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A Villafranchian Hipparion-Bearing Mammal Fauna from Sésklo (E. Thessaly, Greece): Implications for the Question of Hipparion–Equus Sympatry in Europe

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A Villafranchian Hipparion-Bearing Mammal Fauna from Sésklo (E. Thessaly, Greece): Implications for the Question of Hipparion–Equus Sympatry in Europe quaternary Article A Villafranchian Hipparion-Bearing Mammal Fauna from Sésklo (E. Thessaly, Greece): Implications for the Question of Hipparion–Equus Sympatry in Europe Athanassios Athanassiou 1,2 1 Ephorate of Palaeoanthropology–Speleology, Ministry of Culture, Ardittou 34B, 11636 Athens, Greece; [email protected] 2 Museum of Palaeontology and Geology, National and Kapodistrian University of Athens, 15784 Athens, Greece Academic Editors: Maria Rita Palombo and Juan Manuel López García Received: 31 May 2018; Accepted: 2 August 2018; Published: 7 August 2018 Abstract: Recently collected fossil material in the Villafranchian locality of Sésklo, as well as a re-evaluation of a pre-existing, partly-published museum collection, allow the recognition of a lower faunal level in the locality, older than the main Equus-dominated fossil assemblage, dated in the Early Pleistocene (MNQ17). The lower level yielded, instead, an advanced hipparion, referred to the species Plesiohipparion cf. shanxiense, and a small number of associated taxa: an ostrich (Struthio cf. chersonensis), an unidentified proboscidean, the pig Sus arvernensis, two antelopes (Gazella cf. bouvrainae and Gazellospira torticornis), a large bovid (Bovini indet.), and a rhinoceros (Stephanorhinus sp.). The lower-level fauna is dated in the latest Pliocene (MN16) and indicates a rather open and dry palaeoenvironment. The faunal sequence in Sésklo shows that the hipparion did not co-occur with the stenonid horse, at least in this region. Previous reports on sympatry of these taxa may result from faunal mixing, requiring re-examination of the available samples. Keywords: Plesiohipparion; Late Pliocene; Villafranchian; Greece; biostratigraphy; equid sympatry 1. Introduction The fossiliferous locality of Sésklo is located 8 km west of the city of Vólos, the capital of the Prefecture of Magnesia (Thessaly, Greece) and 2 km north-east of the homonymous village (Figure1a). The area is used as a clay pit operated by the cement industrial company “Hercules”, during the quarrying works of which, vertebrate fossils come occasionally to light. The first fossil finds known from the locality were unearthed in 1971 [1,2]. Thenceforth several fossil collections or excavations were carried out in the pit (Figure2) producing a rich fossil vertebrate collection of Villafranchian age. The Sésklo fauna is well known from several papers published during the last 40 years. It is a diverse fauna, comprising twenty-two mammalian taxa [2–10]. Most of the finds belong to the horse Equus stenonis Cocchi, 1867, but the family Bovidae constitutes the most diverse group within the assemblage. The fauna is biochronologically dated in the lower MN17 zone, as defined by Mein [11] (MNQ17 according to Guérin [12], or Saint-Vallier Faunal Unit, according to Italian authors, e.g., [13]), currently placed in the early part of the Early Pleistocene. Previous publications on the Sésklo fauna already mentioned the presence of fossils in at least four different findspots (A, B, G and D in Figure2) within the quarry, deriving from different fieldwork expeditions [2,3,5]. Unfortunately, all material excavated or collected until 1982 was lumped together, forming a mixed sample that contained all specimens deriving from the locality, without any findspot indication. Quaternary 2018, 1, 12; doi:10.3390/quat1020012 www.mdpi.com/journal/quaternary Quaternary 2018, 1, 12 2 of 24 Quaternary 2018, 2, x FOR PEER REVIEW 2 of 24 Figure 1. (a) Map of Greece, indicating the geographic location of the Sésklo locality (asterisk), west of Figure 1. (a) Map of Greece, indicating the geographic location of the Sésklo locality (asterisk), west of the city of Vólos. Locality coordinates: 39.368° N, 22.851° E (WGS84 datum). Contour interval: 200 m. the city of Vólos. Locality coordinates: 39.368◦ N, 22.851◦ E (WGS84 datum). Contour interval: 200 m. (b) Panoramic view of the clay pit seen from the North. (b) Panoramic view of the clay pit seen from the North. Differences in specimen fossilisation and morphology have shown, however, beyond doubt that Differencesa small part in of specimen the material fossilisation derives and from morphology a different have level. shown, Athanassiou however, beyond[5] excluded doubt thatthese a specimenssmall part offrom the the material comprehensive derives from study a different of the Sé level.sklo Athanassioufauna because [5] of excluded their different these specimens physical characters,from the comprehensive as well as for study the reason of the Sthatésklo most fauna of becausethem seemed of their to different belong to physical a large-sized characters, hipparion, as well notas for to thea stenonid reason that horse. most Nonetheless, of them seemed most to of belong them were to a large-sized publishedhipparion, at about the not same to a stenonidtime, together horse. withNonetheless, selected mostmaterial of them from were the main published fossil ataccumulation about the same within time, the together quarry with(findspots selected Β, materialor Δ, or both;from theFigure main 2) fossilas a mixed accumulation sample [3]. within Their the dark quarry colour (findspots hinted to B, the or D similarly, or both; coloured Figure2) clays as a mixed at the basesample of the [3]. basin Their sedimentary dark colour hinted sequence to the (Figures similarly 2b colouredand 3) as clays the most at the probable base of the stratigraphic basin sedimentary level of provenance.sequence (Figures Indeed,2b andone3 of) as the the initially most probable recorded stratigraphic findspots (Γ level, see ofFigure provenance. 2) must Indeed,have been one situated of the ininitially the dark recorded clays. findspotsMuch later, (G, in see the Figure summer2) must of have2005, beenduring situated the ongoing in the dark works clays. of the Much quarry, later, a in new the findspotsummer ofwas 2005, discovered, during the situated ongoing in worksthe lower of the hori quarry,zons of a newthe red findspot clayey was layers discovered, that dominate situated the in basinthe lower fill. horizonsThe site was of the destroyed red clayey by layers the quarry that dominate machinery, the but basin a few fill. Thefragmentary site was destroyedspecimens by were the quarrydelivered machinery, to the present but a fewauthor fragmentary by the quarry specimens supervisor were deliveredA. Christophorides, to the present who author precisely by the mapped quarry theirsupervisor location. A. Christophorides,This findspot was who dubbed precisely SE2 (Figure mapped 2).their The preparation location. This of findspotthe fossil was material dubbed during SE2 (Figurethe following2). The years preparation showed, of thequite fossil surprisingly, material during that SE2 the also following differs years markedly showed, from quite the surprisingly, main fossil accumulationthat SE2 also because differs markedly it exclusively from yielded the main remains fossil attributable accumulation to a becausehipparionine it exclusively horse, instead yielded of Equusremains. This attributable discovery to acorroborated hipparionine our horse, previous instead assumption of Equus. Thisabout discovery the provenance corroborated of ourthe previousdark-coloured assumption sample, about as both the samples provenance contain of thefoss dark-colouredil remains of a sample, very large-sized as both samples hipparion. contain The presentfossil remains paper ofis aa verycomprehensive large-sized description hipparion. Theof these present two paper samples, is a comprehensive which must have description been very of closethese twostratigraphically samples, which and must chronologically, have been very an closed quite stratigraphically probably represent and chronologically, the same andfauna. quite A probablypreliminary represent version the of same the present fauna. A paper preliminary was pr versionesented of in the the present 15th paperCongress was of presented the Regional in the 15thCommittee Congress on ofMediterranean the Regional CommitteeNeogene Stratigraphy on Mediterranean (RCMNS Neogene 2017) Stratigraphy[14]. (RCMNS 2017) [14]. Quaternary 2018, 1, 12 3 of 24 Quaternary 2018, 2, x FOR PEER REVIEW 3 of 24 Quaternary 2018, 2, x FOR PEER REVIEW 3 of 24 Figure 2. (a) Distribution of the fossiliferous sites/findspots within the Sésklo clay pit. The year of Figure 2. (a) Distribution of the fossiliferous sites/findspots within the Sésklo clay pit. The year of Figurediscovery 2. (ofa) Distributioneach one is indicatedof the fossiliferous in parenthese sites/findspotss. Graphical within scale: the 500 Sésklo m. Satellite clay pit. image The yearsource: of discovery of each one is indicated in parentheses. Graphical scale: 500 m. Satellite image source: discoveryGoogle. (b )of Stratigraphic each one is columnindicated of inthe parenthese Sésklo Basins. Graphicalsedimentary scale: fill, 500based m. onSatellite Müller image [15] and source: own Google. (b) Stratigraphic column of the Sésklo Basin sedimentary fill, based on Müller [15] and own Google.data. The (b position) Stratigraphic of the fossiliferouscolumn of the sites Sésklo is approximate. Basin sedimentary Graphical fill, scale: based 5 onm. Müller [15] and own data. The position of the fossiliferous sites is approximate. Graphical scale: 5 m. data. The position of the fossiliferous sites is approximate. Graphical scale: 5 m. Figure 3. The lower part of the Sésklo Basin sedimentary fill, showing the transition from the lower Figure 3. The lower part of the Sésklo Basin sedimentary fill, showing
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