Athanassiou, 2012 - M. Trogontherii Peloponnese

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Quaternary International 255 (2012) 9e28

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Quaternary International

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A skeleton of Mammuthus trogontherii (Proboscidea, Elephantidae) from NWPeloponnese, Greece

Athanassios AthanassiouHellenic Ministry of Culture, Ephorate of PalaeoanthropologyeSpeleology, Ardittou 34B, 11636 Athens, Greece

a r t i c l e i n f o

Article history:Available online 2 April 2011

E-mail address: [email protected].

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a b s t r a c t

Fossil elephant remains were identified in Loussiká, NW Peloponnese, Southern Greece, when tuskfragments were recognized in a bulldozer backfill. An excavation carried out in 2001 and 2003 by theHellenic Ministry of Culture revealed the partial skeleton of an adult male mammoth, referred to theMiddle Pleistocene species Mammuthus trogontherii. The recovered material includes part of the skull,the complete mandible, several vertebrae and ribs, both scapulae, ulna, tibia, most carpal and tarsalbones, metapodials and phalanges. The metrical and anatomical study of the skeleton shows that theliving individual was about 45 years old, stood about 3.80 m high and weighed about 8 t. M. trogontheriiis a very rare species in Southern Europe. The Loussiká skeleton represents the first solid evidence of thespecies’ presence in Southern Greece and considerably expands to the south its palaeobiogeographicrange in the Balkan area.

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1. Introduction

Elephant fossils are common in palaeontological sites, as theirrobust bones and their massive teeth have relatively good preserva-tion potential in the fossil record, even in relatively higher energyenvironments. This resulted in a fairly good fossil record, recoveredfrom numerous localities, that contributed to the current under-standing of evolution and phylogenywithin the family Elephantidae.

In mainland Greece, the recorded fossil elephant-bearing local-ities are close to forty (Doukas and Athanassiou, 2003), commonlysituated in fluvial and lacustrine basins. The straight-tuskedelephant Elephas antiquus is the dominant species, present in morethan twenty sites dated to theMiddleeLate Pleistocene (Doukas andAthanassiou, 2003; Tsoukala et al., 2010). Older, Lower Pleistocenelocalities have yielded numerous remains of the primitivemammoth Mammuthus meridionalis, often in association with thegomphothere Anancus arvernensis. More advanced mammoth findsare, however, very rare and they are geographically restricted to thenorthern part of Greece. The insular proboscidean faunas, knownfrom more than thirty sites, are characterised by the presence ofdwarf endemic forms, derived mainly from continental E. antiquus.

The new site ‘Loussiká’, described in the present paper, is situ-ated in western Achaia, NW Peloponnese (Fig. 1). It was discoveredin 1988 by the archaeologist Andreas Darlas, while prospecting the

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area around the village of Loussiká for prehistoric artefacts. There,in the southern slope of the Serdiní valley, he noticed the presenceof tusk and bone fragments in a bulldozer backfill. The bulldozer,while levelling the ground for agricultural use, had cut across theproximal part of the skull, destroying the sole preserved right tusk,as well as an unknown number of bones. The site was excavatedseveral years later, in 2001 and 2003, by the Ministry of Culture(Ephorate of PalaeoanthropologyeSpeleology, Athens, and ST’Ephorate of Prehistoric and Classical Antiquities, Patras) under thedirection of Dr. A. Darlas, with the participation of the presentauthor. The excavation revealed a partial proboscidean skeletonthat included the skull, the mandible, and part of the axial andappendicular skeleton. The recovered specimens were prepared inthe laboratory of the Ephorate of PalaeoanthropologyeSpeleologyin Athens and are currently stored in the collections of theArchaeological Museum of Patras. The skull still remains in Athensbecause of its fragility and demanding preparation, and it will bealso transported to Patras when the preparation and consolidationare finished.

The elephant skeleton of Loussiká is referred here to the MiddlePleistocene ‘steppe mammoth’ Mammuthus trogontherii (Pohlig,1885). However, thefindingwas preliminarily referred to E. antiquusFalconer and Cautley, 1847 (Doukas and Athanassiou, 2003;Athanassiou, 2010), a quite common Pleistocene elephant species,mainly because of a misinterpretation of the skull morphologyduring the excavation (see Section 7). After the plaster jacketremoval, the preliminary preparation and re-examination of the

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Fig. 1. Topographical map of W. Achaia (SW Greece) showing the geographic position of the new locality ‘Loussiká’ (asterisk; 38� 50 53.800 N, 21� 350 32.100 E, altitude 45 m, WGS84datum) south of the homonymous village. Contour interval: 100 m. Graphical scale: 3 km.

A. Athanassiou / Quaternary International 255 (2012) 9e2810

skull and dental remains, it became evident that the Loussiká skel-eton belongs toM. trogontherii, a very rare species in Greece, knownuntil now only from scanty dental material.

M. trogontherii is usually considered as an intermediate form inthe Mammuthus lineage, temporally and evolutionary placedbetween the species M. meridionalis (Nesti, 1825) and Mammuthusprimigenius (Blumenbach, 1799) (Garutt, 1964; Maglio, 1973;Dubrovo, 1977; Lister, 1996a). An anteroposterior shortening ofthe skull and mandible, and an increase of hypsodonty and molarplate number are the main trends along this evolutionary lineage,which in a short time interval (less than 2.5 million years) isthought to have more or less gradually transformed a woodlanddwelling generalist (M. meridionalis) to a highly specialised, cold-adapted grazer (M. primigenius). However, evidence publishedfairly recently (Lister and Sher, 2001; Lister et al., 2005; Wei et al.,2006, 2010) indicate that this simple process is less plausible, atleast for Europe, as the Mammuthus species appear much earlier inthe East Asian fossil record. This implies repeated migration of newtaxa from Asia to Europe, where they replaced the existing species,possibly after a short period of coexistence. This theory, thoughimplying a complicated evolutionary pattern, is in accordance withsome older published opinions (e.g. Depéret et al., 1923; Osborn,1942; Azzaroli, 1977) that favoured repeated immigrations of newforms into Europe as the cause of Mammuthus species turnover.

Apart from the mammoth skeletal elements, the locality yieldedalso two tortoise plastron fragments, as well as a cervid antlerfragment and a ruminant distal phalanx; the latter is attributable toa small deer, the size of Capreolus. Moreover, a coprolite was foundamong the elephant bones, suggesting the presence of a carnivore(see Section 4.1).

2. Material and methods

The excavatedmaterial was compared to recent Elephas maximusosteologicalmaterial belonging to theMuseumof Palaeontology andGeology, University of Athens, in order to determine the anatomicalposition of each specimen. The osteological description of Loxodontaafricana, published in a series of papers by Smuts and Bezuidenhout(1993, 1994), van der Merwe et al. (1995), Bezuidenhout andSeegers (1996), as well as illustrated fossil skeletons descriptions(e.g. Trevisan, 1954; Kroll, 1991), were also of great help.

The broken or fragile specimens were glued together and/orconsolidated using Paraloid B72, a highly reversible acrylic polymer(resin), which is commonly used in fossil bone preparation andconservation.

All measurements were carried out using callipers or measuringtape (for the larger specimens). Inaccurate measurements, becauseof incomplete preservation, distortion or inaccessibility, that can bereliably estimated are given in brackets. For molar measurementsand calculation of indices I followed Maglio (1973); the lamellarfrequency parameter is, however, measured only labially on theupper molars, and labially and lingually on the lower molars,because all studied teeth are in situ, not allowing measurements attheir base. The lingual side of the upper molars is also not acces-sible, because of the presence of supporting material. The bonemeasurements are anatomically oriented, i.e. parallel or perpen-dicular to the sagittal plane. ‘DAP’ stands for anteroposteriordiameter; ‘DT’ stands for transverse diameter; the ‘height’ ismeasured dorsoventrally, except for the scapula, the metapodialsand the phalanges, were it is meant proximodistally (as theanatomical orientation of these bones is not vertical). Anymeasurement technique that deviates from this general scheme isdescribed in the relevant table. The upper molars are indicated by‘M’, the lower ones by ‘m’.

3. Geological setting

The new locality ‘Loussiká’ is situated in the valley of the smallstream Serdiní, a tributary of the river Peíros, which is the majorstream of the drainage basin. The fossil-bearing deposits are offluvial origin. They consist of clayey sand, sand and coarse sand,often exhibiting cross bedding (Fig. 2), and belong to a depositionalterrace of the Peíros River system, occurring at an altitude of about50 m above sea level. The rather fine-grained sediment suggestsa low-energy fluvial environment. At the top of the section there isa calcrete-rich palaeosol and the recent soil. The terrace depositsoverlie unconformably a Pliocene shallow-marine sequence, whichcovers thewider area of western Achaia, and consists of sandstones,sandy clays and marls, rich in invertebrate fossils, mainly bivalvesand gastropods (Tsoflias and Fleury, 1980). The basement consistsof Late EoceneeOligocene flysch that belongs to the Tripoligeotectonic zone of the Alpine orogene. South of the studied areathere are also some outcrops of Cretaceous limestones of the samegeotectonic zone.

4. Taphonomy

The excavation extended in an area 6 m long (EeW) to 3 mwide(NeS), that is 18 m2. It yielded most of the cranial and axial skel-eton, except for the sacrum, the pelvis and all but one caudal

0

1 m

Pliocene marls

clayey sand

cross bedd sanded

sand, coarse sand

palaeosols

pebbles

soil

Fig. 2. Lithostratigraphy of the fluvial sediments at Loussiká locality. The mammothskeleton was found in the clayey sand layer.

Fig. 3. The distribution of the mammoth skeletal elements at Loussiká (based onoriginal sketches by L. Stavropoulou, redesigned and completed with own data). Someminor fragments are omitted for clarity reasons. Smaller bones that lie under largerones are also not shown (e.g. metapodials under the skull). Graphical scale: 1 m.

Fig. 4. Loussiká mammoth skeletal elements in situ: 1, skull (rostral part is on theright); 2, right hemimandible; 3, left hemimandible (still unexcavated; number isplaced on m2); 4, left ulna; 5, left calcaneus. The vertical white line indicates thedirection of the bulldozer cut (EeW). Note the level difference among the skull, theright and the left hemimandible, indicating rapid sediment deposition and burial.Graphical scale: 30 cm.

A. Athanassiou / Quaternary International 255 (2012) 9e28 11

vertebrae. Most of the long bones are, though, missing, except foran ulna and a tibia. The preservation of the Loussiká skeleton isgenerally good, with the exception of the skull, which suffers fromerosion and root weathering.

The recovered specimenswere not in anatomical association buttotally disarticulated and dispersed; they were scattered mainly inan NEeSW direction, in a layer of fine clayey fluvial sand (Fig. 3).Elements that are anatomically remote from each other were foundlying close together: e.g. the right tibia was lying to the left of theskull, the left ulna to the right of the skull, while the left calcaneuswas found under the right tusk sheath. Other associated elements,as the mandible and the skull, were, though, not widely separated.The recovering of delicate hyoid bones from the ventral side of theskull indicates minimal, if any, translocation of the skull (the hyoidapparatus attaches to the skull through a cartilage and it is easilylost or destroyed during decomposition of the carcass and burial).No specimen shows signs of edge rounding because of rolling, againstrongly suggesting minimal or no transportation.

The skull was found in upright position, sitting on its ventralside (which may imply an original “sudden death posture”, i.e. theanimal lying on its belly; Haynes, 1988). The mandible, broken atthe symphysis level, was found considerably lower (40e70 cm)than the skull, with its rostrum facing SSW, at the opposite direc-tion than the skull (Figs. 3 and 4). The depositional level differenceindicates rapid sediment deposition, covering most of the skeleton,except perhaps for the apex of the skull, which is eroded. Alter-natively, the mandible might be pushed in the sediment bya passing elephant that stepped on it. Most of the skull is pene-trated by roots (Figs. 5 and 8c) that have caused extensive damage,

especially at the less compact and sinuous parts of it. The left tuskwas removed from its alveolus after death and before burial, as thealveolus was found empty. Some bones exhibit post-depositionalbreakage, as the tibia, which was broken by a fault near its distalend (Fig. 5). No recovered specimen shows signs of severe distor-tion or compression.

It is quite possible that several of the remaining bones are morebroadly scattered, beyond the excavated area. Actually, several boneand tusk fragments were recovered from the disturbed sedimentNE of the excavated area, but many others may have been missed,as the disturbed sediment was widely dispersed. The originalposition of these fragments and their taphonomic relation to theskeletal elements that were excavated in situ are unknown.

The overall pattern of the skeletal elements position implies thatthe original orientation of the mammoth corpse before burial wasalso NEeSW, with the head to the NE. The area SWof the excavatedquarry may preserve the distal part of the axial skeleton, as well assome long bones. However, the excavation was not extended to the

Fig. 5. Loussiká mammoth skeletal elements in situ: 1, the occipital region of the skull(partly covered with consolidant-impregnated gauze); 2, the right tibia. Note thepresence of roots that run through the skull and the post-depositional fracture anddisplacement in tibia. Graphical scale: 30 cm.


SW mainly for financial and technical reasons. The observedNEeSW dispersion does not correspond to a hypothetical palae-ocurrent flow direction, as the excavated skeletal elements, withthe exception of the tibia, two thoracic vertebrae with long spinalprocesses and some costae, are not arranged parallel to this direc-tion (Fig. 3). Nonetheless, there is also another bone concentration,in which the elongated specimens (costae, left scapula fragment)are arranged in an NWeSE direction (Figs. 3 and 6). No sortingpattern is observed.

4.1. Biological agencies in skeleton disassociation

A possible explanation for the observed dispersal pattern is thatthe mammoth bones were scattered because of trampling and/orcarnivore activity. Extant elephants are known to be attracted to,examine and manipulate bones of their dead conspecifics usingtheir trunk and feet; they usually kick, drag, toss or carry themaround, often to long distances (Haynes, 1991, p. 157e158; Moss,2000, p. 270e271, photo plates after p. 128). The excavated speci-mens exhibit, however, no trampling marks and no long bonefractures that can be attributed to trampling proboscideans.Carnivore or rodent gnawingmarks are also absent. At least aminorcarnivore activity is, though, indicated by the presence of a bilobe,subspherical coprolite, with one concave and one conical end. Its

Fig. 6. Loussiká mammoth skeletal elements in situ: 1, left scapula; 2, right scapula;note the spine fragment ‘a’ that was broken off from its original position ‘b’ duringdeposition; 3, an accumulation of costae. Graphical scale: 30 cm.

morphology and dimensions (44 mm long, with a diameter of37e43 mm) suggest that it can be referred to a large spotted hyena(Crocuta crocuta), although its attribution to a larger hyaenidspecies as Pachycrocuta brevirostris cannot be excluded (Keiler,2001; Parfitt and Larkin in Lewis et al., 2010).

4.2. Possible human activity

A single bone, the left fourth metatarsal, bears a cluster of linearmarks on its plantar surface that exhibit the following characters:(a) they are subparallel to each other and run perpendicular to thelong axis of the bone; (b) they are sharp with well-defined borders;(c) they occur in a “recessed” area, not exposed to abrasion; (d) theyare concentrated in an area suitable for cutting the plantar muscles.Though these observations are very preliminary, they are consis-tent with human induced cut marks (Shipman and Rose, 1983;Fisher, 1984; Olsen and Shipman, 1988; Blumenschine et al.,1996). However, since no similar marks have been observed onother skeletal elements, this observation is considered a mereindication, not evidence of human activity around the mammothcorpse.

5. Systematics

Order: Proboscidea Illiger, 1811Family: Elephantidae Gray, 1821Genus: Mammuthus Burnett, 1830

Mammuthus trogontherii (Pohlig, 1885)Synonymy:

Elephas antiquus Falconer and Cautley, 1847 (Doukas andAthanassiou, 2003, p. 100, Table 3)Elephas antiquus Falconer and Cautley, 1847 (Athanassiou,2010)Elephas antiquus Falconer and Cautley, 1847 (Tsoukala et al.,2010, Table 1, p. 10)

5.1. Material

The Loussiká partial skeleton consists of the skull and mandible,the anterior part of the vertebral column, several costae, bothscapulae, left ulna, right tibia, most carpals, tarsals and meta-podials, and several phalanges. A detailed list of the recoveredanatomical parts is given in Table 1. Their distribution in themammoth skeleton is presented graphically in Fig. 7.

5.2. Description

5.2.1. SkullThe skull is partially preserved, lacking most of the sinuous

upper part, above the level of the zygomatic arches (vertex,braincase, nasal cavity, upper part of the occipital d Fig. 8). Thepreserved parts are in rather bad condition as they are muchfragmented and suffer from recent root weathering. The tuskalveoli are also damaged, lacking rostral parts, but their generalmorphology is recognizable: they are subcircular in cross sectionand diverge slightly from each other, as they bend laterally in theirrostral regions, remaining, though, very close together. In theiranteriormost preserved part, the tusks appear to have been onlyabout 5 cm apart. Only the basal part of the right tusk remains insitu. In lateral view, the skull appears high and anteroposteriorlyshortened. Both zygomatic arches are present, though badly frac-tured, forming with the tusk alveoli an angle of about 110�. Thelatter are directed downwards, forming with the molar alveolarplane an angle of about 120�. The maxilla is particularly high to

Table 1List of the recovered anatomical parts that belong to the partial skeleton ofMammuthus trogontherii from the locality of Loussiká. The anatomicaldetermination of the specimens marked with a question mark is notabsolutely positive, because of incomplete preservation. A graphicalpresentation of this list is given in Fig. 7.

Anatomical part Side

Skull with M3sTusk fragments dex.?Second upper molar dex.Mandible with m2s, m3sBasihyoidStylohyoid dex.AtlasCervical vertebra 3Cervical vertebra 5Cervical vertebra 7Thoracic vertebra 1Thoracic vertebra 2Thoracic vertebra 3Thoracic vertebra 4Thoracic vertebra 5Thoracic vertebra 6Thoracic vertebra 7Thoracic vertebra 9Thoracic vertebra 10Thoracic vertebra 11Thoracic vertebra 12Thoracic vertebra partThoracic vertebra partThoracic vertebra partThoracic vertebra partCaudal vertebra?More than 15 costaeScapula sin.Scapula dex.Ulna sin.Scaphoid dex.Lunar sin.Lunar dex.Triquetrum sin.Triquetrum dex.Pisiform dex.Trapezium dex.Trapezoid sin.Trapezoid dex.Magnum dex.Hamatum sin.Hamatum dex.Metacarpal II dex.Metacarpal III sin.Metacarpal III dex.Metacarpal IV sin.Metacarpal IV dex.Metacarpal V sin.Metacarpal V dex.Phalanx prox. manus IV sin.Phalanx prox. manus IV dex.Phalanx media manus III sin.Phalanx media manus III dex.Phalanx media manus IV dex.Ischium part sin.Ischium part dex.Femoral greater trochanter epiphysis dex.Tibia proximal part sin.Tibia dex.Calcaneus sin.Astragalus sin.Navicular dex.Cuboid dex.Metatarsal I?Metatarsal IV sin.Metatarsal IV dex.Phalanx prox. pedis IV sin.Phalanx media pedis II?Sesamoid

a

b

Fig. 7. Schematic drawing of a mammoth skeleton, showing the anatomical position ofthe excavated skeletal elements at Loussiká: a, left side; b, right side. Based ona drawing of Palaeoloxodon antiquus by C. Beauval (available at www.archeozoo.org),adapted to the mammoth morphology.


accommodate the hypsodont molars. In caudal view, only the basalpart of the occipital is preserved, with its elliptical foramenmagnum and the massive condyles. The highly pneumatisedoccipital is very much developed laterally and posteriorly (Fig. 8c).Its central part, dorsally to the foramen magnum is denser anddistinctly recessed in relation to the lateral parts. The availablecranial measurements are given in Table 2. The skull remainspartially unprepared, particularly ventrally.

5.2.1.1. Hyoid apparatus. Among the recovered cranial parts are thebasihyoid and a fragment of the right stylohyoid (Fig. 9), two of thefive ossicles (one basihyoid, two thyrohyoids, two stylohyoids) thatconstitute the hyoid apparatus. Another fragment may belong toa thyrohyoid. The hyoid apparatus is situated in the gular regionand it is articulated to the cranial base via the tympanohyal carti-lages (Shoshani and Marchant, 2001; Shoshani et al., 2007). Thebasihyoid is a dorsoventrally flattened, straight bone that becomesthicker laterally and bents slightly posterodorsally to articulatewith the left and right thyrohyoids. The Loussiká specimen is80 mmwide; the maximum andminimum diameters at the middleare 19.0 and 11.5 mm respectively. The stylohyoid is a Y-shapedbone consisting of a superior, a posterior and an inferior ramus. It isvery rare as a fossil find, but it is potentially significant for taxo-nomic and phylogenetic studies (Shoshani and Tassy, 2005;Shoshani et al., 2007). The Loussiká specimen has broken rami, sotheir relative size cannot be assessed. The identification of the rami

http://www.archeozoo.org

Fig. 8. The skull of the Loussiká mammoth: a, anterior view; b, lateral view (rightside); c, caudodorsal view in an early preparation stage (note the presence of twometapodials under the skull). Graphical scale: 20 cm.

Table 2Cranial measurements of the Loussiká mammoth (in mm).

Maximal length (parallel to the alveolar level) >1200Maximal width (at the zygomatic arches) (840)Width of the praemaxillaries (tusk alveoli) (500)Minimal distance between tusks 30Width at the level of the molar alveoli 310Occipital width >870Occipital condyles width (at their lateral borders) 220Foramen magnum height 70Foramen magnum width 91

Fig. 9. Bones of the Loussiká mammoth hyoid apparatus: a, right stylohyoid, lateralview; b, basihyoid, dorsal view. Graphical scale: 20 mm.


was based on their relative robustness, the superior ramus beingthe most robust and the inferior the least. All three rami areelliptical in cross section, more flattened medially. Due to incom-plete preservation none of the characters described by Shoshaniet al. (2007) can be scored on this specimen. A peculiarity is thatthe superior and posterior rami are not aligned but form an open

bent (an angle of 132� between them). A similar morphology isobserved in Mammuthus columbi (Shoshani et al., 2007, Fig. 3), butnot in M. trogontherii from West Runton (Lister and Stuart, 2010,Fig. 24).

5.2.2. MandibleThe mandible (Fig. 10) is very massive. The symphysis is badly

damaged, particularly at its right side. The base of the symphysealprocess is, though, preserved, indicating that the rostral processwas weak. The mandibular corpus is very robust, especially nearthe mandibular angle. Rostrally it is very deep, but it becomesgradually shallower caudally, as its ventral border curves dorsally,forming a rounded mandibular angle. The mandibular ramus ismuch developed anteroposteriorly. It has a thick caudal border thatends to the condylar process. Rostrally the ramus is much thinnerand forms the coronoid process, which is connected to the condylarone by an oblique ridge. The masseteric fossa is wide but rathershallow. Measurements are given in Table 3.

5.2.3. Upper dentitionThe tuskmorphology is known from the right tusk base, retained

in the alveolus, and fromseveral small fragments recovered fromthebulldozerdebris,whichpresumably belong to the same right tusk. In

Fig. 10. Left hemimandible of the Loussiká mammoth: a, occlusal view; b, lateral view.Graphical scale: 10 cm.

Fig. 11. Cross section of a tusk fragment of the Loussiká mammoth exhibiting theSchreger pattern. The Schreger angles are slightly acute. Graphical scale: 10 mm.


cross section the tusk is subcircular, with a maximum diameter of199 mm and a minimum of 180 mm at its base, in the alveolus. Thelargest recovered tusk fragment is 440mmlong,withmaximumandminimum cross section diameters of 203 mm and 183 mm respec-tively. Although it is very short, it exhibits a noticeable curvature.

In the natural (due to breakage) cross sections of two othersmall fragments the Schreger pattern can be observed. TheSchreger pattern is a special character of the proboscidean ivoryand has been used in genus-level taxonomy (Espinoza and Mann,1993; Palombo and Villa, 2001; Trapani and Fisher, 2003). Itconsists of two sets of radiating spiral lines that intersect with eachother, forming the Schreger angles, and reflects macroscopically thedentine internal structure. The Schreger lines in both examinedspecimens form acute angles of 87e89� near the cement/dentinejunction (Fig. 11).

The skull has both M3s in situ, while open alveoli in both sidesindicate the presence of M2s in life. A much worn molar, found

Table 3Mandibular measurements of the Loussiká mammoth (in mm).

Left Right

Maximal length (parallel to the labial wall of the corpus) 690 >610Maximal width of the corpus 185 205Maximal height (at the articulation) 480 (480)Height at the coronoid process

(parallel to its anterior border)e (360)

DAP of the ramus, at the level of the coronoid process(parallel to the labial wall of the ramus)

315 320

Corpus height, at the base of the ramus 250 235Corpus height, at the level of m2 305 e

isolated, may well represent the right M2. It consists of 8 plates anda distal talon. Its length is 153 mm and its width 95 mm. The M3sare little worn, so the distal parts of them are still in the alveoli.Nine plates plus the anterior talon are in use (a tenth plate is hardlyworn), forming broad occlusal surfaces that are 150 mm long and102 mm wide (Fig. 12). As both third molars are in situ theirmaximal dimensions and plate number cannot be measured orcounted; e.g. the width is expected to be larger towards the crownbase. Ametrical estimation based on themorphology of the maxillais given in Table 4. It is not possible to estimate the molar height,but the very high maxillas could accommodate thirdmolars as highas 200 mm. The enamel is rather thick and moderately folded andwrinkled, particularly at the anterior and median region of theocclusal surface, without, though, forming loxodont sinuses. Theenamel at the occlusal surface of the distal, slightlyworn lamellae isalmost unwrinkled. At the distalmost part of the occlusal surfacethe incipient wear produces a transverse series of subcircularenamel islets; the median islets merge together with advancingwear, resulting in a transversely elongated islet.

5.2.4. Lower dentitionThe two hemimandibles bear both second and third molars

(Figs. 10a and 13). The former have triangular occlusal surfaces andare totally worn. They consist of three plates that are worn down tothe roots. The left m3 has the anterior talonid and nine plates in

Fig. 12. Loussiká mammoth upper left M3, occlusal view. Anterior side is on the left.Graphical scale: 30 mm. The molar is in situ; its non-occlusal part and the maxilla arenot shown, because they are covered by supporting material.

Table 4Upper and lower third molar measurements (in mm) of the Loussiká mammoth.Next to the width is given the plate number at the level of which it was measured.Plate number excludes any talons.

Left M3 Right M3 Left m3 Right m3

Length (350) (350) >300a >290a

Width 102 e 100 (at 5th) 98 (at 4th)Plate number >14 >14 >16b >17b

Plate frequency 6.3 6.3 6.1 6.2Enamel thickness 2.6e2.9 2.6e3.0 2.7e3.2 2.7e3.2

a Estimated 370e410.b Estimated 20e22.

Fig. 14. Loussiká mammoth cervical vertebrae: a, atlas, cranial view; b, seventhcervical vertebra, caudal view. Graphical scale: 10 cm.


use, while 16 plates are totally visible. It is estimated that another4e6 plates are still inside the alveolus at the posterior part of themandibular corpus, raising the total plate number to 20e22. Itstotal length could be greater than 400 mm. Based on the consid-erable depth of the mandibular corpus the molar may be as high as200 mm. The occlusal surface is rather broad and oval in shape. Theenamel is fairly thick (thicker than that of the upper molars) andmoderately wrinkled at the medial part of the occlusal surface. Theenamel of the slightly worn distal lamellae (seventh and eighth)forms a transversally elongated islet and two elliptical oneslingually and labially of it. The right m3 is similar to the left, but has10 plates in use and 17 visible. Metrical data are given in Table 4.

5.2.5. Vertebral column and ribsOnly the anterior part of the vertebral column is available. The

vertebrae have the typical elephantid morphology (Bezuidenhoutand Seegers, 1996). The atlas is stoutly built and has much devel-oped lateral wings (Fig. 14a). Apart from atlas, the cervicals wereidentified as third, fifth and seventh, mainly because they do not fitto each other when trying to re-articulate them. The seventh(Fig. 14b) is also morphologically distinct, as it has articular facetsfor the first pair of ribs ventrolaterally in its caudal side and it lacks

Fig. 13. Loussiká mammoth mandibular dentition, occlusal view: a, left m2 and m3; b,right m2 and m3. Graphical scale: 10 cm.

foramens in its transverse processes. The articular surfaces of allavailable cervical vertebrae are fused to the bodies. Measurementsof the cervical vertebrae are given in Table 5.

The preserved thoracic vertebrae are characterised by their longspinal processes, particularly the anterior ones. The spinalprocesses are strongly inclined backwards, forming with thevertebral body plane an angle of 50�e60� (the angle increasescaudally), quite unlike certain E. antiquus specimens, as that fromUpnor that has almost vertical spinal processes (Andrews andForster Cooper, 1928). (The Upnor vertebral column may be,though, morphologically aberrant, as in other E. antiquus skeletonsthis character is less accentuated d e.g. Trevisan, 1954; Maccagno,1962; Melentis, 1963; Kroll, 1991). The exact anatomical position ofthe at least 13 thoracic vertebrae is generally not known, as theywere not found in anatomical association and it is not possible to

Table 5Cervical vertebral measurements of the Loussiká mammoth (in mm).

Atlas Third Fifth Seventh

Maximal height 213 >240 >290 362Maximal width 431 305 e 337Cranial articular width 228 154 144 150Cranial articular height 112 164 168 (140)Caudal articular width 195 171 181 155a

Caudal articular height 101 167 166 159DAP of the corpus

(measured ventrally, at the sagittal plane)53 57 87

DAP of dorsal arc 90

a Excludes the articular surfaces for the first ribs.

Table 6Thoracic vertebral measurements (in mm) of the Loussiká mammoth. The numbering of the “4th” to “12th” is mostly tentative, used to indicate the relative position of thevertebrae, and corresponds to their approximate anatomical position in life. The remaining thoracic vertebrae were fragmentary and not measurable. The articularmeasurements exclude the articulations for the costae.

1st 2nd 3rd 4th 5th 7th 9th 11th 12th

Maximal height 595 (535) >580 e >550 e e e 360Maximal width 344 e (320) (306) 316 e 320 221 243Cranial articular width 157 e 139 130 e 122 e e e

Cranial articular height 149 e 123 141 e 139 e e e

Caudal articular width 156 e 142 139 e e 142 128 127Caudal articular height 130 e (125) (141) e (139) 140 137 123Spinal process “length”

(from the top of the vertebral foramen to the apex of the process)380 510 >450 e >440 e e e 315

DAP of the corpus(measured ventrally, at the sagittal plane)

82 e e 77 (83) 77 (78) (90) (80)


re-articulate them, because most of them have incompletelypreserved bodies or lack one of their body epiphyses (not yetfused). An exception is the second (Fig. 15a) and third, which werefound associated to each other and were identified mainly on theirvery long and robust spinal processes. The best preserved thoracicvertebrae were positioned relative to each other using the char-acters that change gradually in craniocaudal direction: raising ofthe transverse processes; increased inclination of the spinalprocess; raising of the articulations for the costae; changing of thebody shape to more triangular (in cranial or caudal view); changingof the vertebral foramen shape from triangular to elliptical (incranial or caudal view) (Kulczyski, 1955; Bezuidenhout and Seegers,1996). The more caudally placed thoracic vertebrae (starting from

Fig. 15. Loussiká mammoth thoracic vertebrae: a, second, caudal view; b, abouttwelfth, caudal view. Graphical scale: 10 cm.

the “ninth”) (Fig. 15b) have bodies with unfused anterior articula-tions. The posterior articulation is fused in all available thoracicvertebrae. None of the studied thoracic vertebrae exhibits anymarked bilateral asymmetry in the development of the transverseprocesses or the articular facets, as it is often found in individuals ofthe genus Mammuthus (Lister, 2009; Lister and Stuart, 2010).Measurements of the thoracic vertebrae are given in Table 6. Nolumbar or sacral vertebrae were found. A much eroded specimenpossibly represents one of the first caudals.

The costae are fragmentary. The anterior ones are morphologi-cally distinct, as their head forms an almost right angle with theirbody. This angle becomes more obtuse caudally, as the costal headarticulates at a gradually more dorsal area on the vertebral bodies.Due to their incomplete preservation the costae were not identifiedanatomically; their relative position was just characterized asanterior, middle or posterior. A tentative anatomical position of theavailable costal fragments is given in Fig. 7.

5.2.6. Thoracic limbThe thoracic limbs are represented by both scapulae, left ulna,

most of carpal and metacarpal bones and five phalanges. Theirdimensions are given in the Tables 7e11. The scapulae are incom-pletely preserved. The left one preserves only the anterior andcentral part along the spine, lackingmost of the posterior blade part(fossa infraspinata). The spine is almost straight, presenting a weakconcavity cranially. It preserves both hamate and suprahamateprocesses, which form an angle of 35� between them. The rightscapula is more complete, lacking the cranial part of the blade (fossasupraspinata). Both have fusing proximal epiphyses, the fusion linesremaining well visible. Their dimensions are quite similar to themammoth specimens from Steinheim (Dietrich, 1912) and WestRunton (Lister and Stuart, 2010). The latter has, however, quitelarger glenoid cavity. Compared to the Grevená E. antiquus speci-mens (Tsoukala and Lister, 1998), the Loussiká scapulae are pro-portionally wider anteroposteriorly (though equally high) and have

Table 7Scapula measurements of the Loussiká mammoth (in mm).

Left Right

Maximal height (parallel to the spine) 1070 1045Articular height (parallel to the spine) 1010 990Length of the dorso-caudal border

(from the dorsal end of the spine to the caudal angle)e (990)

Length of the caudal border(from the distal articulation to the caudal angle)

e 855

DAP of the distal end 290 e

DT of the distal end (140) e

DAP of the distal articulation 216 e

DT of the distal articulation (135) 145

Table 8Ulna measurements of the Loussiká mammoth (in mm).

Maximal height 975Articular height (from the medial coronoid process

to the distal articulation)825

DAP of the proximal end 300Maximal DT of the proximal end (articular level) 243DT of the proximal articulation 235DAP of the olecranon 230Minimum DAP of the diaphysis 127Minimum DT of the diaphysis 110DAP of the distal end 186DT of the distal end 213DAP of the distal articulation 102DT of the distal articulation 152

Table 10Metacarpal measurements of the Loussiká mammoth (in mm).

Metacarpal II RightMaximal height 192Minimal DT of the shaft 73DAP of the distal end 85DT of the distal end 104DAP of the distal articulation 92DT of the distal articulation 85Metacarpal III Left RightMaximal height 221 220DAP of the proximal end 126 e

DT of the proximal end 82 84DAP of the proximal articulation 112 110DT of the proximal articulation 82 84Minimal DT of the shaft 72 71DAP of the distal end 95 93DT of the distal end 95 99DAP of the distal articulation 95 93DT of the distal articulation 86 85Metacarpal IV Left RightMaximal height 201 205DAP of the proximal end 117 119DT of the proximal end e 99DAP of the proximal articulation 103 104DT of the proximal articulation e 99Minimal DT of the shaft 82 82DAP of the distal end 98 98


distinctly smaller glenoid cavity (articular) dimensions. The sameobservation on the glenoid cavity dimensions applies also whencomparing the Loussiká scapulae to E. antiquus specimens fromItaly (Trevisan, 1954; Maccagno, 1962). Compared to E. antiquusfrom Upnor (Andrews and Forster Cooper, 1928) the Loussikáspecimens are smaller in all dimensions.

The ulna (Fig. 16a) is robust with a massive olecranon. In lateralaspect it is rather straight and not bent as in Loxodonta (asfigured by

Table 9Carpal measurements of the Loussiká mammoth (in mm).

Scaphoid RightMaximal height 142Maximal DAP 119Lunar Left RightMaximal height 83 85Maximal DAP 145 141DAP of the proximal articulation e 114DT of the proximal articulation e 135DAP of the distal articulation 126 121DT of the distal articulation (141) 148Triquetrum Left RightMaximal DAP e 138Maximal DT e 146DAP of the proximal articulation 100 e

DT of the proximal articulation e 118DAP of the distal articulation 116 121DT of the distal articulation e 142Pisiform RightMaximal DAP 144Maximal height 57Minimal DT (at the middle of the bone) 75Trapezium rightMaximal height 97Maximal DT 95Trapezoid Left RightMaximal height 75 71Maximal DAP 126 126Maximal DT 88 85DAP of the proximal articulation 101 100DT of the proximal articulation 66 66DAP of the distal articulation 96 103DT of the distal articulation 71 63Magnum RightMaximal height 123Maximal DAP 155DAP of the proximal articulation >120DT of the proximal articulation 106DAP of the distal articulation e

DT of the distal articulation 77Hamatum Left RightMaximal height 123 (125)DAP of the proximal articulation 122 128DT of the proximal articulation 114 (119)DAP of the distal articulation 136 e

DT of the distal articulation 122 e

DT of the distal end 103 100DAP of the distal articulation 96 96DT of the distal articulation 89 90Metacarpal V Left RightMaximal height 181 181DAP of the proximal end 114 105DT of the proximal end 94 90DAP of the proximal articulation e 95DT of the proximal articulation 83 83DAP of the distal end 116 117DT of the distal end 97 95DAP of the distal articulation 115 117DT of the distal articulation (86) 83

Smuts and Bezuidenhout, 1993, Figs. 12 and 13). The diaphysis isgenerally triangular in cross section, though it gradually becomesalmost square-shaped towards the distal extremity of the bone. Thedistal epiphysis, while firmly attached to the shaft, is not completelyfused, exhibiting a deep cleft caudally, along the fusing line. TheLoussiká ulna is metrically smaller and slenderer than most speci-mens attributed to male M. trogontherii (Steinheim, Untermaßfeld,

Table 11Phalanges manus measurements of the Loussiká mammoth (in mm).

Phalanx prox. IV Left RightHeight 93 100DAP of the proximal end 71 72DT of the proximal end 94 (97)DAP of the proximal articulation 60 58DT of the proximal articulation 78 (82)DAP of the distal end 52 52DT of the distal end 76 77Phalanx media III RightHeight 61DAP of the proximal end 47DT of the proximal end 70DAP of the proximal articulation 43DT of the proximal articulation 53DAP of the distal end 32DT of the distal end 63Phalanx media IV RightHeight 43Maximal DAP 38Maximal DT 50

Fig. 16. Elements of the Loussiká mammoth thoracic limb: a, left ulna, lateral view; b, right pisiform, medial view; c, right scaphoid, lateral view; d, right triquetrum, proximal view;e, right triquetrum, distal view; f, right lunar, proximal view; g, right lunar, distal view; h, right trapezium, lateral view; i, right trapezium, medial view; j, right trapezoid, proximalview; k, right trapezoid, distal view; l, left hamatum, proximal view; m, left hamatum, distal view; n, right magnum, proximal view; o, right magnum, distal view. In proximal anddistal views the anterior side is facing towards the bottom of the figure. Graphical scales: 20 cm (ulna), 5 cm (carpals).


Odessa, West Runton, Kostolac d Dietrich, 1912; Dubrovo, 2001;Lister and Stuart, 2010) and similar or smaller than specimensattributed to E. antiquus (Andrews and Forster Cooper, 1928;Trevisan, 1954; Maccagno, 1962; Melentis, 1963).

The right carpus is complete, represented by all eight bones; theleft lacks the scaphoid, the pisiform, the trapeziumand themagnum.The bones of the right carpus can be re-articulated very well to eachother, despite damage at the anterior part of the magnum, revealing


an aserial carpus structure (the lunar extends well over bothmagnumand trapezoid). The scaphoid (Fig.16c) is triangular in shapeand laterallyflattened. It is stouter than that of Loxodonta (Smuts andBezuidenhout, 1993) and it is very close to the morphology ofM. primigenius, as depicted by Andrews and Forster Cooper (1928, pl.III). Its radial facet is high and ends proximally to a pointed top. Thedistal facet (for the trapezoid) extends proximolaterally till about themiddle of the bone. The triangular-shaped lunar (Fig. 16f and g) iswider than themagnum (situated just below it), also overlapping thetrapezoid. This disposition has been considered characteristic for themammoths (e.g. Trevisan,1954, Figs. 22 and47), though later authorsoftenattribute it to individualvariation (e.g. BedenandGuérin,1975).The triquetrum (Fig.16d and e) is also of triangular shapewith a longapophysis that projects lateropalmarly. Both, left and right, aredamaged, particularly the former, which is broken palmarly. Thepisiform (Fig.16b) is the lateralmost bone of the proximal carpal row.It is elongated, projecting palmarly. The trapezium (Fig. 16h and i),situatedmedially in the distal carpal row, is mediolaterally flattenedand pentagonal in medial aspect, due to a tuberosity at its palmarside. The trapezoid (Fig. 16j and k) has two main triangular articularsurfaces proximally and distally. The distal articular surface extendslaterocaudally to form a facet for the magnum, unlike certainE. antiquus specimens (Andrews and Forster Cooper, 1928, pl. IV;Trevisan, 1954, Fig. 21). Cranially there is only one facet for themagnum, which corresponds to a similar unified facet in themagnum. This is considered as a character of Mammuthus, notobserved inE. antiquus (Dubrovo, 2001). Themagnum(Fig.16nando)is a massive, cuboid bone with a prominent palmar tuberosity. Itsproximal articular facet has almost parallel lateral and medialborders. The distal facet for the secondmetacarpal iswell developed,extending to the whole length of the distal articulation. This facet isusually reduced dorsally in E. antiquus (Beden and Guérin, 1975),though the presence of non-reduced facets is also reported (Trevisan,1954, p. 36e37; Maccagno, 1962, p. 112). The hamatum (Fig. 16l andm) bears two large articular surfaces for the triquetrum and the fifthmetacarpal thatmeet eachother along the lateralmarginof the bone.The contact of these surfaces is very short in E. antiquus (Beden andGuérin, 1975).

The metacarpals (Fig. 17) are elongated, robust, dorsopalmarlyflattened bones, with wide proximal articular facets for the distalcarpal row and a distal trochlea for the articulation of the proximalphalanges. The proximal articulation of the third metacarpal isnoticeable, because it is very prominent, the facets for the magnum

Fig. 17. Loussiká mammoth right metacarpal IIeV series, dorsal view. The MC II is on th

and the hamatumbeing strongly inclined and formingan acute anglebetween them. Metrically the Loussiká metacarpals are smaller andrather less stout than certain E. antiquus ones (Andrews and ForsterCooper, 1928; Trevisan, 1954, Figs. 25 and 26; Tsoukala and Lister,1998). Compared to the metacarpals of M. trogontherii from Stein-heim theyare also smaller, particularly thefifthmetacarpal (Dietrich,1912).

5.2.7. Pelvic limbThe pelvis fragment comes from the symphyseal area of the

ischial bones at the caudal part of the pelvis. The symphysis isfused, but the fusion line is clearly demarcated. The ischium heightat its caudal area (at the ischial tuberosity) is about 33 cm. No othermeasurement can be taken on this specimen. The only femoralremain is the epiphysis of the greater trochanter of the right femur.It has no trace of any incipient fusion with the femoral body. Theright tibia (Fig. 18a) is well preserved, though it is somewhatdeformed at its proximal end. It is robust and fully grown, bothepiphyses being already fused. Of the left tibia only the proximalend is available. Measurements are given in Table 12.

The calcaneus (Fig. 18b) is heavily built with very rugose non-articular surfaces, particularly at the tuber. The facet for thefibula, as well as the lateral part of the ectal facet, are missing. Thesustenactular facet is markedly triangular. The general morphologyof the bone is very similar to that of M. primigenius calcaneus asdepicted by Andrews and Forster Cooper (1928, pl. VII). Metrically(Table 13) it is smaller than the M. trogontherii specimens fromSteinheim and Untermaßfeld (Dietrich, 1912; Dubrovo, 2001), aswell as most of those attributed to E. antiquus (Andrews and ForsterCooper, 1928; Trevisan, 1954, Figs. 25 and 26; Tsoukala and Lister,1998), except for Riano (Maccagno, 1962, p. 116).

The astragalus (Fig. 18e and f) is low and broad, flattened ina dorsoplantar direction. The morphology and the disposition of itsarticular surfaces are very similar to thoseofM.primigeniusaccordingto Andrews and Forster Cooper (1928, pl. VI) and to theWest Runtonspecimen (Lister and Stuart, 2010, Fig. 7). Metrically (Table 13) theastragalus from Loussiká is smaller than the specimens from Stein-heim, Untermaßfeld and West Runton (Dietrich, 1912; Dubrovo,2001; Lister and Stuart, 2010) and larger that the specimens fromLa Fage, attributed toM. aff. trogontherii (Beden and Guérin, 1975).

The other two recovered tarsal bones, navicular and cuboid,belong to the right foot. The navicular (Fig. 18c and d) is flat witha concave proximal and a convex distal articular surface, with four

e right. The figured MC V is the left one (pictured mirrored). Graphical scale: 5 cm.

Fig. 18. Elements of the Loussiká mammoth pelvic limb: a, right tibia, dorsal (anterior) view; b, left calcaneus, cranial view; c, right navicular, proximal view; d, right navicular, distalview; e, left astragalus, proximal view; f, left astragalus, distal view; g, right metatarsal IV, dorsal view; h, right cuboid, proximal view; i, right cuboid, distal view. Graphical scales:20 cm (tibia), 5 cm (tarsals and metatarsal).

Table 13Tarsal measurements of the Loussiká mammoth (in mm).

Calcaneus LeftMaximal height (normal to the cuboid articulation) 222Maximal diameter of the head (tuber calcaneus) 143Maximal DAP (normal to the astragalus articulations) 141Total DT of the articular surfaces for the astragalus 197Maximal diameter of the lateral articular surface for the astragalus (112)Maximal diameter of the medial articular surface for the astragalus 102Astragalus LeftMaximal height (parallel to the calcaneus articulations) (150)


facets; the facet for the cuboid is the largest. At its caudal surfacethe facet for the calcaneus is small. The posteromedial part of thebone has eroded surfaces. It is smaller than the navicular fromUntermaßfeld (Dubrovo, 2001) and very similar in size to the smallernavicular fromStreinheim (Dietrich,1912). The cuboid (Fig.18h and i)is triangular in shape. Unlike the cuboid from Upnor (E. antiquus dAndrews and Forster Cooper, 1928, pl. VIII) it does not have a prom-inent tubercle in its caudal surface. Measurements are given inTable 13.

The fourth metatarsal (Fig. 18g) is a short and robust bone witha very faintly divided triangular proximal articulation for theectocuneiform and the cuboid. The corpus is trapezoid in crosssection, wider dorsally. It is shorter than the metatarsal IV fromSteinheim (Dietrich, 1912) and metrically similar to the same boneof E. antiquus from Riano (Maccagno, 1962). Measurements aregiven in Table 14.

Table 12Tibia measurements of the Loussiká mammoth (in mm).

Left Right

Maximal height e 750DAP of the proximal end 202 205Maximal DT of the proximal end 257 264DAP of the proximal articulation 144 142DT of the proximal articulation 223 238Minimum DT of the diaphysis e 129DAP of the distal end e 160DT of the distal end e >190DAP of the distal articulation e 126DT of the distal articulation e 172

The available phalanges manus and pedis are not differ-entiated morphologically from the typical of the familyElephantidae. Their measurements are given in Tables 11 and 15respectively.

Maximal DAP (normal to the calcaneus articulations) 90Maximal DT (parallel to the calcaneus articulations) 177Navicular RightMaximal DAP (100)Maximal DT 153DAP of the proximal articulation 82DT of the proximal articulation 119DAP of the distal articulation 84DT of the distal articulation e

Cuboid RightHeight 58Maximal DAP 123DAP of the proximal articulation 91DT of the proximal articulation 104DAP of the distal articulation 95DT of the distal articulation 103

Table 14Metatarsal measurements of the Loussiká mammoth (in mm).

Metatarsal IV Left Right

Maximal height 148 144DAP of the proximal end 86 e

DT of the proximal end 83 83DAP of the proximal articulation 82 e

DT of the proximal articulation 83 83DAP of the distal end 83 82DT of the distal end 86 76DAP of the distal articulation 83 82DT of the distal articulation 69 68

Table 16Correlation between combined scapula and ulna articular height (in cm) and themeasured or estimated skeletal shoulder height (in cm) of Mammuthus skeletons.Data taken from Lister and Stuart (2010, Suppl. Table B).

Skeleton Species Gender Scapula þ ulnaarticular length

Skeletal height

Condover M. primigenius _ 155 296Praz Rodet M. primigenius _ 151 280Olyosh M. primigenius \ 120 215Loussiká M. trogontherii _ 184 363a

West Runton M. trogontherii _ 190 369Edersleben M. cf. trogontherii \ 170 345Nogaisk M. meridionalis _ 220 410Georgievsk M. cf. meridionalis _ 193 396

a Estimated from the graphical presentation of the data (Fig. 19).

220

250

280

310

340

370

400

430

shou

lder

heig

ht(c

m)

M. primigenius

M. trogontherii

M. meridionalis


6. Inferred physical and ontogenetic characters

6.1. Stature

A simple method for estimating the shoulder height in life of theLoussikámammothwouldbe to add together the articularheights ofthe thoracic limb bones, and then correct for the non-preserved softtissues and cartilage (e.g. Osborn, 1942, p. 1022). This is because thelongbones ofmost Proboscidea (with the exceptionof archaic small-sized forms) are anatomically placed one under another, due to thecolumnar stance of the limb. However, a humerus has not beenfound among the excavated bones, so this method cannot be fol-lowed here. A less accurate estimation can, though, be based incomparisons of the available limb bone (scapula, ulna and tibia)dimensions with those of other mammoth skeletons of knownshoulder height. The tibia is rather problematic for stature estima-tion, as it stops growing early in ontogeny, not reflecting thesubsequent body growth. The distal epiphysis of ulna remainsunfused until late in life (Roth, 1984; Lister, 1999), constitutinga better predictor. Christiansen (2004) gives correlated ulnadimensions with shoulder height for the M. meridionalis skeletonfromDurfort, mounted at theMuséumNational d’Histoire Naturellein Paris. This skeleton is 383 cm high and its ulna has a maximalheight of 1083 mm. Similarly, the M. trogontherii skeleton fromSteinheim is 370 cm high and its ulna maximal height is 1075 mm(Dietrich,1912). The Loussiká ulna is 10% and9% smaller respectivelyand d hypothesizing an isometric limb bone scaling d we getequally shorter statures of 345 and 336 cm respectively. Nonethe-less, this simplistic approachmayunderestimate the actual height atwithers, as the Loussikámammoth has relatively large scapulas andnot fully fused ulna. Comparing the combined left scapula and ulnaarticular length with corresponding measurements on otherM. trogontherii skeletons taken from Lister and Stuart (2010, Suppl.Table B) a higher shoulder height is predicted. The available data aregiven in Table 16 and presented graphically in Fig. 19. The trend lineillustrated in the graph predicts a shoulder height of 363 cm for theLoussiká skeleton. If the female individuals are not taken intoaccount, then the predicted Loussiká skeleton height increases by5e6 cm. It should be noted however, that Lister and Stuart (2010)consider scapula and ulna as problematic bones for skeletal heightestimation (mainly because of inconsistent measurement methodsamong authors) and, consequently, the estimated stature of 363 cm

Table 15Phalanges pedis measurements of the Loussiká mammoth (in mm).

Phalanx prox. IV Left

Height 96DAP of the proximal articulation 65DT of the proximal articulation 66DAP of the distal articulation 57DT of the distal articulation 52

should be considered as highly approximate. To estimate the liveshoulder height from skeletal height another 6.3% (Osborn,1942) or15 cm (Christiansen, 2004) has to be added, resulting in 386 or378 cm respectively.

6.2. Body mass

The body mass is typically estimated using humerus or femurdimensions (maximum height, diaphyseal circumference), orshoulder height. In the absence of humerus or femur at Loussiká theestimated shoulder height has to be used. This is already prob-lematic at the beginning, as the estimated skeletal height is veryapproximate, and because extant elephants are smaller andanatomically different frommammoths, as well as from each other.Moreover, two elephants of identical shoulder height can differ inbody mass by a factor of 2 (Roth, 1990) rendering any estimationeven less accurate.

Two approaches were used for the estimation of the Loussikáindividual body mass from the shoulder height. The first is a linearregression based on a data sample ofmale E. maximus (Christiansen,

190100 120 140 160 180 200 220 240

scapula + ulna height (cm)

Fig. 19. Graphical presentation of the correlation between combined scapula and ulnaarticular height and the measured or estimated skeletal shoulder height of Mammu-thus skeletons (see also Table 16). The two female skeletons are marked as such.Comparative data (open circles) taken from Lister and Stuart (2010, Suppl. Table B). Theestimated position of the Loussiká skeleton is marked with a black circle.


2004, Table 2) and figured in Fig. 20. Using a live shoulder height of380 cm, a body mass of 7850 kg is estimated for the Loussikámammoth. The second approach estimates the bodymass using theequation BM¼ 5.07� 10�4� SH2.803, where BM is the bodymass inkg and SH the shoulderheight in cm (Christiansen, 2004, p. 527). Theequation is based on a sample of male L. africana from Uganda. Fora shoulder height of 380 cm the equation yields a body mass of8633 kg. In conclusion, based on the available metrical data andhaving inmind the rough approximation of the results, a bodymassof about 8 t is estimated for the Loussiká mammoth.

6.3. Gender determination

The extant elephants exhibit evident sexual dimorphism inseveral characters, the most obvious of which is that the males areconsiderably larger than females. In fact elephants count among themost sexually dimorphic mammals: the males are about 20e40%larger in linear dimensions and can be almost twice as heavy as thefemales (Haynes, 1991; Sukumar, 2003). Moreover, the tusks offemale individuals are much shorter, thinner and less curved. Othersexual dimorphic characters include the shape of pelvis andmandible (relativewidth, relative corpusheight, developmentof thesymphyseal process), aswell as themore rapid and continuous bodysize growth of males till late age. Similar dimorphism patterns havealso been observed in fossil proboscideans, as mammoths(M.primigenius,M. columbi) andmastodonts (Mammut americanum)(Haynes, 1991; Averianov, 1996; Lister, 1996b). Inference about theLoussiká mammoth gender can be drawn from skeletal size,symphyseal process development, presence of unfused epiphysesand tusk size (as a reasonably complete pelvis is not available).

The estimated skeletal shoulder height of the Loussiká individualis placed in the range of male skeletons ofM. trogontherii (Lister andStuart, 2010, Table 1), being 10 cm higher than the Azov II skeleton,which is referred to a very large female. The limb bone dimensionsare generally smaller than corresponding male M. trogontheriidimensions published in the literature (see Sections 5.2.6 and 5.2.7).

4000

4500

5000

5500

6000

6500

7000

7500

8000

8500

260 280 300 320 340 360 380 400

body

mas

s(k

g)

shoulder height (cm)

Fig. 20. Linear regression of male Elephas maximus shoulder height to body mass,based on Christiansen (2004, Table 2). The estimated position of the Loussiká skeletonis marked with a black circle.

This would allow the attribution of this skeleton to a rather smallmale. However, an assignment to an exceptionally large female,though improbable, cannot be literally rejected; no clear-cutboundary can be drawn between male and female dimensionalranges, particularly when temporal and/or geographical variation ispossible among compared fossil samples.

The inferred small symphyseal process of the Loussiká mandiblehas been considered as a female character, nonetheless not unam-biguously, as there are several recorded exceptions: Averianov(1996, Table 26.1) reports female woolly mammoth specimenswith developed process, as well as male specimens with smallsymphyseal process. Thus this character cannot be conclusive.

Another sexually dimorphic character is the presence of not fullyfused bones inmale individuals of advanced age, as they continue togrow as adults (Roth, 1984; Haynes, 1991). The occurrence of fusingor unfused limb bone epiphyses in a mammoth in its forties (seeSection 6.4), as the Loussiká individual, is a clear indicationof amale.

The maximum base diameter of the preserved tusk fragments atLoussiká (199e203 mm) is placed among the high values reportedfor Mammuthus samples. A large sample of M. primigenius tusksgave maximum base diameters well smaller than 200 mm formales (a value comparable to that of Loussiká) and 90 mm forfemales (less than half the diameter of the Loussiká tusk)(Averianov, 1996, based on Vereshchagin and Tikhonov, 1986). Theavailable published measurements on M. trogontherii specimensare rather few. The tusk of the West Runton mammoth, which isassigned to amale individual, has a base diameter of 217mm (Listerand Stuart, 2010, Table 2), slightly larger than Loussiká. Other maleM. trogontherii specimens measure 210/175 mm (max/min basediameter) (Steinheim skeleton, Dietrich, 1912) and 215 mm (Azov Iskeleton, Baigusheva and Garutt, 1987, cited by Lister and Stuart,2010). Specimens assigned to female M. trogontherii from Edersle-ben and Novogeorgievsk, have base diameters of 138 and 149 mmrespectively (Lister and Stuart, 2010). The maximum base diameterof specimens attributed to M. meridionalis can be larger (e.g.238 mm in L’Aquila, Maccagno, 1962, Fig. 17) 230e250 mm inChilhac (Boeuf, 1983, p. 179). The large tusk diameter of the Lous-siká individual is a clear indication of a male animal, as no femalewould possess such a robust tusk. This character, together with theevidence of continuing growth mentioned above, are consideredhere as the most reliable of the available anatomical evidence forgender determination and thereby the Loussiká mammoth isreferred to a male.

6.4. Ontogenetic age

The ontogenetic age of living and fossil elephantid individuals iscommonly estimated by identifying the cheek teeth in use andobserving their wear state. The method is based on the almosthorizontal eruption and progression of upper and lower molari-form teeth, which gradually bring the more distal molar lamellaeandmolars at the occlusal area. This peculiar type of molar eruptionis characteristic of the Elephantidae. Laws (1966) established anontogenetic scheme of thirty age groups (IeXXX) for Loxodonta,defined by mandibular dental progression stages, and assigneda corresponding age in years for each group. Though widely used,Laws’method has received some criticism, as the resulting ages lessthan 30 years have been considered as overestimated (e.g. Lark,1984; Jachmann, 1985). An amendment of the method for the agegroups of 10e30 years was published by Jachmann (1988). Despitethe possibility of introduced errors (which can simply result fromvariations in diet among studied populations that may produceslightly different occlusal wear patterns in individuals of the sameage) Laws’ method remains a useful tool in estimating the onto-genetic age of an African elephant.


The Laws’ method has been applied in fossil elephantid species,as they also exhibit the same horizontal molar progression. As fossilelephantids do not necessarily have the same plate number in eachmolar as Loxodonta, a proper adjustment has to be done where themethod refers to the number of plates in use. A comparison of theocclusal surface of the Loussiká mandible (Figs. 10a and 13) to Laws’(1966) figures leads to the conclusion that the studied individual isplaced to the age group XXII. In this group the m2 has only 2e3remaining enamel lamellae or it is shed, leaving an open alveolus,while the m3 has 6e7 lamellae in use. Given a mean Africanelephant m3 plate number of 13 (Maglio, 1973, Table 11) and anestimated plate number of 20e22 for the Loussiká m3s, the 9e10lamellae in use in the Loussiká mandible are equivalent to sixlamellae in Loxodonta. The almost totally worn m2s of the Loussikámandible are also consistent with the m2 state in this age group.

Laws (1966) assigns an age of 39 � 2 years to group XXII.Assigning this age to the Loussiká individual would, however, be anunderestimation of its real age at death, as a species’ longevity isknown to scale positively to body size (Eisenberg, 1990; Maiorana,1990). M. trogontherii was a much larger animal than L. africana, soit is expected to have a longer lifespan. A 40 years old male Africanelephant weights about 5 t (Laws, 1966, Fig. 14), while the bodymass of the Loussiká individual is estimated to 8 t (see Section 6.2).Using the formula given by Maiorana (1990) (longevity scales asa 0.25 power of body mass) an age of death of about 45 years can beestimated for the Loussiká mammoth.

Studying Asian elephant samples in a similar way to Laws(1966), Roth and Shoshani (1988, Table 2) cite an age of 37 yearsfor a specimen, which has more than half of m2 in wear as well asa slightly worn m3. This information that refers to another extantspecies (E. maximus) corroborates the estimated age of the some-what older Loussiká individual.

An estimation of the individual ontogenetic age can also bebased on the epiphyseal synostosis state. This is possible becausethe ontogenetic growth of elephants does not cease when the adultage is reached d as it is common among mammals d but theycontinue to grow as adults (Roth, 1984; Haynes, 1991). As the bonegrowth principally takes place between the body and the epiph-yses, several epiphyseal sutures remain open until late in life. Thefusion sequence, as well as the ontogenetic stage duringwhich eachfusion occurs, are generally consistent among individuals andamong extant and fossil species (though minor differences dooccur) (Roth, 1984; Haynes, 1991; Lister, 1999). However, they differconsiderably between the genders, as the males have much moreprolonged growth period in order to acquire their much greaterbody size. For any given age class based on dental eruptionsequence, as those of Laws (1966) mentioned above, femalesgenerally have more already fused epiphyses than males.

The Loussiká partial skeleton does comprise limb bones, theepiphyseal synostosis state of which can contribute to the onto-genetic age estimation. These are the scapulae, the ulna, thetrochanteric epiphysis of the femur and the tibia. The latter is theonly fully grown, with fused epiphyses, the former two have fusingepiphyses, while the trochanteric epiphysis is completely detached(found isolated). Studies in extant elephant samples (Roth, 1984;Haynes, 1991, Appendix) have documented the fusing sequencesof the limb bone epiphyses: Both tibial ends stop growing in Lox-odonta males during the age classes XVIIIeXX and distal ulna fusesduring or after XXII. The ossified scapular cartilage also fuses late inlife (after class XXI) in Loxodonta (Roth, 1984; Smuts andBezuidenhout, 1993) and during or after the XXIIeXXIII classes inM. primigeniusmales (Lister, 1999). The trochanteric epiphysis fusesduring the classes XXIIeXXV in extant female Elephas, during XXI ina single male Loxodonta specimen and during XXIIIeXXV inM. primigenius (Roth, 1984; Lister, 1999).

The vertebral body epiphyses, several of which remain unfusedin the Loussiká skeleton, do not completely fuse to the bodies untilvery late in life (sixth decade in both males and females) in extantAfrican elephant populations (Haynes, 1991). This allows thevertebrae to continue to grow anteroposteriorly, even when theindividuals have stopped growing in terms of shoulder height.

In conclusion, the synostosis state observed in the Loussikáskeleton is concordant with the fusing sequences of E. maximus, L.africana and M. primigenius and also corroborates the ontogeneticclassification in the age group XXII, as suggested by the dentalprogression state.

7. Discussion

Elephants of the mammoth lineage appeared in Eurasia duringthe Late Pliocene, about 3.5 million years ago. The taxonomy of theearly forms is still ambiguous, as they are only known from scantyand incomplete samples, but they are recently grouped under thespecific nameMammuthus rumanus (Stef�anescu,1924) (Markov andSpassov, 2003; Lister et al., 2005). The Pleistocene mammoths d

M. meridionalis, M. trogontherii and M. primigenius d constitute anevolutionary line exhibiting anatomical trends as increasing molarhypsodonty and lamellar frequency, enamel thinning, cranialanteroposterior shortening and heightening and changes in bodysize. The main anatomical characters of the mammoths includea high, domed skull, narrow praemaxillary bones, twisted tusks andwide molars with weak enamel folding (Osborn, 1942; Maglio,1973; Lister, 1996a). Another Pleistocene elephant is the straight-tusked E. (Palaeoloxodon) antiquus, which coexisted in EuropewithM. trogontherii andM. primigenius. E. antiquus is distinguishedfrom the mammoths by the presence of a two-domed frontopar-ietal crest in its skull, the weakly curved, untwisted tusks, thepresence of median loxodont enamel sinuses in the molars, and therather thicker molar enamel. In the case of isolated molars,however, the distinction between E. antiquus and M. trogontheriican be difficult, due to their similar hypsodonty and lamellarfrequency. This difficulty is accentuated by the great morphologicaland metrical variation that characterises the elephants in general,a fact that led Soergel (1913) to establish ‘varieties’ of intermediatemorphology in these two species: ‘Elephas trogontherii var. anti-quus’ and ‘Elephas antiquus var. trogontherii’.

The postcranial elements are difficult or impossible to distin-guish, as they are similar in size and morphology. Several authorshave tried to find distinguishing criteria based on the shape and therelative development of the autopodial (mainly astragalus andcalcaneus) facets (e.g. Andrews and Forster Cooper, 1928; Neuville,1946; Trevisan, 1954; Melentis, 1963; Beden and Guérin, 1975).Using these criteria, particularly those reported by Andrews andForster Cooper (1928), the Loussiká postcranial elements aregenerally similar to those of Mammuthus (but not without excep-tionsd see Sections 5.2.6 and 5.2.7). However, there is accumulatingevidence against the validity of such anatomical distinctions: Bedenand Guérin (1975) have observed that the facet morphology changesduring ontogenetic development of individuals that belong toa single species, so they attribute these morphological differences tointraspecific variation. Lister and Stuart (2010) also found that nopublished character can by reliably used to separate Mammuthusfrom Elephas (Palaeoloxodon). Moreover, a recent re-examination ofthe dental material from Megalópolis (unpublished data), a fossilfauna where the criteria of Melentis were based on, as well asgeochemical data (Iliopoulos et al., 2010), have shown that only onespecies, E. antiquus, exists in the sample studied by Melentis.

The proboscidean skeleton excavated at Loussiká was prelimi-narily attributed to E. antiquus (Doukas and Athanassiou, 2003;Athanassiou, 2010), so far the only known MiddleeLate Pleistocene


elephant species in Southern Greece, because of the followingreasons:

(a) The skull vault is damaged (eroded) dorsally in a way thatreveals a horizontal section of the caudally expanded occipitalbone (Fig. 8). During the excavation it was thought that thissection is positioned much higher in the skull and the caudally(at both sides of the external occipital crest) expanded occipitalbone was erroneously interpreted as the two-bulged fronto-parietal crest of E. antiquus.

(b) The lowerm3morphology is characterised by high hypsodonty,fairly folded, rather thick enamel and medium lamellarfrequency, all seen in E. antiquus. Also, the incipiently wornlamellae formawidemedianenamel loopand smaller labial andlingual ones, as it is usually the case in this species. Theappearance of the occlusal surface is broad, but very close to themaximumwidth cited for E. antiquus (Maglio, 1973, Table 18).

During the recent preparation of the skull, it was observed thatthe whole part above the nasal cavity was destroyed before finalburial (a usual damage in fossil elephant skulls), so a frontoparietalcrest or vertex is not preserved. Moreover, the tusk sheaths werefound to run almost parallel to each other and are situated veryclose together. This morphology, in spite of the absence of thevertex and tusks, points towards the genus Mammuthus andexcludes and attribution to the straight-tusked elephant, which hasmarkedly diverging tusk alveoli (Osborn, 1942; Maglio, 1973).

Subsequent observations on two tusk fragments, exhibiting theSchreger pattern came to corroborate the unexpected cranialevidence. The Schreger pattern has been used in the proboscideangenus-level taxonomy, as well as for the forensic discrimination ofivories that come from fossil or extant elephants. The acute anglesmeasured perpendicular to the tusk axis and close to the cement/dentine junction (Fig. 11) are consistent with an attribution toMammuthus, in contrast to Elephas (Palaeoloxodon), which exhibitsobtuse outer Schreger angles (over 93�), similar to the extantgenera (Palombo and Villa, 2001).

The attribution to Mammuthus is also compatible with the molarmorphology, as described in Sections 5.2.3 and 5.2.4, particularly thewide crown, as well as the absence of loxodont sinuses in the enamelloops. The molars, mainly the third ones, have been used extensivelyin mammoth species-level taxonomy, as their morphology directlyreflects the species’ dietary adaptations. Unfortunately, the preser-vation in situ of the third molars at Loussiká does not allow certainparameters to be measured or counted accurately. The high lamellaenumber (about 20e22) and their high hypsodonty are advancedcharacters that preclude the attribution of the skeleton to a lessderived species as M. meridionalis. The latter has a maximum platenumber of 14, and a hypsodonty index less than 165 (Maglio, 1973,Table 30), or even lower (99e146) in specimens from Valdarno, itstype locality (Lister and Stuart, 2010, p. 190). Late forms ofM.meridionalisddatedat the latest EarlyPleistoceneor the transitionto theMiddle Pleistocene and often placed in the separate subspeciesArchidiskodon meridionalis tamanensis Dubrovo, 1963 d showadvanced dental characters (higher hypsodonty, higher lamellarfrequency, thinner enamel d Dubrovo, 1977; Baigusheva and Titov,2010), but again do not reach the high plate number and level ofhypsodonty seen at Loussiká.

On the other hand, the Loussiká dentition is not so progressiveas in the most advanced species of the Mammuthus lineage,M. primigenius. The lamellar frequency of 6.1e6.3 plates per 10 cmof molar length (which may be better attributed to the large molarsize, not to low plate number) is outside the variation ofM. primigenius (more than 6.5 in Maglio, 1973, Table 32; more than7.4 in Lister, 1996a, Fig. 19.4), a species well known for its tightly

packed enamel plates. A related metrical parameter is the enamelthickness, which is much thinner (less than 2.0 mm according toMaglio, 1973 and Lister and Stuart, 2010, p. 191) in M. primigenius,but as high as 3.2 mm in Loussiká m3s. Another distinguishingfeature between the Loussiká mammoth and M. primigenius is thelarger size of the former. Although there are large-sized woollymammoths, their maximal dental size (upper and lower molarlength less than 285 and 320 mm respectively; Maglio, 1973,Table 32) is clearly smaller than the estimated for Loussiká.

Comparing the Loussiká dental specimens to the sample fromSüßenborn, Germany, the type locality of M. trogontherii, it isobserved that they fallwell into the rangeof variation of this species.Guenther (1969) counts 14e21.5plates inbothM3sandm3s, placingthe Loussiká molars among the maximal values of Süßenborn. Thesame is true for crown height (maximal values 209mmand 172mmfor M3 and m3 respectively), though these parameters are roughlyestimated on the Loussiká molars (especially the lowers). The sameauthor reportsM3 length andwidth ranges of 235e375 and 75e120respectively, and m3 length and width ranges of 260e390 and75e115 respectively. Again the studied specimens are closer to themaximum values. Maglio (1973, Table 31) reports similar ranges(thoughwith lower maxima, especially for the parameter of length)and also gives a lamellar frequency range of 5.0e8.2 and 5.0e7.2 forM3 and m3 respectively, with Loussiká molars placed very close tothe mean values. Other important dental samples of typicalM. trogontherii are those from Mosbach, Germany, and Tiraspol,Moldova; their metrical characters do not differentiate essentiallyfrom Süßenborn (Guenther, 1969; Dubrovo, 1975; Table 2). Incomparison to the recently studied West Runton mammoth (Listerand Stuart, 2010), the Loussiká dental material is slightly dimen-sionally smaller (as it is also the case with the postcranial skeleton)and has similar plate number and frequency.

The lateMiddle Pleistocenemammoths are oftenmore advancedin dental characters (higher plate number, higher lamellarfrequency, thinner enamel etc.) and tend to be smaller in body size.They have been placed in a distinct subspecies, M. trogontherii cho-saricus Dubrovo, 1966 (often given specific rank), in a distinctspecies,M. intermedius (Jourdan,1861) (Labe andGuérin, 2005), or inM. primigenius; their taxonomic status still remains ambiguous(Listeret al., 2005; PalomboandFerretti, 2005). Good samples of thismorphotype have been described from the former Soviet Union,Germany and Italy (Siegfried, 1956; Ambrosetti, 1964; Palombo,1972; Dubrovo, 1977; Kotsakis et al., 1978). The Loussikámammoth differs from these samples by its generally larger dentaland body size, lower lamellar frequency and thicker enamel.

The metrical comparison shows, in conclusion, that the partialskeleton from Loussiká is dentally more advanced thanM. meridionalis, less derived and larger than M. primigenius, andcorresponds well to the upper range of the dental sample fromSüßenborn, type locality ofM. trogontherii.

8. BiostratigraphyePalaeoecology

The site of Loussiká did not yield any biochronologic evidence,as the accompanying fauna is very poor and atypical taxonomically.The biochronology of the locality has to be based exclusively on themammothfind. Theearliest remainsofM. trogontherii inEuropedatefrom the Early to Middle Pleistocene transition, probably over-lapping chronologically with the latest M. meridionalis. The typicalM. trogontherii morphotype persisted through the early MiddlePleistocene, with smaller, ‘chosaricus’-type forms appearing duringthe later Middle Pleistocene. The good metrical and morphologiccorrespondence of the Loussikámammothwith the typematerial ofM. trogontherii and other samples referred to the typical form of thespecies can justify the stratigraphic placing of the site in the lower


Middle Pleistocene, despite its reduced body size in comparisonwithother skeletonsdated to this time span. The rather small staturedifference can be well attributed to intraspecific variation, particu-larly when the geographic distance between the compared samplesis evidently long.Moreover, the Loussiká skeleton comprises severalnon-fused epiphyses, clearly indicating that this individual was stillgrowing at the time of death. Also the retention of an ancestralcharacter, as the thick enamel, if not of local trophic significance, canbe considered as indicative of an older geochronologic age. There-fore, an early Middle Pleistocene age is more probable for Loussiká.

The typical M. trogontherii from Central-East Europe is consid-ered as a steppe dweller d thus its vernacular name ‘steppemammoth’d as it is well adapted to open landscapes and aridity. Itinitiates an evolutionary line that ended to the extremely speci-alised, cold-adapted grazer, the woolly mammoth. Recent palae-oecological data referring to the environment of the West Runtonmammoth (Stuart and Larkin, 2010) have shown thatM. trogontheriimay be associated with a temperate forest, but this is still a soleindication referring in a northern regionwith anyway cool climate.

The site of Loussiká has not produced any palaeoecologicalevidence, which could be used for the reconstruction of the palae-oenvironment. Nonetheless, the nearby lacustrine basin of Mega-lópolis, situated in central Peloponnese, has yielded a goodpalaeoenvironmental record for the Middle Pleistocene, mainlybased on pollen (Okuda et al., 2002). The basin is filled with cyclicbedsof lignite, depositedduring interglacials, anddetritus, depositedduring the glacials. All the lignite seamsyieldpollenof temperate oakforest (as well as an ‘interglacial’ mammal fauna rich in E. antiquus,Hippopotamus sp. and deer), whereas the prevailing plant genus inthe detrital beds is the shrub Artemisia, a strong indication of semi-arid steppe. The Megalópolis palaeoenvironmental record suggeststhat cold and arid steppic conditions did prevail during glacialperiods in a quite southern region as central Peloponnese. This pieceof evidence fits well with the ‘unexpected’ discovery of a cold-adapted animal at Loussiká. The presence of M. trogontherii in Pelo-ponnesecan, therefore, begeochronologically spottedmorepreciselyto a cold episode of the early Middle Pleistocene, as those of theMarine Isotope Stages (MIS) 14, 16 or 18, the climatic conditions ofwhich would allow the southern expansion of the steppemammothpopulation.

Fig. 21. Map of the Mediterranean region showing the geographical distribution of the Mamsouthernmost sites. The question mark denotes samples of problematic taxonomic determLenardi�c (1994), van der Made and Mazo (2001), Palombo and Ferretti (2005), Albayrak (2NASA World Wind. The mammoth figure in taken from Osborn (1942, p. 1052).

9. Palaeobiogeographic implications

Despite its apparent expansion during the glacials,M. trogontheriiseems to have beenmuch less abundant in regions of less continentalclimatic conditions, as theWestern and Southern Europe, away fromits core region in Eastern and Central Europe. In MediterraneanEurope inparticular (Iberian, Italian andBalkanpeninsulas) it is quiteuncommon, usually identified on the presence of isolated molarswith trogontherioid morphology. In Italy, the species is totallyunknown southern of Rome (nonetheless the more derived lateMiddle Pleistocene mammoths expanded to the South till Capri,40.5�N) (Palombo and Ferretti, 2005). In Spain, though also rare, it isreported to have reached as south as the GuadixeBaza basin (about37.5�N) (van derMade andMazo, 2001). The Balkan Peninsula is alsopoor in M. trogontherii. Radulesco et al. (1965) describe dentalmaterial from the Brasov area in Romania, while Lenardi�c (1994)studied an advanced ‘chosaricus’-like skull from Croatia. A recentimportant discovery in Eastern Serbia (at Kostolac) refers toa completeM. trogontherii skeleton (Korac et al., 2010).

The presence of the species in Greece was already reported byPsarianos (1958) and Marinos (1964) in the north of the country(localities Sotír in Western Macedonia and Phílippi in EasternMacedonia) based on a total of three isolated molars. The poorphotographic documentation (without a metrical scale) and theabsence of any metrical data do not allow a re-evaluation of thistaxonomic attribution. The Philippi specimens are morphologicallyvery similar toM. trogontherii, though they could also be referred toan early M. primigenius (a species also reported from Phílippi). Thespecimen from Sotír, presumably a left m3, exhibits weak enamelplication and virtually absent loxodont sinus, but the crown israther narrow for a mammoth. More recent material coming fromthe same locality (a sand pit) is also referred to Mammuthus(Velitzelos and Schneider, 1973). However, the provided photo-graphic and metrical data (narrow crowns, wrinkled enamel etc.)point clearly to E. antiquus, as is also correctly observed by Tsoukalaet al. (2010). In conclusion, the only until now plausible evidence onthe presence ofM. trogontherii in Greece comes from the peat basinof Phílippi, situated in the NE of the country. The studied skeletonfrom Loussiká expands considerably to the south the species’biogeographical distribution in Greece.

muthus trogontherii-bearing localities according to available data. Loussiká is among theination. Data according to Marinos (1964), Radulesco et al. (1965), Horowitz (1977),009), Mol and Lacombat (2009), Korac et al. (2010). Composite satellite image source:


Dental finds attributable toM. trogontherii are also reported fromother countries of the EasternMediterranean region. Albayrak (2009)cites four sites inTurkey, one ofwhich is the typical locality of Elephasarmeniacus Falconer,1857, a synonym ofM. trogontherii. Another site,Dursunlu, located in SW Turkey near Konya, is almost as far south asLoussiká. A few other dental specimens from Syria and Israel (local-ities Latamne, Evron, Benot Ya’akov) are referred to M. trogontherii(Horowitz, 1977, based mainly on the studies of Hooijer). Maglio(1973) considered these specimens as Elephas namadicus(¼E. antiquus), but Hooijer’s specific determinations have beenaccepted by other scholars (Adam, 1988, pp. 15e16; Guérin et al.,1993). In a later review of the Levantine proboscidean remainsTchernov and Shoshani (1996) consider these samples as taxonomi-cally problematic, but they do assign some dental fragments fromLatamne toM. trogontherii (Tchernov and Shoshani,1996, Table 21.1).Considering these controversial opinions and the generally frag-mentary state of the available material, it seems that the presence ofM. trogontherii in the Levant is rather ambiguous. Consequently thebiogeographic distribution of the species can be securely extended tothe south to Southern Turkey and Southern Greece in EasternMediterranean (about 38�N), and Southern Spain (Granada) in theWestern Mediterranean (about 37.5�N) (Fig. 21).

10. Conclusions

Given the rarity of associated skeletons referred to M. trogon-therii, the Loussiká skeleton is an important find, preservingmost ofthe cranial and postcranial elements of this mammoth. It is themost complete and well-preserved find of this species in theEastern Mediterranean region and one of the few known skeletonsin Europe. The study of its anatomical and metrical charactersshowed that the skeleton belongs to amale, about 45 years old, thatweighted about 8 t and stood in life about 3.80m high. Dentally, it isvery similar to the type sample of the species.

M. trogontherii is extremely rare in Greece and was previouslyunknown in the southern part of the country. Its presence extendsconsiderably to the South the species’ known geographical distri-bution in the Balkan Peninsula.

Based on the specific determination of the find, and taking intoaccount its particular anatomical characters, the fossiliferouscontinental deposits of Loussiká can be dated to the early MiddlePleistocene, quite probably to a cold stage.

Acknowledgements

The Loussiká excavation was financed by the Hellenic Ministryof Culture (Ephorate of PalaeoanthropologyeSpeleology, Athens,and ST’ Ephorate of Prehistoric and Classical Antiquities, Patras);the Municipality of Olenía contributed to the excavation teamwiththree workers. The team consisted of A. Darlas (archaeologist,director), A, Athanassiou (geologistepalaeontologist), P. Poly-doropoulos (preparator), D. Bouzas (geologist), L. Stavropoulou(designer), S. Stamatopoulos, G. Martzaklis, A. Chrysikopoulos, N.Spiliopoulos and G. Tsironis (workers). The finds were prepared inthe laboratory of the Ephorate of PalaeoanthropologyeSpeleologyby P. Polydoropoulos, P. Ghioni, V. Papamikou and V. Klaridi. Thegeologist E. Ypsilanti helped with the measurements. A recent re-examination of the Loussiká partial skeleton (currently in thecollection of the Archaeological Museum of Patras) was facilitatedby E. Kollia and A. Paraskevopoulos, archaeologists of the ST’Ephorate of Prehistoric and Classical Antiquities. M. Golfinopoulou(ST’ Ephorate of Prehistoric and Classical Antiquities) provideda copy of one of the excavation plans. G. Iliopoulos (University ofPatras) offered hospitality during my stay in Patras and provideduseful comments and discussions on the elephant osteology and

taphonomy. An anonymous (Bavarian State Collection for Palae-ontology and Geology) and Yasemin Tulu (Calvert Marine Museum)greatly helped in the search of hard to find publications. Twoanonymous reviewers are especially thanked for critically readingthe manuscript and providing useful comments.

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