KYPT-824) are referred to this species. They are rhomboid in general shape with a medially projecting process, which creates a proximomedial and a distomedial concavity in the bone outline. The non-articular part

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of the bone is robust, particularly in its proximal part, where it projects well above the articular surface. The distal end is less stout and tapers distally to a pointed end. The articular surface is triangular, mediolaterally concave in its medial part and convex in its lateral part. Its lateral margin is slightly convex, almost straight, while its proximal

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and distal margins are sigmoid (convex laterally, concave medially). Dimensionally (Table 3) they are close to the minimal values of the Untermaßfeld, Germany

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(Kahlke, 1997, p. 336) and other European samples (Mazza, 1995, p. 179). KYP1-772 appears to be too short (its height is similar to that of specimens attributed to H. amphibius), but this might be due to high individual variation in this bone. The tibia is a short and robust bone, represented by two complete specimens, one from KYP1 (KYP1-559) and one from KYP4 (KYP4-314; Fig. 7O), as well as by partially preserved specimens (notably KYP1-795). Its proximal end is greatly expanded mediolaterally and craniocaudally with regard to the distal end, due to its very prominent condyles. The proximal articular surfaces are separated caudally by a deep notch. The body of the tibia is triangular in cross section, with almost flat or slightly convex medial side, and concave craniolateral and caudal sides. The tibial tuberosity is very strong. The distal end is only slightly larger than the body and it

ACCEPTED MANUSCRIPT accommodates the articular cochlea, which is rather shallow laterally. As in the case of the femur, the Kyparíssia tibiae are short, similar to some H. amphibius specimens (compare e.g., Faure, 1985), but they are wider than them quite like the smallest H. antiquus (Fig. 11B, C). Measurements are given in Table 3. The hippopotamus astragalus is a stout tarsal bone. There are two specimens in the

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Kyparíssia sample, one from KYP1 (Fig. 9H) and one from KYPT (Fig. 7K). They appear somewhat different, as the former is proportionally wider, and has less

pronounced distal sagittal keel, and more massive distal articulation, but otherwise

they share the same general morphology. The proximal articulation is wide and forms

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an open-notched trochlea with prominent sagittal ridges, which articulates to the distal end of the tibia. The trochlea is markedly asymmetrical, the lateral ridge being considerably higher and stronger that the medial. The medial ridge ends plantarly to a

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very prominent, rounded knob. The distal articulation consists of two craniocaudally convex, but mediolaterally concave, facets, for the articulation with the central (navicular) and the fourth (cuboid) tarsal bones, respectively. These facets form a prominent sagittal keel between them. The medial one is confluent with the plantar facet for the calcaneus. The latter occupies most of the plantar face of the astragalus;

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it is separated from the proximal trochlea by a deep groove that continues distomedially and terminates above the distal articular surface. In cranial (dorsal) view the proximal and the distal articular surfaces are clearly separated; a deep median depression is formed between them and extends sagittally to the proximal

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trochlea. The medial and lateral faces are dominated by the corresponding sides of the proximal trochlea. The lateral face also bears distally a small elliptical facet for the

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calcaneus. Measurements are given in Table 3. Compared to already published astragalus specimens, the Kyparíssia astragali are rather small, particularly the KYPT-838, which resides between the H. antiquus and the H. amphibius metrical ranges given by Faure (1985) for most parameters, except for the dorsoplantar ones (DAP), which fall within the H. antiquus range. KYPT-838 appears to have more H. amphibius-like proportions than KYP1-794, as the astragali of the latter species generally tend to be narrower with respect to their height than those of the former one (Faure, 1985; Mazza, 1995). The relative size of both specimens with regard to H. antiquus and H. amphibius samples is given in Fig. 11D, being comparable to the smallest H. antiquus and close to the H. amphibius range.

ACCEPTED MANUSCRIPT The calcaneus is rather elongate but strong (Fig. 7J). The calcaneal tuberosity is hardly differentiated from the body of the bone in lateral or medial aspects, but it is much wider when seen cranially or caudally. The proximalmost end of the tuberosity bears a wide and shallow, trochlea-like groove. The distal part of the calcaneus bears the articular facets for the fibula, the astragalus and the fourth tarsal bone (cuboid).

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The largest of the astragalus facets is found on the sustentaculum, extending medially from the body, facing distally and having subtriangular outline (it is wider caudally). Four smaller facets are found on the distally-situated coracoid process: two of them

situated on its medial side articulate with the astragalus, and the other two are found

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on the cranial and caudal side of the coracoid process’s distal end, articulating to the fibula and the fourth tarsal respectively. Both available calcaneal specimens are of

small dimensions (Table 3). KYP1-718 is juvenile, being more slender and lacking its

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unfused proximal end. KYP4-332 is smaller than any H. antiquus specimen measured by Mazza (1995) and Kahlke (1997), but it is very similar to a specimen referred by Mazza (1995) to H. tiberinus, a junior synonym of H. antiquus. Its height is also shorter than the minimum for this species given by Faure (1985). However, its DT and DAP dimensions fall well within the range of H. antiquus, being larger than the

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maximum of the extant species. In the scatter plot of Fig. 11E, KYP4-332 is placed between the H. antiquus and H. amphibius samples. The fourth tarsal (cuboid) is large and massive, with a high plantar side. The proximal face of the bone has a central strongly concave articular surface for the astragalus and

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a somewhat smaller dorsally-sloping elongate surface for the calcaneus. They are separated from each other by a prominent ridge. The medial side bears a circular

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articular facet for the central tarsal (navicular) at its plantar part, as well as a dorsoplantarly elongate one for the third tarsal bone (lateral cuneiform). On the distal side there are two articular surfaces, a circular one for the fourth metatarsal, and a smaller semicircular one for the fifth metatarsal. Metrically the single available specimen KYP1-786 is placed among the largest specimens of H. antiquus, according to the data given by Mazza (1995) and Kahlke (1997), while it is slightly larger than the dimensional ranges given by Faure (1985) for the same species. The metatarsals are similar to the metacarpals in general shape (Fig. 9F–H), but their proximal ends are more expanded volarly by means of a non-articular apophysis. The third and fourth ones are the longest and widest, with dorsoplantarly flattened bodies. The third has a triangular concave proximal articulation for the third tarsal (lateral

ACCEPTED MANUSCRIPT cuneiform). The same articulation in the fourth metatarsal is larger, circular, and articulates with the mediodorsal part of the fourth tarsal (cuboid). The second metatarsal (Fig. 9F) is much smaller, curves slightly medially, and has a very narrow proximal end that bears two articular facets for the first and second tarsals. The fifth metatarsal (Fig. 9G) is also small, with a body of subtriangular cross section, and

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semicircular proximal articulation for the fourth tarsal. All metatarsals bear small facets on the medial and/or lateral sides of their proximal ends for the articulation with their adjacent metatarsals. Measurements are given in Table 3. In metrical

comparisons the studied metatarsals appear to be rather large with respect to the

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metacarpals from Kyparíssia, placed well within the ranges of published H. antiquus samples (Fig. 11F–H), particularly those from KYP1. A certain degree of morphologic and metrical variation is evident (e.g., Fig. 9F). The KYPT specimens

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are somewhat smaller, but still usually larger than the largest H. amphibius (Fig. 9F– H).

The phalanges of the pes are similar morphologically to those of the manus, and in some cases are not clearly separable from them. As the studied specimens were found

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isolated, their exact anatomical position may be ambiguous.

As mentioned in Section 3, skeletal elements from the sites KYP1, KYP3 and KYP4 may belong to individual skeletons. This cannot be tested in all cases, as anatomically adjacent elements are not always available. A perfect articular facet match is observed

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at least between the intermediate carpal KYP1-777 and the radio-cubitus KYP1-773, as well as among the right-side pelvic-limb autopodial elements from KYP1 (KYP1-

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794, KYP1-786, KYP1-790, KYP1-781, KYP1-774) figured in Fig. 9H. The latter specimens, together with axial skeleton elements from the same site, also have in common the presence of lesions (exostoses and erosions), strongly indicating their belonging to a single skeleton. Similarly, the anatomically adjacent hippopotamus specimens that derive from the layer C of KYPT (Fig. 2) articulate very well to each other (e.g., Fig. 9A, G). However, there are also examples of clearly non-matching articular surfaces (e.g., the femur KYP1-766 and the tibia KYP1-559) implying the presence of more than one individual in the relevant site.

4.4.2. Discussion

ACCEPTED MANUSCRIPT The hippopotamus is one of the most common species in the fossil fauna of Kyparíssia. In the Megalopolis Basin it is also known from material excavated in 1902 (Melentis, 1966c), as well as from the very recent systematic excavation at Marathousa 1 (Konidaris et al., this issue). Hippopotamus was common in the western and southern parts of Europe during the Pleistocene, contracting and expanding its

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geographic range in response to the succession of glacial and interglacial climates, and eventually became extinct during the last glacial. The early very large-sized

forms, usually referred to as H. antiquus Desmarest, 1822, were replaced by smaller

ones during the Middle Pleistocene, eventually reaching a size comparable to that of

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the recent African species H. amphibius Linnaeus, 1758, and usually identified as

such. The fossil samples also exhibit a certain degree of morphologic and dimensional variability, leading some scholars to propose additional species, such as H. incognitus

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Faure, 1984 and H. tiberinus Mazza, 1991. On the other hand certain authors consider all European Pleistocene hippopotamuses as belonging to a single gradually evolving species, namely H. amphibius, in which they recognize two subspecies (H. amphibius antiquus and H. amphibius amphibius) (e.g., Kahlke, 1997; 2001). A two-species approach, H. antiquus – H. amphibius, is currently the most widely accepted,

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supported by morphological observations on craniomandibular material (Caloi et al., 1980). The other two species defined more recently on European samples have been considered as local forms placed well within the morphological and metrical variation of H. antiquus and H. amphibius (Petronio, 1995). Since the species-level taxonomy

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of European hippopotamuses is still debated, we provisionally utilise the two-species (antiquus–amphibius) taxonomic scheme in the present study, following Petronio

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(1995).

Due to the lack of craniomandibular specimens in the studied material, which are the only skeletal elements that offer taxonomically diagnostic characters (Caloi et al., 1980), the specific determination has to be based on biometry alone. As it is evident from the biometrical comparison of certain skeletal elements (Figs 10, 11), the Kyparíssia hippopotamus is a middle-sized one, dimensionally close to the smallest individuals referred to H. antiquus in most metrical parameters. It is actually quite similar in size to most samples referred by Mazza (1995) to “H. tiberinus” (Figs 10, 11) (currently considered as a junior synonym of H. antiquus). The metrical similarity to the smallest H. antiquus specimens is systematic, pertaining to most bones. This applies to the stylopodial bones (humerus, femur) and tibia, which are particularly

ACCEPTED MANUSCRIPT short. On the contrary, the abaxial metacarpals (second and fifth), as well as the available metatarsals from KYP1, are comparatively long (Figs 10, 11). Despite its intermediate body size falling in the tricky area between the two Pleistocene species, the hippopotamus sample from Kyparíssia as a whole fits better to the species H. antiquus, particularly in the transverse dimensions, indicating more robust elements

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than those of the extant species. Its smaller dimensions with regard to other samples of H. antiquus might be evolutionary, as the Middle Pleistocene hippos tend to be smaller than the Early Pleistocene ones, or related to the local climatic conditions

during the specific time period when these animals were living, as the hippopotamus

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body-size variations have been potentially linked to climatic changes (Mazza and

Bertini, 2013). Compared to the already published material from the Megalopolis basin (Melentis, 1966c; Konidaris et al., this issue) the studied sample from

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Kyparíssia is very similar dimensionally, except for the single postcranial specimen from Marathousa 1, which is larger (Fig. 10F, H; Fig. 11A, G).

5. Biochronology

The insufficient stratigraphic data of the studied material, and the presence of at least

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two fossiliferous horizons may be considered problematic for the biostratigraphy of the studied sites in Kyparíssia mine. However, given the rather fast sedimentation rate of 21 cm/kyr calculated by van Vugt et al. (2000) for the Megalopolis Basin, which was corroborated recently by Tourloukis et al. (this issue), and the stratigraphic

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distance of less than 15 m between the lowest and the top fossiliferous horizons, a chronostratigraphic range of about 70 kyr can be estimated for the Kyparíssia fauna,

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which is not expected to be important biostratigraphically in the Middle Pleistocene. Turtles and birds are of minimal importance for determining the age of the locality, as they have not changed morphologically since the Pleistocene. The extant boar Sus scrofa appears in the fossil record before the beginning of the Middle Pleistocene, possibly even earlier than the Jaramillo subchron, as indicated by closely related forms (Sus gr. scrofa; Martínez-Navarro et al., 2015). The stratigraphic range of Hippopotamus antiquus is usually considered as Early – early Middle Pleistocene, its first appearance possibly dated at about 2.1 Ma (Bellucci et al., 2014; Pandolfi and Petronio, 2015; Pandolfi et al., 2015). Early finds from the Peloponnesian locality of Elis, about 80 km NW of Megalopolis, may be of similar age or even somewhat older (Thenius, 1955; Reimann and Strauch, 2008). The upper biostratigraphic limit of the

ACCEPTED MANUSCRIPT species is also not well defined, as, under the two-species concept followed here, it was replaced during the 700–400 ka interval by the extant species H. amphibius, at least in Western Europe. Marra et al. (2014) and Pandolfi and Petronio (2015) are more precise on this limit regarding the Italian faunas, and place the last occurrence of H. antiquus in MIS15 (at about 600 ka). The biochronology of the genus in SE

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Europe is, however, not well known. The presence of a large-sized hippo at Marathousa 1, a site provisionally dated to MIS12 or close to the MIS12–MIS11

transition (between ~0.48 and ~0.42 Ma) (Jacobs et al., this issue; Konidaris et al.,

this issue; Tourloukis et al., this issue) shows at least that H. antiquus has survived till

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more recent times in this region. Mazza and Bertini (2013) observe a marked decrease in the body size of European hippopotami at the beginning of the Middle Pleistocene, that led to populations referred to as “H. tiberinus” (according to Mazza, 1995) or

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“H. ex gr. H. antiquus” (according to Mazza and Bertini, 2013). If this observation (which is based on rather few well-dated fossil samples) is correct, it would mean that the age of the Kyparíssia fauna is constrained to the early Middle Pleistocene. However, the occasional presence of large hippopotami in similar aged localities, including the adjacent Marathousa 1 mentioned above, shows that this hypothesis

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needs further investigation. The available data on the stratigraphy, biometry and taxonomy of the European (and Greek in particular) hippopotamus samples are still inadequate for a detailed reconstruction of the genus’ morphological and metrical variation through space and time.

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In conclusion, the presence of Sus scrofa and Hippopotamus antiquus in the studied material indicates a latest Early to early Middle Pleistocene age for Kyparíssia (about

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1.0 to 0.4 Ma). This rather wide timeframe is somewhat constrained by the study of the rest of the taxa included in the Kyparíssia fauna (Athanassiou, this issue). The complete faunal list (Table 4) points to an age between 0.8 and 0.4 Ma.

6. Palaeoecology

Contrary to their uncertain biochronologic indications the studied samples offer sufficient clues on the palaeoenvironment of the locality. Considering the two chelonian species identified at Kyparíssia, it is safe to assume that they inhabited similar ecological niches as their extant representatives. Therefore we can infer the presence of ponds and rivers because of the occurrence of Emys orbicularis. Testudo marginata is, though, flexible ecologically, as it can inhabit forested, woodland, as

ACCEPTED MANUSCRIPT well as open areas (Ernst and Barbour, 1989). The same conclusion on the presence of a freshwater body is drawn by the avifauna, which consists exclusively of water-birds. The majority of the identified avian taxa are adapted to a wide range of climatic zones. The mute swan (Cygnus olor) is currently mostly found in freshwater marshes and lakes and feeds mainly on vegetative parts of aquatic plants, but will also

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consume aquatic invertebrates and small amphibians (Carboneras, 1992). Similarly, the common teal (Anas crecca), prefers freshwater environments, with emergent

vegetation and feeds primarily on seeds of aquatic plants and grasses but also on

aquatic invertebrates (Carboneras, 1992). The mallard (Anas platyrhynchos) lives off

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rich shallow wetlands with emergent vegetation and has a highly opportunistic diet consuming aquatic and terrestrial plants, invertebrates, amphibians and fish

(Carboneras, 1992). The gadwall (Mareca strepera) is found in shallow wetlands and

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feeds mostly on seeds, leaves, stem and roots of aquatic plants (Carboneras, 1992). The shoveler (Spatula clypeata) is found in a variety of freshwater wetlands, and consumes mostly small sized invertebrates and occasionally plants (Carboneras, 1992). The garganey (Spatula querquedula) is found mostly on swamps and small lakes and feeds mostly on aquatic invertebrates, but will also consume small fish,

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amphibians and aquatic plants (Carboneras, 1992). The common coot (Fulica atra) prefers slow moving bodies of water, is primarily vegetarian but will also consume molluscs, insects and even small fish and amphibians (Taylor, 1996). The darter (Anhinga sp.) lives on freshwater lakes, swamps and slow moving rivers, requires

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some tall vegetation such as trees for preening and sunning, and feeds mainly on fish, amphibians and aquatic invertebrates (Orta, 1992). The darter is not presently found

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in Greece, having been restricted to warmer regions, but went extinct relatively recently in Turkey (Orta, 1992). It is also known from Marathousa 1 (Michailidis et al., this issue) and was part of the warm-adapted fauna of the Megalopolis palaeolake. The great cormorant (Phalacrocorax carbo) is found on lake and sea shores and consumes mostly fish but will also consume crustaceans and amphibians (Orta, 1992). It has a very wide range of climatic zone tolerance, from the arctic to the tropical, and is often found alongside darters (Orta, 1992). The majority of birds identified from Kyparíssia are adapted to living by shallow freshwater bodies, with emergent vegetation and at least some isolated trees. The lake system, with its richly vegetated shores would have been large enough to support populations of fish, amphibians and several aquatic invertebrates. The character of the

ACCEPTED MANUSCRIPT avifauna described here, is identical to the one identified from Marathousa 1 (Michailidis et al, this issue), revealing a similar ecosystem during the Middle Pleistocene, recorded on both localities. The mammalian taxa identified in this study can also give some clues to the reconstruction of the local palaeoenvironment. Sus scrofa can be found today in quite

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diverse habitats with respect to temperature and humidity, but it is a predominantly forest species, favouring temperate deciduous or mixed forests, woodlands and thickets (Faure and Guérin, 1984; Guérin and Faure, 1997), particularly in the

European ecological context. It is also very versatile in its diet, being an omnivore

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that can feed on roots, tubers, fruits, nuts, seeds, but also on small animals and

carrions (Guérin and Faure, 1997; Powell, 2004), that are usually more plentiful in a richly vegetated environment. The abundant representation of Hippopotamus in the

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fossil sample infers the presence of a large body of water. Hippopotamus is dependant on the presence of fresh water throughout the year, thus indicating also a warm– temperate climate with prevailing temperatures well above the water freezing point. Moreover, the subsistence of H. antiquus almost exclusively on aquatic plants, as evidenced by stable isotope analyses of a Spanish sample (Palmqvist et al., 2003),

population.

7. Conclusions

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indicates that the freshwater-covered area was quite expanded, in order to sustain a

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The lignite mine of Kyparíssia in the northern part of the Megalopolis Basin has yielded a diverse fossil fauna collected during field surveys and small-scale

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excavations over the years 2004–2008 and 2012. The fauna is dominated by mammals, of which Hippopotamus and Sus are described here, but turtles and birds are also present and rich in species diversity. The chelonian assemblage includes a terrestrial and an aquatic (freshwater) species, while the recovered bird specimens all belong to aquatic taxa, implying the presence of a water body as a dominant environmental feature in the region. The strong presence of Hippopotamus is another piece of evidence towards the same conclusion. As a whole, the studied taxa point to a predominately forested environment around an expanded perennial lake with rich aquatic and waterside vegetation. The occurrence of certain large-sized herbivores in the fauna, such as a horse and a giant deer, also indicate the existence of more open areas in close distance to the palaeolake (Athanassiou, this issue). This reconstruction

ACCEPTED MANUSCRIPT of richly vegetated surroundings with wide open areas is in accordance with the published data on the palaeoenvironment of the Megalopolis Basin (Mädler, 1971; Sickenberg, 1976; Schütt et al., 1985; Nickel et al., 1996; see also Field et al., this issue, and Konidaris et al., this issue). The identified mammalian taxa are consistent with a Middle Pleistocene (Galerian)

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age of the locality, as already concluded in previous and ongoing studies (Sickenberg, 1976; van Vugt et al., 2000; Konidaris et al., this issue). The small body size of

Hippopotamus antiquus from Kyparíssia may, though, indicate an age closer to the

vaguely known last appearance of this species, possibly constraining it to the 0.7–0.4

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Ma range. The currently collected faunal data do not allow any biostratigraphic distinction of the studied sites within the Kyparíssia mine according to their

taxonomic content. However, the age difference among the sites is not expected to be

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significant.

According to the currently available data the Kyparíssia fauna appears to be similar to the samples yielded at other findspots of the Megalopolis basin, located within the Marathousa member, including the recently excavated Marathousa 1 (Konidaris et al., this issue). The detailed study of the stratigraphy and the taphonomy of the locality, as

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well as the collection of better samples, remain to be carried out, if a large-scale systematic excavation could be organised within the Kyparíssia mine in the near future. The sites KYP3 and KYP4 are still there and are considered promising for

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future research.

Acknowledgements

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The authors wish to thank Dr. G. Lyras, D. Bouzas, Dr. V. Giannopoulos, Dr. D. Minou, as well as many graduate and postgraduate students who participated to the Kyparíssia fieldwork. The Public Power Corporation (∆ΕΗ) offered valuable technical help during the fieldwork in the mine. Dr. A. Gamauf (Natural History Museum of Vienna) allowed use of their comparative collections. This research was financially supported in part by the Hellenic Ministry of Culture, Ephorate of Palaeoanthropology–Speleology. The excavation at Kyparíssia T site was financed by the ERC STG no. 283503 (‘PaGE’) awarded to K. Harvati. The authors also thank the two anonymous reviewers, whose constructive comments and corrections greatly improved the original manuscript.

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Athanassiou, A., 2016. Pleistocene vertebrates from the Kyparíssia lignite mine, Megalópolis, S. Greece. Closing Symposium, ERC Starting Grant Project “PaGE”, Paleoanthropology at the Gates of Europe, Tübingen, Germany, Abstracts, 14–15. Athanassiou, A., this issue. Pleistocene vertebrates from the Kyparíssia lignite mine, Megalopolis Basin, S. Greece: Rodentia, Carnivora, Proboscidea, Perissodactyla, Ruminantia. Quat. Intern. this issue. Athanassiou, A., Tourloukis, V., Thompson, N., Lychounas, A., Panagopoulou, E., Harvati, K., 2016. Hippopotamus (Artiodactyla, Mammalia) and other vertebrate remains from the Kyparíssia-T site, Megalópolis, S. Greece. Closing Symposium, ERC Starting Grant Project “PaGE”, Paleoanthropology at the Gates of Europe, Tübingen, Germany, Abstracts, 12–13. Baumel, J.J., King, A.S., Breazile, J.E., Evans, H.E., Vaden Berge, J.C. (Eds.), 1993. Handbook of Avian Anatomy: Nomina Anatomica Avium. Second Edition. Publications of the Nuttall Ornithological Club, No. 23. Harvard University, Cambridge, Massachusetts. Bochenski, Z.B., Tomek, T., 1997. Preservation of bird bones: Erosion versus digestion by owls. Int. J. Osteoarchaeol. 7, 372–387. Bürchner, L., 1903. Wichtige Funde fossiler Knochen in Arkadien. Ber. Naturwiss. Ver. Regensburg 9, 119–123. Behrensmeyer, A.K., 1978. Taphonomic and ecologic information from bone weathering. Paleobiol. 4, 150–162. Bellucci, L., Bona, F., Corrado, P., Magri, D., Mazzini, I., Parenti, F., Scardia, G., Sardella, R., 2014. Evidence of late Gelasian dispersal of African fauna at Coste San Giacomo (Anagni Basin, central Italy): Early Pleistocene environments and the background of early human occupation in Europe. Quat. Sci. Rev. 96, 72–85. Caloi, L., Palombo, M.R., Petronio, C., 1980. Resti cranici di Hippopotamus antiquus (= H. major) e Hippopotamus amphibius conservati nel Museo di Paleontologia dell’Università di Roma. Geol. Rom. 19, 91–119. Carboneras, C., 1992. Family Anatidae (ducks, geese and swans). In: del Hoyo, J., Elliott, A., Sargatal, J. (Eds.), Handbook of the Birds of the World, vol. 1. Lynx Edicions, Bacelona, pp. 528–628. Del Hoyo, J., Elliott, A., Sargatal, J. (Eds.), 1992. Handbook of the Birds of the World, vol. 1. Lynx Edicions, Barcelona. Ernst, C.H., Barbour, R.W., 1989. Turtles of the World. Smithsonian Institution Press, Washington DC. Faure, M., 1985. Les hippopotames quaternaires non-insulaires d’Europe occidentale. Nouv. Arch. Mus. Hist. Nat. Lyon 23, 13–79. Faure, M., Guérin, C., 1983. Le Sus scrofa (Mammalia, Artiodactyla, Suidae) du gisement pléistocène supérieur de Jaurens à Nespouls, Corrèze, France. Nouv. Arch. Mus. Hist. Nat. Lyon 21, 45–63. Faure, M., Guérin, C., 1984. Sus strozzii et Sus scrofa, deux mammifères artiodactyles, marquers des paléoenvironments. Palaeogeogr., Palaeoclim., Palaeoecol. 48, 215–228. Field, M.H., Ntinou, M., Tsartsidou, G., van Bergehenegowuen, D., Tourloukis, V., Thompson, N., Karkanas, P., Panagopoulou, E., Harvati, K., this issue. A

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palaeoenvironmental reconstruction (based on palaeobotanical data and diatoms) of the Middle Pleistocene elephant (Palaeoloxodon antiquus) butchery site at Marathousa, Megalopolis, Greece. Quat. Intern. this issue. Galobart, À., Ros, X., Maroto, J., Vila, B., 2003. Descripción del material de hipopótamo (Hippopotamus antiquus Desmarest, 1822) de los yacimientos del Pleistoceno inferior de Incarcal (Girona, NE de la Península Ibérica). Paleont. i Evol. 34, 153–173. Guérin, C., Faure, M., 1997. The wild boar (Sus scrofa priscus) from the postVillafranchian Lower Pleistocene of Untermaßfeld. In: Kahlke, R.D. (Ed.), Das Pleistozän von Untermaßfeld bei Meiningen (Thüringen), Teil 1. Monographien des Römisch-Germanischen Zentralmuseums Mainz. Habelt, Mainz, pp. 375–383. Hünermann, K.A., 1975. Sus scrofa Linné aus dem Pleistozän von WeimarEhringsdorf. Abh. Zentr. Geol. Inst., Paläontol. Abh. 23, 251–263. Hünermann, K.A., 1977. Sus scrofa L. aus dem Jungpleistozän von Taubach bei Weimar in Thüringen. Quartärpaläont. 2, 225–235. Hünermann, K.A., 1978. Das Wildschwein (Sus scrofa L.) aus dem Jungpleistozän von Burgtonna in Thüringen. Quartärpaläont. 3, 123–128. I.C.V.G.A.N., 2005. Nomina Anatomica Veterinaria. International Committee on Veterinary Gross Anatomical Nomenclature, Hannover. Jacobs, Z., Li, B., Karkanas, P., Tourloukis, V., Thompson, N., Panagopoulou, E., Harvati, K., this issue. Dating of Middle Pleistocene Marathousa 1 (Greece) lacustrine sediment using multiple-aliquot pre-dose multi-elevatedtemperature post-infrared infrared stimulated luminescence (MET-pIRIR). Quat. Intern. Kahlke, R.D., 1997. Die Hippopotamus-Reste aus dem Unterpleistozän von Untermaßfeld. In: Kahlke, R.D. (Ed.), Das Pleistozän von Untermaßfeld bei Meiningen (Thüringen), Teil 1. Monographien des Römisch-Germanischen Zentralmuseums Mainz. Habelt, Mainz, pp. 277–374. Kahlke, R.D., 2001. Schädelreste von Hippopotamus aus dem Unterpleistozän von Untermaßfeld. In: Kahlke, R.D. (Ed.), Das Pleistozän von Untermaßfeld bei Meiningen (Thüringen), Teil 2. Monographien des Römisch-Germanischen Zentralmuseums Mainz. Habelt, Mainz, pp. 483–500. Konidaris, G.E., Athanassiou, A., Tourloukis, V., Thompson, N., Giusti, D., Panagopoulou, E., Harvati, K., this issue. 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. Quat. Intern., DOI:10.1016/j.quaint.2017.12.001. König, H.E., Liebich, H.G. (Eds.), 2004. Veterinary Anatomy of Domestic Mammals. Schattauer, Stuttgart. Löhnert, E., Nowak, H., 1965. Die Braunkohlenlagerstätte von Khoremi im Becken von Megalopolis/Peloponnes. Geol. Jb. 82, 847–867. Mädler, K., 1971. Die Früchte und Samen aus der frühpleistozänen Braunkohle von Megalopolis in Griechenland und ihre ökologische Bedeutung. Geol. Jb. Beiheft 110, 1–79. Marra, F., Pandolfi, L., Petronio, C., Di Stefano, G., Gaeta, M., Salari, L., 2014. Reassessing the sedimentary deposits and vertebrate assemblages from Ponte Galeria area (Rome, central Italy): An archive for the Middle Pleistocene faunas of Europe. Earth Sci. Rev. 139, 104–122.

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Martínez-Navarro, B., Madurell-Malapeira, J., Ros-Montoya, S., Patrocinio Espigares, M., Medin, T., Hortolà, P., Palmqvist, P., 2015. The Epivillafranchian and the arrival of pigs into Europe. Quat. Intern. 389, 131– 138. Mazza, P., 1995. New evidence on the Pleistocene hippopotamuses of Western Europe. Geol. Rom. 31, 61–241. Mazza, P.P.A., Bertini, A., 2013. Were Pleistocene hippopotamuses exposed to climate-driven body size changes? Boreas 42, 194–209. Melentis, J.K., 1961. Die Dentition der pleistozänen Proboscidier des Beckens von Megalopolis im Peloponnes (Griechenland). Ann. Géol. Pays Hellén. 12, 153– 262. Melentis, J.K., 1963. Die Osteologie der pleistozänen Proboscidier des Beckens von Megalopolis im Peloponnes (Griechenland). Ann. Géol. Pays Hellén. 14, 1– 107. Melentis, J.K., 1966a. Die pleistozänen Cerviden des Beckens von Megalopolis im Peloponnes (Griechenland). Ann. Géol. Pays Hellén. 16, 1–92. Melentis, J.K., 1966b. Die pleistozänen Nashörner des Beckens von Megalopolis in Peloponnes (Griechenland). Ann. Géol. Pays Hellén. 16, 363–402. Melentis, J.K., 1966c. Über Hippopotamus antiquus Desmarest aus dem Mittelpleistozän des Beckens von Megalopolis in Peloponnes (Griechenland). Ann. Géol. Pays Hellén. 16, 403–435. Melentis, J.K., 1966d. Sus scrofa L. aus dem Jungpleistozän des Beckens von Megalopolis in Peloponnes (Griechenland). Ann. Géol. Pays Hellén. 16, 436– 445. Melentis, J.K., 1966e. Die Boviden des Jungpleistozäns des Beckens von Megalopolis im Peloponnes (Griechenland). Ann. Géol. Pays Hellén. 16, 446–472. Melentis, J.K., 1966f. Über Equus abeli aus dem Mittelpleistozän des Beckens von Megalopolis im Peloponnes (Griechenland). Ann. Géol. Pays Hellén. 17, 158– 168. Melentis, J.K., 1966g. Clemmys caspica aus dem Pleistozän des Beckens von Megalopolis im Peloponnes (Griechenland). Ann. Géol. Pays Hellén. 17, 169– 181. Michailidis, D., Konidaris, G.E., Athanassiou, A., Panagopoulou, E., Harvati, K., this issue. The ornithological remains from Marathousa 1 (Middle Pleistocene; Megalopolis Basin, Greece). Quat. Int. this issue. Mitzopoulos, H., Orphanides, T.G., von Heldreich, T., 1862. [General report on the accessions in the collections of the Physiographic Museum in Athens during the academic year 1860–1861]. Athens. (in Greek) Nickel, B., Riegel, W., Schönherr, T., Velitzelos, E., 1996. Environments of coal formation in the Pleistocene lignite at Megalopolis, Peloponnesus (Greece) — reconstructions from palynological and petrological investigations. N. Jb. Geol. Pal. Abh. 200, 201–220. Okuda, M., van Vugt, N., Nakagawa, T., Ikeya, M., Hayashida, A., Yasuda, Y., Setoguchi, T., 2002. Palynological evidence for the astronomical origin of lignite–detritus sequence in the Middle Pleistocene Marathousa Member, Megalopolis, SW Greece. Earth Planet. Sci. Let. 201, 143–157. Orta, J., 1992. Family Anhingidae (darters). In: del Hoyo, J., Elliott, A., Sargatal, J. (Eds.), Handbook of the Birds of the World, vol. 1. Lynx Edicions, Barcelona, pp. 354–361.

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Palmqvist, P., Gröcke, D.R., Arribas, A., Farina, R.A., 2003. Paleoecological reconstruction of a lower Pleistocene large mammal community using biogeochemical (δ13C, δ15N, δ18O, Sr:Zn) and ecomorphological approaches. Paleobiol. 29, 205–229. Panagopoulou, E., Tourloukis, V., Thompson, N., Athanassiou, A., Tsartsidou, G., Konidaris, G.E., Giusti, D., Karkanas, P., Harvati, K., 2015. Marathousa 1: a new Middle Pleistocene archaeological site from Greece. Antiquity 343 Project Gallery, http://antiquity.ac.uk/projgall/panagopoulou343. Pandolfi, L., Petronio, C., 2015. A brief review of the occurrences of Pleistocene Hippopotamus (Mammalia, Hippopotamidae) in Italy. Geol. Croat. 68, 313– 319. Pandolfi, L., Grossi, F., Frezza, V., 2015. New insight into the Pleistocene deposits of Monte delle Piche, Rome, and remarks on the biochronology of Hippopotamus (Mammalia, Hippopotamidae) and Stephanorhinus etruscus (Mammalia, Rhinocerotidae) in Italy. Estud. Geol. 71, e026. Papadopoulos, P., Lüttig, G., Vinken, R., 1997. Geological map of Greece 1 : 50,000, Megalopolis sheet. Institute of Geology and Mineral Exploration, Athens. Petronio, C., 1995. Note on the taxonomy of Pleistocene hippopotamuses. Ibex 3, 53– 55. Powell, D.M., 2004. Pigs (Suidae). In: Hutchins, M., Kleiman, D.G., Geist, V., McDade, M.C. (Eds.) Grzimek’s Animal Life Encyclopedia, 2nd Edition, Gale, Farmington Hills, MI, pp. 275–290. Reimann, C.K., Strauch, F., 2008. Ein Hippopotamus-Schädel aus dem Pliozän von Elis (Peloponnes, Griechenland). N. Jb. Geol. Pal. Abh. 249, 203–222. Reynolds, S.H., 1922. A monograph on the British Pleistocene Mammalia – vol. III, part I: Hippopotamus. Palæontographical Society, London. Schütt, H., Velitzelos, E., Kaouras, G., 1985. Die Quartärmollusken von Megalopolis (Griechenland). N. Jb. Geol. Pal. Abh. 170, 183–204. Sickenberg, O., 1976. Eine Säugetierfauna des tieferen Bihariums aus dem Becken von Megalopolis (Peloponnes, Griechenland). Ann. Géol. Pays Hellén. 27, 25–73. Skuphos, T.G., 1905. Über die palæontologischen Ausgrabungen in Griechenland in Beziehung auf das Vorhandensein des Menschen. Comptes Rendus du Congrès International d’Archéologie, Athènes, pp. 231–236. Taylor, P.B., 1996. Family Rallidae (rails, gallinules and coots). In: del Hoyo, J., Elliott, A., Sargatal, J. (Eds.), Handbook of the Birds of the World, vol. 3. Lynx Edicions, Barcelona, pp. 108–209. Thenius, E., 1955. Hippopotamus aus dem Astien von Elis (Peloponnes). Ann. Géol. Pays Hellén. 6, 206–212. Theodorou, G., 2014. Megalopolis – 112 years after the first excavation by National and Kapodistrian University of Athens (NKUA) and the post-lignite era. VIth International Conference on Mammoths and their Relatives, Grevena–Siatista (2014). Sci. Ann., Sch. Geol., Aristotle Univ. Thessaloniki, 102, 195–196. Thompson, N., Tourloukis, V., Panagopoulou, E., Harvati, K., this issue. In search of Pleistocene remains at the Gates of Europe: directed surface survey of the Megalopolis Basin (Greece). Quat. Intern., DOI:10.1016/j.quaint.2018.03.036. Tourloukis, V., Muttoni, G., Karkanas, P., Monesi, E., Scardia, G., Panagopoulou, E., Harvati, K., this issue. Magnetostratigraphic and chronostratigraphic constraints on the Marathousa 1 Lower Palaeolithic site and the Middle

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Pleistocene deposits of the Megalopolis basin, Greece. Quat. Intern., DOI:10.1016/j.quaint.2018.03.043. Tsiftsis, E.V., 1987. Geology and Hydrogeology of the Megalopolis Basin, Peloponnese, Greece. PhD, University of Bristol, Department of Geology, Bristol. Tsoukala, E., 1989. [Contribution to the study of the Pleistocene fauna of large mammals (Carnivora, Perissodactyla, Artiodactyla) from Petralona Cave, Chalkidiki (N. Greece)]. PhD thesis, Aristotle University of Thessaloniki, Thessaloniki. (in Greek) Tsoukala, E., Chatzopoulou, K., 2005. A new Early Pleistocene (latest Villafranchian) site with mammals in Kalamotó (Mygdonia Basin, Macedonia, Greece) – preliminary report. Mitt. Komm. Quartärforsch. Österr. Akad. Wiss. 14, 213– 233. Van Vugt, N., de Bruijn, H., van Kolfschoten, T., Langereis, C.G., 2000. Magnetoand cyclostratigraphy and mammal-fauna’s of the Pleistocene lacustrine Megalopolis Basin, Peleponnesos, Greece. Geol. Ultraject. 189, 69–92. Vinken, R., 1965. Stratigraphie und Tektonik des Beckens von Megalopolis (Peloponnes, Griechenland). Geol. Jb. 83, 97–148. Vlachos, E., Delfino, M., 2016. Food for thought: Sub-fossil and fossil chelonian remains from Franchthi Cave and Megalopolis confirm a glacial refuge for Emys orbicularis in Peloponnesus (S. Greece). Quat. Sci. Rev. 150, 158–171.

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Fig. 1. Map of Greece, indicating the geographic location of the Megalopolis Basin (ellipse) in Arcadia, central Peloponnesus. The oblique satellite image ‘A’ shows the distribution of lignite mines within the basin (1: Chorémi, 2: Marathoúsa, 3: Thoknía, 4: Kyparíssia) as well as the position of the Kyparíssia fossiliferous outcrops (asterisk). The positions of the main fossiliferous sites within the Kyparíssia mine, situated north of the homonymous village, are given as red dots on the satellite image ‘B’. Relief map source: Wikimedia Commons. Satellite images source: Google. Note the different orientation of the satellite image ‘B’. (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.)

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Fig. 2. Stratigraphy of the site Kyparissia T (KYPT). A: Stratigraphic column of the exposed section. The bone drawings indicate the presence and the relative abundance of fossils. B: Photograph of the section showing the main accumulation of fossils. Scale bar (in the centre of the photograph): 20 cm. C: Layer geometry over a section

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length of 8 m in S–N direction showing the horizontal distribution of fossil

specimens. The numbers indicate Hippopotamus specimens and refer to their

specimen numbers (omitting the KYPT prefix). Chelonian, avian and cervid skeletal fragments are indicated by triangles, lozenges and circles, respectively. The

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highlighted area of layer C corresponds to the main accumulation of fossils (see also

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Fig. 3).

Fig. 3. The main fossil accumulation from KYPT, partially prepared and still in a plaster jacket, as seen from below, exhibiting a high degree of syndepositional fragmentation. The specimen numbers of certain visible finds are given (omitting the

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KYPT prefix).

Fig. 4. Chelonian specimens from Kyparissia. Emys orbicularis (Linnaeus, 1758), blue in the online version of the figure. Carapace: A, KYPT-833: nuchal; B, KYPT-

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856: partial neural; C, KYPT-834: suprapygal; D, KYP3-37, partial costal of a juvenile; E, KYP-10: partial costal; F, KYP-360: peripheral; G. KYP3-656: peripheral; Plastron: H, KYP-512: left epiplastron; I, KYP3-66: right hyoplastron

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fragment; J, KYP1-663: left hypoplastron; K, KYP-361: partial right xiphiplastron; L, KYP3-39, partial right xiphiplastron. Testudo marginata Schoepff, 1793, brown in the online version of the figure. Carapace: M, KYP-744: nuchal; N, KYP-78: partial costal; O, KYP-753: partial anterior peripheral; P, KYP1-673: bridge peripheral; Q, KYP3-87: 7th right peripheral; R, KYP-748: two articulated posterior peripherals, probably the 8th and 9th from the right side; Plastron: S, KYP-513: left xiphiplastron. Carapace: 1, dorsal; 2, drawing of the dorsal; 3, visceral views. Plastron: 1, visceral; 2, ventral; 3, drawing of the ventral views. Scale bar equals 5 cm. (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.)

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Fig. 5. Avian specimens from Kyparíssia. A: distal left humerus of Cygnus olor, KYP3-621, caudal aspect; B: distal right ulna of Cygnus olor, KYP3-621, ventral aspect; C: clavicula of Cygnus olor, KYP3-621, caudal aspect; D: complete left humerus of Anhinga sp., KYPT-804, caudal aspect left, cranial aspect right; E:

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complete right humerus of Fulica atra, KYP-516, caudal aspect left, cranial aspect right; F: distal left humerus of Spatula clypeata, KYPT-819, cranial aspect; G:

complete left tarsometatarsus of Phalacrocorax carbo, KYPT-812, dorsal aspect left, plantar aspect right; H: cervical vertebra of Cygnus olor, KYP3-660; I: distal left

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tibiotarsus of Cygnus olor, KYP3-621, cranial aspect; J: proximal right humerus of

Cygnus olor, KYP3-621, caudal aspect; K: proximal right carpometacarpus of Anas

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platyrhynchos, KYP4-338, ventral aspect; L: complete left coracoideum of Spatula querquedula, KYP-750, dorsal aspect; M: distal right coracoideum of Mareca strepera, KYP-509, dorsal aspect; N: distal left femur of Anas crecca, KYP-51, caudal aspect; O: distal right ulna of Anas platyrhynchos, KYP-511, dorsal aspect left,

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ventral aspect right. Scale bar equals 5 cm.

Fig. 6. Suid specimens from Kyparíssia referred to Sus scrofa Linnaeus, 1758. A: right proximal radius and ulna (KYP3-658); B: left distal humerus (KYP-667); C: left dentary fragment with the m3 (KYP3-81); D: part of a right lower canine (KYP-700),

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lingual view. Scale bar equals 5 cm.

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Fig. 7. Dental and osteological material from Kyparíssia referred to Hippopotamus antiquus Desmarest, 1822. A: right I2 (KYPT-842), mesial view; B: dentary fragment with p3–m1 (KYP-418), lingual view; B΄: the same specimen in occlusal view; C: left i1 (KYP3-102); D: right c (KYP3-101); E: right c (KYP3-46); F: fourth(?) lumbar vertebra (KYP1-564), caudal view; G: thoracic vertebra (KYP3-614), cranial view; H: cervical vertebra (KYP3-616), cranial view; I: sacrum (KYP3-544), ventral view; I΄: the same specimen in dorsal view; J: right calcaneus (KYP4-332), medial view; K: astragalus (KYPT-838), cranial view; L: right radius–ulna (KYP1-773), cranial view; M: proximal part of a left radius–ulna (KYP1-478), cranial view; N: left humerus

ACCEPTED MANUSCRIPT (KYP1-767), caudal view; O: right tibia (KYP4-314), cranial view; P: right femur (KYP4-768), cranial view. Scale bar equals 10 cm.

Fig. 8. Metrical comparison of the Hippopotamus lower first molar (m1, KYP-418)

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from Kyparíssia (black dot) with published H. antiquus (lozenges) and H. amphibius (triangles) samples (data from Mazza, 1995). The green and blue lines represent the metrical ranges of H. antiquus and H. amphibius respectively, according to Faure

(1985). Measurements in mm. (For interpretation of the references to colour in this

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figure legend, the reader is referred to the Web version of this article.)

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Fig. 9. Osteological (autopodial) material from Kyparíssia referred to Hippopotamus antiquus Desmarest, 1822. A: left carpals III (capitatum) and IV (hamatum) (KYPT808, KYPT-813, respectively), in proximal view, shown articulated to each other; B: right second metacarpal (KYPT-869), in dorsal view; C: right third metacarpal (KYPT-815), in dorsal view; D: left fourth metacarpal (KYPT-836), in dorsal view; E: right fifth metacarpal (KYPT-825), in dorsal view; F: one right and two left second

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metatarsals (KYP-425, KYP-424 and KYP1-680, respectively, from left to right), in dorsal view (KYP1-680 is broken proximally); G: left metatarsals III–V (KYPT-865, KYPT-866, KYPT-868, respectively, from left to right) that quite probably belong to the same individual, in dorsal view at their inferred anatomical positions; H: right

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autopodial elements that quite probably belong to the same individual: astragalus (KYP1-794), fourth tarsal (KYP1-786), third metatarsal (KYP1-790), fourth

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metatarsal (KYP1-781), fifth metatarsal (KYP1-774), proximal phalanges (KYP1-782 and KYP1-783), middle phalanx (KYP1-787), in dorsal view at their inferred anatomical positions. Scale bar equals 10 cm.

Fig. 10. Metrical comparison of Hippopotamus thoracic limb elements from Kyparíssia (black dots) with published H. antiquus (lozenges) and H. amphibius (triangles) samples (data according to Melentis, 1966c; Mazza, 1995; Kahlke, 1997; Galobart et al., 2003; Tsoukala and Chatzopoulou, 2005; Konidaris et al., this issue). H. antiquus lozenge colour varies according to locality, where known (grey: Untermaßfeld, Germany; green: Incarcal, Spain; orange: Megalopolis, Greece; blue:

ACCEPTED MANUSCRIPT Kalamotó 2, Greece; red: Marathousa 1, Greece), while yellow indicates the specimens attributed by Mazza (1995) to H. tiberinus, which is considered here as a junior synonym of H. antiquus, following Petronio (1995). The green and blue lines represent the metrical ranges of H. antiquus and H. amphibius respectively, according to Faure (1985). Measurements in mm. (For interpretation of the references to colour

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in this figure legend, the reader is referred to the Web version of this article.)

Fig. 11. Metrical comparison of Hippopotamus thoracic and pelvic limb elements from Kyparíssia with published H. antiquus and H. amphibius samples (data

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according to Melentis, 1966c; Mazza, 1995; Kahlke, 1997; Galobart et al., 2003;

Tsoukala and Chatzopoulou, 2005). H. antiquus lozenge colour varies according to

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locality, where known (grey: Untermaßfeld, Germany; green: Incarcal, Spain; orange: Megalopolis, Greece), while yellow indicates the specimens attributed by Mazza (1995) to H. tiberinus, which is considered here as a junior synonym of H. antiquus, following Petronio (1995). The green and blue lines represent the metrical ranges of H. antiquus and H. amphibius respectively, according to Faure (1985). Measurements in mm. (For interpretation of the references to colour in this figure legend, the reader

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is referred to the Web version of this article.)

ACCEPTED MANUSCRIPT Table 1. Lower cheek teeth measurements (in mm) of Hippopotamus antiquus from Kyparíssia.

L p3 41.5 —

W p3 26.5 —

L p4 35.0 —

W p4 31.0 —

L m1 51.0 —

W m1 33.5 —

L m2 — 65.2

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lower dentition KYP-418 KYP1-685

H m2 — > 56

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Table 2. Thoracic limb measurements (in mm) of Hippopotamus antiquus from Kyparíssia.

L med — 415 413

DAPp — > 178 184

DTp — — 155

DAPm 69 76 71

DTm 67 60 69

radius KYP-636 KYP1-478 KYP1-773

L — — 300.0

L med — — 288.0

L lat — — 265.0

DAPp — — 73.0

DAPpa 70.0 66.0 68.0

DTp — — 107.0

ulnar KYPT-826

H 59.5

DAP 48.0

DT 81.0

intermedium KYP1-777 KYPT-801

H 78.5 68.0

DAP 75.0 68.0

DT 82.0 73.0

carpal III KYP1-788 KYPT-813 KYPT-818

H 61.0 56.5 56.1

DAP 99.0 88.0 86.0

DT 84.5 52.9 52.7

carpal IV KYP1-784 KYP4-594 KYPT-808

H 63.0 52.0 56.5

DAP 89.0

DT 84.0 86.0 77.5

MC II KYP1-779 KYPT-869

L 142.0 134.0

DAPp 50.0 43.0

DTd 149 149 151

DTda 107 103 —

DTpa 113.0 107.0 105.5

DAPd — — 91.0

DAPda — — 90.0

DTda — — 92.0

DTm 38.0 40.3

DAPd 41.0 40.0

DTd 48.0 44.1

DAPda 41.0 38.6

M AN U

TE D EP

AC C

74.5

DAPd 141 144 142

RI PT

L — > 448 > 450

SC

humerus KYP1-470 KYP1-767 KYP1-793

DTp 44.0 38.5

DAPpa 42.0

DTpa 41.0 38.2

DAPm 22.0 27.3

DTda 44.0 44.0

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DAPp 55.0 56 —

DTp 58.4 57.6 56.6

DAPpa 54.5 55 —

DTpa 58.4 57.6 56.6

DAPm 23.0 26 27

MC IV KYP1-789 KYP3-545 KYPT-836 KYP-165 KYP-525 KYP3-588

L 153.5 147.0 143.0 139.6 137.4 —

DAPp 56.7 58.0 54.0 — 56.0 —

DTp 61.3 61.0 56.0 57.0 58.0 60.2

DAPpa — — 54.0 — — —

DTpa 61.3 61.0 56.0 57.0 58.0 60.2

DAPm 25.5 28.0 29.0 26.2 25.5 25.7

MC V KYP1-780 KYPT-825

L 118.5 113

DAPp 46.0 49.8

DTp 48.0 42

DAPpa 45.0 47

DTpa 43.0 41.5

M AN U

TE D EP AC C

DTm 43.7 46 46

DAPm 25.8 32

DAPd 43.0 — —

RI PT

L 156.0 161 161

DTd 50.3 — —

DAPda 43.0 43.5 42.8

DTda (48.5) 54.8 54.5

DTm 51.0 51.0 46.0 41.5 16.5 47.0

DAPd 44.2 — — 43.0 44.6 —

DTd 57.5 — — 48.0 55.7 —

DAPda 44.2 49.5 43.0 43.0 44.2 —

DTda 52.7 56.0 52.2 (46) 52.0 —

DTm 41.5 42

DAPd 42.0 —

DTd 48.0 —

DAPda 42.0 41

DTda 43.0 45

SC

MC III KYP3-454 KYPT-815 KYPT-867

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Table 3. Pelvic limb measurements (in mm) of Hippopotamus antiquus from Kyparíssia.

DTp 195.0 192.0

Caput DAP 98.0 89.0

Caput DT 96.0 90.0

DAPm 68.0 70.5

DAPd 199.0 —

DTd 170.0 —

DTda 168.0 —

patella KYP1-772 KYPT-824

H 111.0 127.0

Ha 89.0 88.0

DAP 72.0 66.0

DT 112.5 109.0

DTa 101.0 92.0

tibia KYP1-559 KYP4-314 KYP1-795

L 365.0 340.0 —

DAPp 137.0 139.0 137.0

DTp 164.0 173.0 167.0

DTm 69.0 63.0 —

DAPd 82.5 73.5 —

DTd 115.0 110.0 —

astragalus KYP1-794 KYPT-838

H 118.0 104.0

H med 93.5 88.0

H lat 107.5 92.0

DAP med 77.0 72.4

DAP lat 75.7 62.0

DTp 96.5 88.0

DTpa 86.0 84.0

DAPda 58.5 52.0

DTda 95.0 88.0

calcaneus KYP4-332

H 198.0

H med 149.0

DAPmax 82.0

DTmax 88.0

DTp (tuber) 71.0

DTmin 37.0

tarsal IV KYP1-786

H 90.0

DAP 81.0

DT 90.0

H dorsal 51.5

MT II KYP1-680 KYP-425

L > 105 90.9

DAPp (44) 40.5

DTp (36) 28.8

DAPpa — —

DTpa — —

DAPm 24.1 26.5

DTm 34.7 28.4

DAPd (42) 37.8

DTd (51.5) 39.8

DAPda (40) (35.5)

DTda (46) 36.4

MT III KYP1-790 KYPT-865 KYPT-820

L 144 138 —

DAPp 61 61 —

DTp 51.5 42 49

DAPpa 44 39 40

DTpa 51.5 42

DAPm 23 23 —

DTm 46 45 —

DAPd 42 — —

DTd 54.5 — —

DAPda 42 41.5 —

DTda 50.5 49.8 —

MT IV KYP1-781

L 148

DAPp 76

DTp 57

DAPpa 55

DTpa 57

DAPm 24.5

DTm 48.3

DAPd 43

DTd 55

DAPda 43

DTda 51

M AN U

TE D

DAPp (tuber) 70.0

EP

AC C

DTm 71.0 72.0

RI PT

L 529.0 535.0

SC

femur KYP4-768 KYP1-766

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142

—

51

—

49

26

45

—

54

42.5

50.5

MT V KYP1-774 KYPT-868

L 106.5 99.3

DAPp 50.5 45.5

DTp 33 32.5

DAPpa 39 41.5

DTpa 35 30

DAPm 24 28

DTm 30 29

DAPd 43 —

DTd 39 39

DAPda 40.5 39

DTda 39.5 37

AC C

EP

TE D

M AN U

SC

RI PT

KYPT-866

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Table 4

Comprehensive faunal list of Kyparíssia with taxon representation in each site (KYP1–KYP4, KYPT), as well as in the unstratified material (KYP). Combined data from Athanassiou (this issue) and the present study. Note that ´Cervus´ peloponnesiacus is distinguishable from Cervidae indet. only in the unstratified material (KYP), which includes antlers. Family

Genus and species

Testudines

Emydidae

Anseriformes

KYP

KYP1

Emys orbicularis

+

+

+

Testudinidae

Testudo marginata

+

+

+

Anatidae

Anas crecca

+

Anas platyrhynchos

+

Cygnus olor

Spatula clypeata

KYP4

KYPT +

+

+

+

+

+

Rallidae

Fulica atra

+

Suliformes

Anhingidae

Anhinga sp.

+

Phalacrocoracidae

Phalacrocorax carbo

+

Rodentia

Castoridae

Castor fiber

+

Carnivora

Canidae

Canis sp.

+

Hyaenidae

Hyaenidae indet.

Felidae

Panthera sp.

+ +

M AN U

Gruiformes

TE D

Spatula querquedula

KYP3

SC

Mareca strepera

KYP2

RI PT

Order

+

+

Felis sp.

Proboscidea

Elephantidae

Perissodactyla

Rhinocerotidae

Suidae

Stephanorhinus sp.

+

Equus sp.

+

+

+

Hippopotamidae

Hippopotamus antiquus

+

+

Cervidae

Praemegaceros verticornis

+

+

Cervus elaphus

+

Dama sp.

+

´Cervus´ peloponnesiacus

+

Cervidae indet.

+

Bison sp.

+

Bovidae

+

+

+

Sus scrofa

AC C

Artiodactyla

+

EP

Equidae

Palaeoloxodon antiquus

+ +

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

AC C

EP

TE D

M AN U

SC

RI PT

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60

210

humerus 190

50

DTd

DTda

170 45

150

130

35

A 110 115

125

135

145

155

165

30 110

175

115

120

125

130

M AN U

DAPd

SC

40

135

E

140

145

150

155

L

70

210

metacarpal III

humerus

65

190

60

DTd

DTp

170

150

55 50

TE D

45

130

B

110 340

390

440

490

L lat

40 135

F 140

145

150

155

160

165

170

175

180

185

540

L

65

metacarpal III

EP

radius 140 130

110 100 90 80 230

250

AC C

120

270

290

310

60

55 DTda

150

50

45

40

C 330

G 35 135

350

140

145

150

155

L

160

165

170

175

180

185

L

55

80

metacarpal II

75

50

metacarpal IV

70 65 DTp

45

DTp

DTp

RI PT

metacarpal II 55

60 55

40 50 45

35

D 30 110

115

120

125

130

135 L

140

145

150

H

40

155

35 120

130

140

150 L

160

170

70

130

metacarpal IV 60

110

55

100

50

90

45

80

40

70

A

35 120

130

140

150

160

60 150

170

170

190

210

M AN U

L 250

E

230

250

270

H

60

femur

metatarsal III

230

55

210

DTda

50

190

45

170

TE D

40

B

150 480

530

580

630

680

L 130

35 115

730

F 120

125

130

135

140

145

150

155

L

60

tibia 120

100

90

80 300

350

AC C

110

400

55

50 DTda

EP

metatarsal IV

DTd

45

40

C

450

G 35 115

500

120

125

130

135

L

140

145

150

155

160

L

130

50

astragalus

metatarsal V

120

45

110

40 DTda

DTda

SC

DT

120

RI PT

calcaneus

65

DTp

DTda

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100

35 90

30

80

D

H

70

25 70

80

90

100 H med

110

120

130

80

85

90

95

100 L

105

110

115

AC C

EP

TE D

M AN U

SC

RI PT

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AC C

EP

TE D

M AN U

SC

RI PT

ACCEPTED MANUSCRIPT

AC C

EP

TE D

M AN U

SC

RI PT

ACCEPTED MANUSCRIPT

AC C

EP

TE D

M AN U

SC

RI PT

ACCEPTED MANUSCRIPT

AC C

EP

TE D

M AN U

SC

RI PT

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AC C

EP

TE D

M AN U

SC

RI PT

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ACCEPTED MANUSCRIPT

45

RI PT

m1

35

30

45

50

55

M AN U

L

TE D

40

EP

35

SC

25

AC C

W

40

AC C

EP

TE D

M AN U

SC

RI PT

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