Lister Et Al., 2012 - M. Trogontherii Drmno, Serbia

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Quaternary International 276-277 (2012) 129e144

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Quaternary International

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A skeleton of ‘steppe’ mammoth (Mammuthus trogontherii (Pohlig)) from Drmno,near Kostolac, Serbia

Adrian M. Lister a,*, Vesna Dimitrijevi�c b, Zoran Markovi�c c, Slobodan Kne�zevi�c d, Dick Mol e

a Palaeontology Department, The Natural History Museum, Cromwell Road, London SW7 5BD, UKb Laboratory of Bioarchaeology, Department of Archaeology, Faculty of Philosophy, University of Belgrade, SerbiacNatural History Museum, Belgrade, SerbiadDepartment of Geology, Faculty of Mining and Geology, University of Belgrade, SerbiaeNatural History Museum Rotterdam, c/o Gudumholm 41, 2133 HG Hoofddorp, The Netherlands

a r t i c l e i n f o

Article history:Available online 16 March 2012

* Corresponding author.E-mail address: [email protected] (A.M. Lister).

1040-6182/$ e see front matter � 2012 Elsevier Ltd adoi:10.1016/j.quaint.2012.03.021

a b s t r a c t

The Kostolac mammoth was discovered in 2009 in Pleistocene deposits adjacent to the Drmno open-castlignite mine in the Serbian Danube Basin. On the basis of cranial and dental features, the individual isidentified as the so-called ‘steppe’ mammoth, Mammuthus trogontherii. The remains are those of an oldmale of estimated age around 62 years, and comprise one of the most complete and best-preservedknown skeletons of this species, and the first from the region. Skeletal height is estimated as aroundfour metres, and body mass 9.5 t. The excellent preservation of the skeleton provides new informationabout the osteology of M. trogontherii, an evolutionary intermediate between the better-known ancestralmammoth Mammuthus meridionalis and woolly mammoth Mammuthus primigenius. The find is alsoremarkable for the articulated condition of the skeleton, the animal occupying a crouching posture whichis probably little-altered from its original death position. This and the depositional environment of theskeleton, a broad, fast-flowing river, suggest that the animal died in relatively shallow water and wasvery rapidly buried in river sediments. Based on the known European record of typical (large-sized)M. trogontherii of this kind, the age of the Kostolac skeleton and its enclosing sediments is between 1.0and 0.4 Ma.

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1. Introduction

The Kostolac mammoth was discovered on May 27, 2009, in anopen lignite mine adjacent to the famous Roman site of Vimina-cium, approximately 90 km east of Belgrade, Serbia (Kora�c et al.,2010; Fig. 1). The area, within the Danube basin, lies on theboundary between the Balkan Peninsula and the Pannonian plain.The Kostolac coal basin, due east from the Velika Morava river, hasbeen exploited since 1870. The mammoth skeleton was discoveredon the north-east rim of the Drmno pit, the youngest and largest ofthe open pits, near the mouth of the Mlava River, on low hilly landthat rises over the flat landscape of the alluvial plains. On the westside, above the left bank of the Mlava, the high ground of Pozar-eva�cka greda spreads in a SSEeNNW direction. The mammoth wasfound at 44�4305000 N, 21�1402100 E, Gauss-Krueger grid reference4 954 024, 7 518 937, at a height of approximately 58m above meansea level, and a depth of 27 m below the modern surface.

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The skeleton, remarkable for its completeness and excellentstate of preservation, is one of very few known skeletons of theMiddle Pleistocene ‘steppe’ mammoth, Mammuthus trogontherii.This species occupies an important position in the evolution of thegenus Mammuthus, as it is considered to be an intermediatebetween the widespread, and better-known, Early Pleistocenespecies Mammuthus meridionalis, and the familiar woollymammoth, Mammuthus primigenius (Lister et al., 2005). Anyinformation on its anatomy and adaptations is therefore of interest.The Kostolac mammoth is also the first skeleton of M. trogontheriifrom the Mediterranean Basin, and its taphonomic situation isremarkable, strongly suggesting that it has been preserved in deathposition.

2. Regional geological setting

The geology of the Kostolac basin comprises sediments of LateMiocene to Quaternary age. Late Miocene sediments formed in aformer gulf of the Paratethys at the southern edge of the Pannonianprovince. The Pontian of the Paratethys is the chronostratigraphic

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Fig. 1. Geographical position of the Kostolac basin, and location of mammoth site at the “Drmno” open pit coal mine.

A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144130

equivalent of the Messinian. Its mollusc fauna includes freshwaterforms such as unionids and Viviparus, indicating local freshening ofenvironment due to inflow of river water from the land. This type offacies is known in the literature as the “Kostolac brackish” Pontian(Stevanovi�c, 1951, 1977, 1990). Here are found deposits of softbrown coal (lignite) and layers of silt, marly clays, sands andgravelly sands. Coal is exploited in quarries near Kostolac town andthe villages of �Cirikovac, Klenovnik and Drmno. To the south, nearthe village of Poljane on the heights of Po�zareva�cka greda, LateUpper Miocene deposits of Pannonian age are found, with lesserdeposits of coal (Milo�sevi�c and Mileti�c, 1975).

Quaternary deposits cover all the land in a wide area aroundDrmno. They include Pleistocene and Holocene sediments. Pleis-tocene features are represented by pre-loess proluvialedeluvialdeposits of the Kli�cevac Formation, loess, and aeolian sand (Raki�c,1978).

The Kli�cevac Formation, named after the settlement of Kli�cevac,is located on the right bank of the Danube ENE of Drmno. It liesuncomformably over Neogene sediments and below Pleistoceneloess and other Quaternary features. This formation consists ofloessoid sandy-clay silt and silty sands, gravels, sandy limestones,sandstones and tufa. Themost widespread are sediments depositedon hills with differing slopes, in the form of a polygenetic curtain,although some were deposited in a subaquatic environment.Subaquatic sediments discovered in the deepest erosion channels

in the areas of Re�cice and Kli�cevac are formed from brown ironsands with gravel and silt lenses, in which ostracods and a mixedfauna of freshwater and terrestrial molluscs (Pisidium amnicum,Planorbis sp., Unio sp., Clausilia sp., etc) are found (Raki�c, 1978).

Aeolian sediments are themost widespread Pleistocene featuresin the broad area of Drmno. Loess covers thewhole land surface andis represented by terrestrial andwetland loess. Along the right bankof the Danube near Kli�cevac there are deposits of aeolian sand.

Holocene sediments are various. Alluvial deposits in river valleysof theDanube and its tributary theMlava have the greatest thickness.They comprise different facies including the sediments of floods, stillwaters and marshes. Elsewhere, contemporary deluvial anddeluvialeproluvial deposits occur, as well as anthropogenic depositssuch as littering and large areas of dug material from open pits.

3. Stratigraphic context of the mammoth skeleton

In the geological section at the mammoth location, Neogene andPleistocene deposits are seen (Fig. 2). The Neogene is representedby coal deposits of Pontian age, developed as the brackish Kostolactype. These layers, below the mammoth, are folded in a lowsyncline, different in structure from the upper, horizontal Pleisto-cene sediments.

The coal deposits, around 50 m in depth, consist of gray fine-grained mica and silt sands with thin intercalations of somewhat

Fig. 2. Stratigraphic column of the mammoth site at the “Drmno” open pit coal mine.1e3: Neogene, Late Upper Miocene (Pontian). 1. Coal e lignite, coal clays; 2. Gray siltysand with intercalation of grayeyellow sand; 3. Orange and brown sand with cross-bedded stratification. 4e5: Quaternary, Pleistocene. 4. Alluvial sediments. 4a:Riverbed sediments: gravel, yellow sand, gray sandy gravel, and clays, with mammothremains at 58 m AMSL; 4b: Wetland sediments: silts, silty sands with mollusc fauna(Unio sp., Planorbis sp., etc.), micromammals, elephantid, Megaloceros giganteus. 5.Loess sediments. 5a: Marshy loess: gray-brownish clayey silts; 5b: Land loess: gray-yellow silt. 6: Holocene. Anthropogenic deposits (archaeological layers and humus).

A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 131

harder grayeyellow sands. These are deltaic sediments formed ina coastal area of the Paratethys gulf where a river emptied from thesouth.

Younger deposits, lying directly beneath the mammoth layer,are formed of orange and brown medium-grained and coarse-grained sands with cross-bedded stratification. These are sedi-ments of a deltaic front formed in a phase of strong river flow andwithdrawal of the Paratethys’ coastal line. Samples from thesesediments have yielded rodent and insectivore remains includingOccitanomys sp., Kowalskia sp., Desmanodon sp. and Sorex sp.

The overlying Pleistocene sediments are divided into a lowercomplex of river sediments, and an upper complex of loess. Thelower complex comprises two horizons, with a total thickness inthe studied section of about 10 m.

The skeleton of themammoth was found in the lower horizon ofthe lower complex, which lies unconformably over the Pontianpalaeorelief (Fig. 2). It is formed from yellow, brownish-yellow andgray gravels and sandy gravels, and brownish-reddish, yellow andgray dusty sands, silt lenses and clays. The gravels are slightlyrounded and contain quartz, hornstone and metamorphite, as wellas smaller fragments of Mesozoic sandstones, limestones andigneous rocks. Sands and gravels are cross-bedded in places. Basedon lithostratigraphic characteristics it can be concluded that thesesediments were deposited by a river trending SSEeNNW, withvarying but often strong flow. The river valley had a substantialbreadth of approximately 600 m. The mineral composition of theclasts suggests that the Pleistocene river flowed from the zone ofSerbian-Macedonian crystal mass to the NNW. The skeleton(Figs. 3e5) lay close to the edge of the channel, oriented NW (head)to SE (tail).

The upper horizon is represented by a flood deposit. Thiscomprises sandy clays and gray-brownish silts, with lenses ofoxbow-lake dark gray silty clays. Fossils of molluscs, micro-mammals and large mammals are found in these sediments.Among the molluscs species of Unio and Planorbis predominate.The rodents comprise Spermophilus sp., Cricetus sp. and Pitymys sp.These animals commonly inhabited the steppe areas of present-day Serbia in the Quaternary. Large mammal remains includea fragmentary elephantid molar and a radius and ulna of the giantdeer Megaloceros giganteus.

The upper complex of Pleistocene sediments comprises loess,and is also divided into two horizons. The older horizon is awetlandloessoid sediment with a terrestrial and freshwater mollusc fauna.The younger horizon includes sediments of land-slope loess. Thethickness of the loess complex varies from 10 to 15 m.

Anthropogenic Holocene deposits are found at the surface of thesection (Fig. 3a).

4. Excavation and preservation of the skeleton

The skeleton (Fig. 3) was unearthed by a large excavation digger.As a result, the left side of the skull was destroyed and the bones ofthe left front leg damaged. The digger bisected the skull longitu-dinally and pulled away its left side, together with the left tusk,lower jaw and part of the right tusk. The right half of the skullremains whole and in place, although its forehead is cracked andsunken (Fig. 6a). The middle part of the right tusk, up to the alve-olus, was detached and broken into pieces. The distal part of theright tusk remains in situ (Fig. 3b). The left front leg bones weredetached from their position and damaged.

When the digger stopped, many fragments of the tusks and skullthat had been detached were collected but it was not possible toassemble the fragments. Upper and lower third molars were saved,as well as fragments of the bones of the left front leg: distal scapula,proximal radius and proximal ulna. Subsequently, the backbone,the bones of the right front leg and both hind legs were carefullyuncovered in situ. The bones of the legs are folded beneath thebackbone, scapula and pelvic girdles (Fig. 7). The excavation wasperformed by the archaeological team of the Roman site of Vim-inacium and experts from the Natural History Museum in Belgrade.Immediately after excavation, thorough conservation of the skel-etonwas undertaken and a protective structure constructed over it.Due to these precautions the skeleton survived, as it was onunstable ground and the bones were prone to fast disintegration.Figs. 3a,b and 5 were taken during the course of the excavation;Figs. 7e9, 12 and 17 at the end of the excavation; and Figs. 6, 10, 11,13e15 after the cover had been constructed, conservation of theskeleton completed and the skull and tusk reconstructed.

Fig. 3. a: skeleton in situ, view from SW (photo courtesy of Viminacium archaeological team). Stratified river sediments are seen in the section behind the skeleton. In thebackground to the left, a profile of loessic deposits is seen, with Holocene cover on top. This includes anthropogenic deposits with brick graves of the Roman necropolis ofViminacium. b: Close-up of the mammoth skeleton. The isolated right tusk-tip is seen to the left.


The skull preserves, on the right side, the zygoma, auditoryopening, orbital region, and the lateral wall of the braincase (Figs. 6and 8; Table 1). The right side of the nare (trunk attachment) andfrontal are crushed and buckled downwards. Only parts of the righttusk are preserved: a fragment of the proximal end remaining inthe alveolus, and a separate distal portion, some 85 cm in lengthand ending very close to the original tip (Fig. 9). This portionappears to be in life position relative to the skull (Fig. 3b); if so, theoriginal length of the tusk, from the alveolus to the distal tip, can bemeasured as approximately 260 cm. Other pieces of tusk werefound loose. Of the mandible only one articular condyle and thesymphysis region are preserved.

Apart from the tail, the vertebral column is complete and, withsome disarticulation and movement, in anatomical order (Figs. 7and 10). The atlas is pressed around the occipital condyles of theskull. The axis and following five cervical vertebrae are in line

behind it. They are inclined downward at some 45� to the line inwhich the skull and the distal end of the right tusk lay. The first andsecond thoracic vertebrae continue the row. The left and right firstribs are articulated between the last cervical and first thoracicvertebrae. The proximal part of the left second rib is articulatedbetween the first and second thoracics. This rib lacks part of itsbody, but its middle part is in situ. The following four thoracicvertebrae, from the third up to the sixth, are displaced, as a group,to the right (Figs. 7 and 10). The remaining thoracics are all inarticulation. The middle thoracics (seventh up to tenth) lie ina straight line that is parallel to the line in which the skull and thefragment of the tusk lay. The posterior thoracics, lumbar and sacralparts of the backbone are inclined from that line and bent to theright. The caudal vertebrae were displaced during excavation(Fig. 11). A total of 13 right and 11 left ribs were uncovered duringthe excavation. Two ribs were found several metres apart to the

Fig. 4. Field drawing of the mammoth skeleton in situ (drawing by Dragana Petrovic).


right side of the skeleton, and four ribs several metres away from itsrear side (Figs. 4 and 12). The sternum is preserved, showing thepresternum and mesosternum fused together. The sacrum hasfallen beneath the pelvis.

The right scapula is significantly rotated in such away that it liesnext to the skull (Fig. 8). The right humerus is perfectly preserved,

Fig. 5. Skeleton in situ, view from NW during excavation, with cranium and rightscapula in the foreground. Photo courtesy of Viminacium archaeological team.

with the articulated ulna and radius bent beneath it (Fig. 13). Thebones of the right front foot are probably beneath the distal end ofthe ulna and radius, buried deep in the sediment. It was notpossible to excavate deeper beneath the skeleton and uncover thecarpals and further bones of the distal leg, since this would mostprobably cause the collapse of the skeleton.

The bones of both hind legs are well preserved. There isremarkable symmetry between the position of the left and rightlimb bones (Fig. 7). The caputs of the femora had fallen out ofthe acetabula, but remained just beneath them. The femora aredirected forwards and diverge somewhat laterally from the overallmidline of the skeleton. Beneath them, the tibiae and fibulae areflexed backwards and lie approximately parallel to the main axis ofthe skeleton. The right patella is still in position in the knee joint;the left one fell out in the course of the excavation. The calcaneaand astragali of both feet are visible (Fig. 7). In the right foot some ofthe smaller tarsal bones are also visible, together with at least eightsesamoid bones, each 40e45 mm in diameter. The metatarsal andfirst phalange of the fifth digit are visible, as well as a furthermetatarsal and a first phalange. In the left foot, the navicular isvisible, together with two metatarsals and at least six sesamoids.Since these elements are well preserved and perfectly articulated itis expected that further metatarsals and phalanges remain buriedin the sediment.

5. Taphonomy of the skeleton

The skeleton lies in a coarse, yellow river sand with variably-shaded bands of finer sand, coarser sand and fine gravel, some-times cross-bedded (Figs. 2 and 3). It is largely complete, with themajority of bones articulated and positioned as in the live animal.The bones of the hind legs, in particular, impressively recall theposture of a living animal. It is likely that the animal was kneeling atthe find-spot, and eventually died in that posture. This indicatesthat the skeleton is in primary position and that the body wasburied under the sandy sediments shortly after the animal died.The conclusion of rapid burial is strengthened by the observation

Fig. 6. a: Cranium of the Kostolac mammoth in dorsal view, after conservation and restoration. Left side of the skull is restored according to the preserved right side. b: Right lateralclose-up of cheek region of the Kostolac mammoth cranium, showing jugal bone (j) and narrow infraorbital process (i).


that the skeleton was not disturbed or damaged by predators,which would undoubtedly have been attracted by the dead body ifit had been exposed for any period of time. The skeleton wasprobably complete before its discovery and excavation.

Although the animal was of an advanced age, a survey of theskeleton revealed no evident signs of pathology. No significantarthritic growth or exostosis was seen on any bone, although theseare common in skeletons of old elephantid individuals. Thissuggests that the animal was in good health. The posterior articularfacets of the fourth thoracic vertebra (T4) (Fig. 14) show strong left-right asymmetry, although in T3 and T6 they are roughlysymmetrical, and in all other vertebrae are not visible. This asym-metry is, however, very common in elephantids (e.g. Lister, 2009,Fig. 17B), and there is no reason to consider it pathological. Thevertebral spines are straight, and there are no perforations at theirbase as seen in some Mammuthus skeletons (e.g. Lister, 2009,Fig. 17A). Of course, the animal might have suffered from a diseasethat does not leave pathological traces on bones, so illness cannotbe completely excluded as possible cause of death.

A second possibility to be considered is that the animal died ofstarvation due to the advanced state of wear of its molars. Each

Fig. 7. Skeleton in situ, view from NE. Pelvis and hind legs in the foreground. Note thesymmetrical position of the left and right hind limb bones. Below the distal tibiaeproximal tarsal bones are seen, while further tarsal bones disappear into the sediment.Photo courtesy of Viminacium archaeological team.

lower molar, in particular, has only about half of its enamel platesleft, and they are obliquely worn and narrowed because of theirproximity to the crown base (Fig. 15a). That elephants do die frominability to feed with worn-out teeth is well-known (Shoshani,1991). An animal the size of the Kostolac mammoth would haveneeded well in excess of the 200 kg of fresh food per day typical ofAfrican elephants today, which may chew for 15 or more hours perday (Sukumar, 2003, p.197). The lower molars are at the 29th of 30eruption and wear stages observed by Laws (1966) for Loxodontaafricana e implying that few African elephants survive beyond thisstage. However, the upper molars of the Kostolac mammoth stillretained a significant proportion of their crown, and the occlusalsurfaces of the upper and lowermolars are of similar dimensions. Inthe absence of quantitative published data on the dental wear stageat which elephants are threatened with starvation, it is difficult toreach a firm conclusion on this possibility.

Modern elephants that are severely injured, diseased or starvingfrequently go to water, where they commonly die. In either of theabove scenarios, the mammoth may have stood in shallow water inthe river, and then collapsed to its knees before expiring. Theillustrations of numerous elephant carcasses that died in Africa dueto drought (Beard, 2000) show in many cases that the animal diedin a crouching position with its legs folded under it, very similar tothe situation in the Kostolac mammoth.

Fig. 8. Close-up of the cranium and right scapula, viewed anteriorly and to the right.Photo courtesy of Viminacium archaeological team.

Table 1Measurements of cranium and mandible from Kostolac (mm). Because of breakage to the left side of the cranium, width measures were estimated by doubling the dimensionfrom the midline (estimated from the midline of the axis vertebra) to the relevant point on the right side of the cranium. Abbreviations in brackets follow Beden (1979), plus:m-l, medio-lateral; d-v, dorso-ventral.

Length ofcranium fromback of occiputto front ofpreorbital rim

Length fromfront of auditoryopening tolowest pointinside orbit (Ab)

Height of orbit,measured onits inside (Ac)

Internalheight ofnasalopening (Ld)

Internal widthof nasalopening (Ih)

Width ofcraniumaroundzygoma (Iq)

Width ofcraniumaroundocciput (Ia)

Minimalwidthof braincase

Minimumdepth (d-v)of zygomaticarch

Minimumwidth (m-l)of zygomaticarch

Diametersof mandibularcondyle

850 417 152 197 600 900 1080 440 64 30 118 � 89


A final possibility is that the animal was in good condition butdied due to a mishap in the river itself. Living elephants commonlyfrequent water and are good swimmers. It is likely that M. trogon-therii were similarly capable of walking or swimming through thecurrent of a river. This could have endangered the mammoth inseveral ways, for example if it became embroiled in a whirlpool, orif the mammoth fell through the ice in winter. The animal mighthave struggled against the water flow and/or soft substratum until,exhausted, it gave up the struggle, knelt down and eventually died.Sukumar (2003, p. 266) lists ‘drowning in swiftly flowing water’ asone of the causes of death in living elephants.

Fig. 9. The terminal portion of the right tusk, viewed from the direction of the skel-eton. Photo courtesy of Viminacium archaeological team.

Whatever the reason for the mammoth’s demise, the posture,articulation and excellent condition of the preserved remainsindicate that it was buried rapidly, while the skeleton was still heldtogether by flesh. The strong river flow indicated by the enclosinglithology suggests that the river was carrying substantial quantitiesof sand and gravel, and it is likely that the carcass itself acted asa sediment trap, hastening its own burial.

Movement and slight dispersal of the bones probably happenedduring the process of burial while the flesh was disintegrating,influenced by the water flow around the massive dead body. Somemovement may also have occurred post-depositionally, i.e. after ithad been covered in wet sand. The depression and cracking of thefront of the skull may have happened due to trampling by othermammoths, or post-depositional sediment movement. The post-cranial skeleton mainly shows disturbance of the anterior elementsof the skeleton, namely the scapula, ribs and vertebrae. The rightscapula is rotated and displaced anteriorly, presumbly due to waterand sediment flow after sufficient soft-tissue deterioration. Thethird to sixth thoracic vertebrae are displaced sideways from themain line of the backbone, probably as a consequence of pressureon this region of the back during the process of bodily decompo-sition. The hind part of the backbone is inclined to the right fromthe line in which lay the skull and the front part of the skeleton, atan angle of approximately 60�. Displacement of bones mainlyaffected the ribs, probably loosened by the collapse and subsequentinclination of the backbone, and thenmoved by river flow. The hindlimbs are relatively undisturbed, but the caputs of the femora hadfallen out of the acetabula, a consequence of the weight of thebones pulling them down after decomposition of the flesh anddeterioration of the ligaments that held the joint. These processeswould have been aided by movement of the soft substratum.

Beside the right side of the skeleton, close to its front leg bones,twomore animal bones were found: a deer ulna and a horse radius.

Fig. 10. The anterior part of the vertebral column, viewed from the left. The cervicalvertebrae are seen to the left of the image, followed by the thoracic vertebrae in threesharply disjointed and differently-oriented groups. Photo: Hans Wildschut.

Fig. 11. Four caudal (tail) vertebrae found in articulation. Photo: Hans Wildschut.

Fig. 12. The skeleton during excavation, viewed from behind. The displacement ofseveral ribs posteriorly and to the left is clear. Photo courtesy of Viminacium archae-ological team.

Fig. 13. The bones of the right leg, showing the radius and ulna folded preciselybeneath the humerus. Photo: Hans Wildschut.


By its size and morphology the deer ulna is identified as red deer,Cervus elaphus, while for the horse radius only a generic identifi-cation (Equus sp.) is possible. To check whether these bones indi-cated further skeletal remains at the site, the excavation area wasenlarged several metres to the north-east, i.e. along the right side ofthe mammoth skeleton. No other bones were found. These isolatedbones must represent parts of disarticulated carcasses that hadbeen transported during the same sedimentary event thatpreserved the mammoth. Conceivably their entrapment in thatposition was related to the current flow and/or sedimentationaccumulating around the mammoth carcass.

A year before the skeleton was discovered, on August 4th 2008,a mammoth tibiawas found at Kostolac, at a depth of 26m from thesurface, probably in a similar level to the skeleton discovered later.The state of preservation is similar to that of the mammoth skel-eton. There are mineral particles attached to the surface of thebone, indicating that it was buried in sand. The tibia belonged to animmature animal, judging by its unfused epiphyses, yet it is of largesize e total length 850 mm, identical to the tibiae of the skeleton.This would be conformable with M. trogontherii and suggests thatthe discovered skeleton might not be the only one buried andpreserved in the sand, and that future excavations in the Kostolacopen mine might reveal more finds.

6. Morphometric and taxonomic study

6.1. Taxonomic identity of the mammoth

The skull and tusks of the Kostolac individual are very incom-pletely preserved, yet they show features suggesting generic

determination. In particular, the long and parallel tusk sheaths(premaxillaries), and the narrow forehead between the temporallines (Fig. 6a), are typical ofMammuthus and exclude Palaeoloxodon.The cranial vertex, although damaged, appears to show a singlepeak characteristic of Mammuthus, rather than the double peak ofPalaeoloxodon. Further taxonomic characters are provided by themolar teeth.

The crowns of the left and right upper, and left lower, molars arepreserved almost in their entirety (Fig. 15), as well as the anteriorand posterior portions of the right lower molar with a breakinbetween. Several features indicate clearly that these are the third(last) molars. First, the tapering form at the posterior end of thecrown. Second, in the lower molars, a significant length of crown is

Fig. 14. Fourth thoracic vertebra (T4), in posterior view, showing the vertebral canaland base of the thoracic spine. The strong lefteright asymmetry between the posteriorarticular facets is clear.

Fig. 15. Molar teeth of the Kostolac mammoth. a, c: Left lower third molar in occlusal and mview. Anterior is to the left. Scales 5 cm.


preserved despite the tooth being in an advanced state of wear; thisis possible only in M3 because forward movement of the tooth hadceased. Finally, the presence of a single, curtain-like longitudinalroot is typical of M3 in advanced wear.

The two candidate genera in the European Pleistocene arePalaeoloxodon and Mammuthus. The Kostolac molars, althoughtheir advanced wear state precludes clear observation of all rele-vant features, show none of the characteristics of Palaeoloxodon andare fully conformablewithMammuthus (Fig.15). First, themassivityof the molars, with M3 widths of 136 mm (right) and 129 mm (left)argues against Palaeoloxodon, as this genus (especially the Euro-pean Palaeoloxodon antiquus) is characterised by relatively narrowcrowns. The individual lamellae are of simple form, narrow andwith their two enamel bands parallel, as in Mammuthus, and lackthe ’cigar shape’ and blunt medial and lateral ends common inPalaeoloxodon. The subdivision of the plates in early wear, typicallyinto subequal rings in Mammuthus but with a long central ringflanked by much shorter lateral and medial ones in Palaeoloxodon,cannot be assessed in the Kostolac individual as all lamellae are infull wear. However, there is no sign of the characteristic, pointedmidline sinus, and tight folding, common in the enamel of Palae-oloxodon. The moderate enamel folding in M3 and M3, and midlineswelling seen especially in some lamellae of the M3, are normal forelephantid molars (including Mammuthus) worn nearly to thecrown base, and do not have taxonomic significance.

The species of Mammuthus represented can be assessedprimarily through the metric features of the molars (Table 2).

Upper and lower molars have both lost their anterior partsthrough normal wear in life. In both M3s, 16 plates plus a smallposterior platelet are preserved, but the anterior two plates areworn down to a dentine platform. Paired roots (sensu Sher andGarutt, 1987) extend to the front of the preserved crown, indi-cating that the unpaired anterior root has been lost by wear, andsuggesting that four (or possibly more) plates have been lost to

edial view. Anterior is to the right. b, d: Left upper third molar in occlusal and lateral

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wear, giving a likely total of around 20 plates, or possibly more. Inthe lowers, the left M3 preserves only the posterior 11 plates and itis not possible to reconstruct the original count.

A lamellar frequency of ca. 6.8 for each M3 is recorded as anaverage between the base and top of the crown, medial and lateralsides, according to the method of Maglio (1973). In the M3s, LF canbemeasured only at the crown base and therefore gives a low value(medial and lateral sides averaged) of 4.8e5.4, which can, however,be compared with analogous measurements on other samples(Table 1). Average enamel thickness across the occlusal surface is2.7e2.8 mm for the M3s and 2.9e3.1 mm for the M3s.

A final metric of taxonomic significance is the crown height ofunworn plates. Due to wear, in the M3s (Fig. 15c) this cannot beestimated at all, and in the M3s is measurable only in the posteriorpart of the crown. The maximum crown height in the right M3 ismeasured on the third plate from the posterior end (plateletexcluded), which is just coming into wear and measures 136 mm,giving a hypsodonty index 1.24 relative to the maximum crownwidth of 110 mm. This value, however, is clearly far below the fullcrown height of the molar which would have been maximal in theanterior to middle part of the tooth; the molar in lateral view(Fig. 15d) shows a steeply rising posterior face suggesting an orig-inal crown height of ca. 180 mm, giving a hypsodonty index of atleast 1.6.

These parameters strongly indicate referral of the Kostolacmammoth to the speciesM. trogontherii. Identity with the ancestralmammoth M. meridionalis is excluded by the plate count of morethan 16 and the reconstructed high crown, as well as by theelevated lamellar frequency (cf. Table 2). Plate counts of 20 canoccur at the low end of the range of variation within the woollymammoth M. primigenius, but the lamellar frequency at Kostolac istoo low, and the enamel too thick, for referral to that species. Bodysize is a malleable feature which should be used with care intaxonomic assignment, but the very large size of the Kostolacskeleton (see below) is more conformable with M. trogontherii(especially earlier European populations) than M. primigeniuswhich is generally smaller.

6.2. Body size and mass

Measurement data for the postcranial bones are given inTables 3e12. Based on published skeletons of M. meridionalis,M. trogontherii and M. primigenius tabulated by Lister and Stuart(2010), a ratio diagram (Fig. 16) clearly indicates the broad simi-larity in size and proportion of the Kostolac individual to othermaleskeletons of M. trogontherii, as well as to those of M. meridionalis.

For the estimation of shoulder height from isolated limb bones,Lister and Stuart (2010) developed regression equations based ona series of mounted skeletons. The three most reliable estimators,humerus, radius and femur lengths, give skeletal shoulder heightsfor the Kostolac skeleton of 369, 365 and 386 cm, respectively, withan average of 374 cm (Table 13). Different authors allow 5e7% extrafor flesh; at 6%, the Kostolac skeletal height of 374 cm gives a valueof ca. 396 cm. We suggest four metres as an approximate estimatefor the live shoulder height of the Kostolac Mammoth.

The body mass of the Kostolac mammoth can be estimated intwo ways. The first method uses regression equations of body massagainst shoulder height for living elephants. Using the 396 cmshoulder height estimated above, Roth’s (1990) regression equa-tions based on three L. africana populations give similar results,indicating a live weight of between 9.4 and 9.7 t for the Kostolacmammoth (Table 14). A second method is the use of individualbone lengths for estimating body mass. Using a combined sampleof modern Elephas maximus and L. africana, Christiansen (2004)derived prediction equations of body mass. Table 15 shows the

Table 3Measurements of vertebrae of the Kostolac mammoth (mm). Measurements that could be taken were limited by accessibility in the in situ skeleton. m-l, medio-lateral; d-v,dorso-ventral; a-p, antero-posterior.

Element Length of thoracic spinefrom top of neural canal

Maximumwidth

Posteriorarticular width

Posteriorcentrum width (m-l)

Posteriorcentrum height (d-v)

Centrumlength (a-p)

Atlas (1st cervical) 514Axis (2nd cervical) 182Third thoracic 560 149 163 98Fourth thoracic 570 154 163 100Fifth thoracic 580 363 133 93Sacrum (5 fused vertebrae) 500

Table 4Measurements of scapulae of the Kostolac mammoth. Equivalent measurement codes from Roth (1982) are given. m-l, medio-lateral; a-p, antero-posterior.

Description Total length(including dorsalepiphysis) ¼ TOT

Total length(excluding dorsalepiphysis) ¼ DIA

Length fromdorsal edge toclosest edge ofglenoid cavity

Glenoid todorso-posteriorcorner of scapula,parallel toposterior edge

Maximum widthof glenoid epiphysis

Articular widthof glenoid(a-p) ¼ LGA

Articular depthof glenoid(m-l) ¼ TSA

L scapula 321 219 152R scapula 1110 900 1075 685 334 230 151


results of applying these equations to limb bone lengths of theKostolac mammoth. Estimates range from 8 to 11 t, with an averageacross bones of 9.4 t. The concordance between the mass estimatebased on shoulder height, and those based on limb bone lengths, isexcellent, and indicates a live weight for the Kostolac mammoth ofca. 9.5 t. It should be remembered, however, that in livingelephants, an animal’s body mass can vary by up to 10% in the shortterm, depending on daily cycles of feeding, drinking, and defe-cating, and even more on a seasonal basis, due to food availabilityand reproductive cycles such as musth (males) and pregnancy(females). The estimated figure should be regarded as a centralestimate only.

6.3. Age and gender

The age of the mammoth at death can be estimated by refer-ence to the dental ageing scheme devised for African elephants (L.africana) by Laws (1966) and revised by Jachmann (1988), basedon mandibular teeth. The M3s of the Kostolac mammoth(Fig. 15a,c) have only the posterior 11 plates remaining, all ofwhich are in wear, and the crown is worn to a height of only 7 cm(left molar) and 5 cm (right molar). For a likely original platenumber of around 20 (see above), this indicates that ca. 11/20 or55% of the plate count remains, equivalent to ca. 6 plates ofL. africanawith a typical full plate number of 11 in M3 (Laws,1966).By reference to Laws’ scheme, this indicates age class XXIX, or an

Table 5Measurements of humeri of the Kostolac mammoth (mm). Equivalent measurement cod

Element Total length ¼ LONG Articular length ¼ ARTS Width ofepiphysis

L humerusR humerus 1215 1170 343

Table 6Measurements of ulnae of the Kostolac mammoth (mm). Equivalent measurement codes

Description Total length Articular length

L ulnaR ulna 1080 875

age of around 57 in ‘African elephant years’. A correction can bemade for the larger body size in M. trogontherii, which scales withpositive allometry to longevity among mammals. By applying theformula of Blueweiss et al. (1978) (longevity in days a body massin grams0.17), and using a longevity of 60 years and a mass of 6 t forliving bull elephants, and the body mass of 9.5 t for the Kostolacmammoth as above, a longevity of 65 years is obtained. An esti-mated absolute age for the Kostolac mammoth is therefore57 � 65/60 ¼ 62 years.

Consistent with this estimate of advanced age, all limb-boneepiphyses on the Kostolac mammoth are fused to their diaphysisshafts. This includes the scapular margin, the proximal femur andthe distal radius/ulna, which are the last to fuse in both livingelephants andmammoths (Roth,1984; Lister,1999). In the skull, thezygomatic arch, which also fuses very late in ontogeny (M. Ferretti,pers. comm.), is completely fused in the Kostolac mammoth.

The male gender of the Kostolac mammoth is clear fromseveral lines of evidence. First, the large size of the skeleton,comparable to known male M. trogontherii individuals such asWest Runton and Steinheim (Fig. 16), suggests male gender, sincethere is clear sexual dimorphism in body size among both livingelephants and mammoths (e.g. Averianov, 1996). Skull robustnessis also markedly stronger in male individuals, but the breakage ofthe Kostolac cranium makes this difficult to assess. The size of thetusks, however, provides a further clue. Two loose chunks of tuskfrom the Kostolac excavation indicate an original diameter,

es from Roth (1982) are given. m-l, medio-lateral.

proximal(m-l)

Minimum diaphysiswidth ¼ TSM

Width of distalepiphysis ¼ TSD

Width of distalcondyle ¼ TTC

352 277160 364 273

from Roth (1982) are given. m-l, medio-lateral.

Width of proximalepiphysis (m-l)

Proximal articularwidth (m-l)

ca. 330 279

Table 8Measurements of pelvic bones of the Kostolac mammoth (mm). ‘L’ are codes from Lister (1996a).

Element Maximumtransverse widthof pelvic girdle ¼ L1

Diagonal heightof aperture fromsacral scar toventral midline ¼ L2

Maximumtransverse diameterof pelvic aperture ¼ L3

Width of iliumfrom lateralmaximum tonearest pointof aperture ¼ L4

Minimum widthof ilium ¼ L5

Maximum widthof ilium

Antero-posteriordiameter ofacetabulum

Pelvis 1670 450 540 630 244 1040 192

Table 9Measurements of femora of the Kostolac mammoth (mm). Equivalent measurement codes from Roth (1982) are given. m-l, medio-lateral.

Element Total length ¼ articular length Minimum (along shaft)maximum (around shaft)diaphysis width ¼ APM

Proximal width(m-l) ¼ TSW

Width of femoralhead (m-l) ¼ HDD

Width of distalepiphysis (m-l) ¼ TSI

Width of distalarticulation

L femur 1470 ca. 196 437 187 301 261R femur 1470 186 450 298 271

Table 10Measurements of tibiae of the Kostolac mammoth (mm). Equivalent measurement codes from Roth (1982) are given. m-l, medio-lateral.

Element Total length Length to edge ofmedial proximal condyle

Minimum (along shaft)maximum (around shaft)diameter of diaphysis

Width of proximalepiphysis (m-l) ¼ TSM

L tibia 850 755 140 273R tibia 855 775 130

Table 11Measurements of fibulae of the Kostolac mammoth (mm). m-l, medio-lateral; a-p, antero-posterior.

Description Total length Minimum (along shaft)maximum (around shaft)diaphysis width

Width of distalepiphysis, perpendicular tolong axis of distal end (m-l)

Depth of distal epiphysis,parallel to long axisof distal end (a-p)

L fibula 805 49 130 ca. 68R fibula 790 49 130 ca. 98

Table 7Measurements of radii of the Kostolac mammoth (mm).

Element Total length Proximal width parallel to posterior edge Proximal articular width Proximal depth perpendicular to posterior edge

L radius 154 138 108R radius 925


presumably close to the base, of around 200 mm. In a large sampleof adult M. primigenius tusks from Siberia, Vereshchagin andTikhonov (1986) give a range of 47e90 mm in females and89e180 mm in males for tusk diameter at the base. The maleM. trogontherii from West Runton (UK), Azov I (Russia) and

Table 12Measurements of foot bones and patella of the Kostolac mammoth (mm). Astragalusmeasured parallel to ventral surface; calcaneummeasured parallel or perpendicularto main vertical medial facets. d-v, dorso-ventral; a-p, antero-posterior; m-l, medio-lateral.

Description Maximumlength (d-v)

Maximumdepth (a-p)

Maximumwidth (m-l)

L patella 137R patella 164R astragalus ca. 186L calcaneum 266 195 165R calcaneum ca. 260 194 176

Steinheim (Germany) have diameters of 210, 215 and 217 mm,respectively (Dietrich, 1912; Baigusheva and Garutt, 1987; Listerand Stuart, 2010). Conversely in the female M. trogontherii fromEdersleben (Germany) and Novogeorgievsk (Ukraine), the figuresare 138 and ca. 149 mm, respectively (Zakrevska, 1935; Garutt andNikolskaya, 1988). These data support the male gender of theKostolac M. trogontherii.

Most diagnostic is the form of the pelvic girdle. In Mammuthus,the pelvic aperture is relatively smaller, and the neck of the iliumrelatively wider, in males than in females, and the sexes can beseparated by the ratio between these two measurements. Theresult holds true for M. primigenius, M. meridionalis and Mammu-thus columbi, and is conformable in available individuals ofM. trogontherii (Lister and Agenbroad,1994; Lister, 1996a; Lister andStuart, 2010). In the Kostolac pelvis (Fig. 17), the ratio betweentransverse aperture width and neck width is 540/244 ¼ 2.21, whilebetween vertical aperture width (measured from the sacral scar tothe ventral midline) and neck width it is 450/244 ¼ 1.84. Thesevalues clearly fall within the male range of other Mammuthus

Limb size and proportion of Mammuthus species

scapula humerus radius femur tibia fibula60

70

80

90

100

110

120

Nogaisk

Odessa

Azov I Georgievsk

Steinheim

Scoppito

KOSTOLAC

West RuntonFarnetaDurfort

Azov II

Gelsenkirchen

Pyatiryzhhsk

Borro al Querco

Savignano

Fig. 16. Ratio diagram of limb bone lengths in Eurasian male Mammuthus skeletons. Red: M. meridionalis; green, M. trogontherii; blue, M. primigenius. Kostolac mammoth in bold;other lines are dashed and dotted to aid visual separation. Ratios calculated as a percentage of West Runton measurements; see Lister and Stuart (2010, Supplementary Table B) forraw data and sources. Only fully- or almost fully-grown individuals are included. M. primigenius individuals not separately named, but range from Sevsk (the smallest) to Siegsdorf(the largest). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

Table 13Estimation of Kostolac mammoth shoulder height from lengths of long bones, based on linear regression equations derived frommounted skeletons ofMammuthus (Lister andStuart, 2010). a, b ¼ slope and intercept of regression equation.

Element Equation R-square 95% confidence interval/prediction interval

Kostolac skeletal shoulderheight estimate þ/� 95%confidence/prediction interval (cm)

Kostolac live shoulderheight (cm), adding6% for flesh

‘All’ humerus a ¼ 2.9701, b ¼ 85.4976 0.8613 3.29/11.49 369 � 12.1/42.4 391‘Best’ radius a ¼ 4.0195, b ¼ �67.6722 0.9018 3.19/10.57 365 � 11.6/38.6 387‘All’ femur a ¼ 2.9458, b ¼ �438.9187 0.8861 3.09/10.27 386 � 11.9/39.6 409Average 396

Table 14Estimation of bodymass (in kg) of the Kostolacmammoth based on predictive equations from shoulder height of modern African elephant populations. Roth’s (1990) data fromLaws et al. (1975), Johnson and Buss (1965), Tumwasorn et al. (1980) and others; Christiansen’s (2004) data from Benedict (1936) and others. m ¼ mass (kg), sh ¼ shoulderheight (cm).

Source population Code in Roth (1990) Code in Christiansen (2004) Equation Body mass (kg) at 396 cm shoulder height

L. africana males, Uganda c 1 m ¼ 5.07*10�4sh2.803 9691L. africana males, Uganda e m ¼ 3.06*10�4sh2.890 9841L. africana males, Uganda f m ¼ 1.81*10�4sh2.97 9394Average 9642

Table 15Estimation of body mass (in kg) of the Kostolac mammoth based on predictive equations from total bone lengths of modern elephant populations, after Christiansen (2004,Tables 3 and 4). Equations are of the form log (mass in kg) ¼ a þ b (log X), where X is bone variable (mm). %SEE and %PE are, respectively, the standard error and percentprediction error of the regression. Where both left and right bones are preserved in the Kostolac skeleton, their mean length was taken.

Bone length a b %SEE/%PE Kostolac measure (mm) Estimated body mass (kg)

Humerus �4.145 2.635 11.52/6.74 1215 9613Radius �3.838 2.634 10.11/6.64 925 9437Ulna �4.135 2.674 8.41/5.34 1080 9471Femur �5.568 3.036 14.54/6.15 1460 10,939Tibia �3.064 2.378 12.47/6.93 852 8028Fibula �3.086 2.422 18.68/11.4 798 8763Average 9375


Fig. 17. Pelvic girdle of the Kostolac mammoth in posterior view. The relatively smallpelvic canal and wide ilium signify male gender.


(Fig. 18), and corroborate the value of this feature for genderdetermination in M. trogontherii.

6.4. Further morphological features

Damage to the cranium precludes a thorough consideration ofits anatomy, but one visible feature is of interest. The zygomaticprocess of the maxillary forms, in its lower part, a bony bridgebounding the infraorbital canal. This ‘infraorbital process’ is prim-itively robust among elephantids, but is narrower and thinner inMammuthus, especially in M. primigenius (M. Ferretti, pers. comm.,2005). In the Kostolac cranium, the process is remarkably slender(Fig. 6b). In the West Runton (UK) M. trogontherii skull, by contrast,the infraorbital process is relatively wide (80 mm at its narrowest

1.4 1.6 1.8 2.0 2.2

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MALES

M. trogontherii

M. primigenius

Fig. 18. Ratio between pelvic aperture width and ilium shaft width. Open symbols: males; filprovided by cranial morphology, tusk size or body size (see Lister, 1996a). The identity of the(Lister, 1996b; van Essen, in litt.). Based on original measurements; the ratio of the femalea photograph (Zakrevska, 1935).

point), but quite thin e this apparently ‘mosaic’ condition betweenprimitive and derived conditions within Mammuthus (Lister andStuart, 2010) might reflect the relatively early age of W. Runton(ca. 700 ka) within European M. trogontherii. Alternatively thedifference of form between the Kostolac and W. Runton specimensmight simply reflect individual variation; further material isrequired to test between these alternatives.

Only parts of the mandible are preserved at Kostolac, but theanterior portion shows a relatively short symphysis and modestrostrum, conforming to the advanced or derived condition withinMammuthus, similar toM. primigenius andmostM. trogontherii, andcontrasting with the longer symphysis of most M. meridionalis (cf.Lister and Stuart, 2010). Ferretti and Debruyne (2011) discuss thetaxonomic distribution of the anterior mandibular foramina. Forthe lateral mental foramina, 86.4% of M. meridionalis specimens(n ¼ 22) possessed 1 or 2 foramina on each hemi-mandible, andonly 13.6% possessed 3 or more. In M. primigenius, by contrast(n ¼ 54), the figures are 44.4% and 55.6%, respectively. Averagefrequency therefore increased between these taxa. The Kostolacmandible shows two foramina on the lateral side, as does the WestRuntonM. trogontherii mandible, a condition commonly also foundin the other two species; a larger sample ofM. trogontheriiwould berequired to test if the species is statistically intermediate.

The presence of a mental foramen on the medial side of theramus is considered by Ferretti and Debruyne (2011) to be a syna-pomorphy of Elephantinae (including Elephas and Mammuthus).Among mammoths, however, it is absent from two specimens ofMammuthus rumanus, but present in 73% of M. meridionalis(n ¼ 30), 69% ofM. trogontherii (n ¼ 18), and 100% ofM. primigenius(n ¼ 173). The frequency of the foramen therefore increasesthrough the sequence. In the Kostolac mandible, the right ramusshows a single pit (14 � 20 mm) leading to two foramina. On theleft ramus, there is a single wide foramen leading into themandibular canal that can be seen due to breakage of themandibular wall. The presence of the foramen at Kostolac and also

2.4 2.6 2.8 3.0 3.2

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Eder

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ver

Gydan

Lena

Aa Kerk

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Oyosh

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FEMALES

led symbols: females. For the majority of specimens, independent evidence of gender isfemale Edersleben (Germany) skeleton as M. trogontherii or M. meridionalis is debatedNovogeorgievsk specimen may be slightly exaggerated as measurements taken from


atWest Runton (Lister and Stuart, 2010) increases the proportion inM. trogontherii to 73% (n ¼ 20), identical to that of M. meridionalis.

The vertebral column of the Kostolac skeleton is the mostcomplete known in M. trogontherii. Another, in the cf.M. trogontherii skeleton from Novogeorgievsk (Ukraine), is statedby Zakrevska (1935) to possess 19 preserved thoracic vertebrae andfour sacrals. At Kostolac, there are 7 cervical vertebrae, 19 thoracics,four lumbars, and a sacrum comprising five fused vertebrae. Eigh-teen thoracic vertebrae have been found in some skeletons ofM. primigenius (e.g. Berezovka: Zalensky, 1903), living elephantstypically have 20 (Mol and van Essen, 1992), and variability inM. trogontherii is uncertain.

The two proximal caudal vertebrae are preserved, and six others,incomplete and separated from the skeleton. The combined lengthof these eight vertebrae is approximately 80 cm, indicatinga complete tail substantially longer than this. This suggestsa significantly longer tail than in M. primigenius, where the totallength of the tail vertebrae in the Beresovka carcass was measuredas 60 cm, with the tail in the flesh measured as 36 cm (Herz, 1902;Zalensky, 1903), exceptionally short for an elephantid and inter-preted as an adaptation to cold climate (Lister and Bahn, 2007).

The feet are unfortunately not fully exposed, so it is not possibleto assess the structure of the carpus, which shows significantvariation within the genus Mammuthus (Lister, 1996b). An inter-esting feature, however, is seen in the left hind foot, where themetatarsal of the first digit is complete and well preserved. Thedistal end of mtI shows no articular surface for the attachment ofa first phalange, indicating that there are only four toes on the hindfoot in this specimen, as observed frequently in other probosci-deans. On the caudal side of the distal end a small articular surfacecan be noted for the attachment of a sesamoid.

7. Geological age of the remains

The geological context, and limited other fauna found in themammoth layer, do not, at the present state our regional under-standing, provide evidence on the precise age of the skeleton.

The identification of the Kostolac skeleton as M. trogontheriidoes, however, place some constraints on the age of the deposit. InEurope this species is characteristic of the early Middle Pleistocene(Cromerian Complex), where it is well-known from localitiesincluding West Runton (UK), Süssenborn and Mosbach (Germany)and Tiraspol (Moldova), spanning approximately MIS 19e13 (ca.780e500 ka). At some localities it has been identified earlier,together with the more primitive M. meridionalis, such as at Dorn-Durkheim, Germany (just below the Brunhes/Matuyama boundary)and at Sinyaya Balka, Russia, in deposits ca. 1.0 Ma in age (Listeret al., 2005).

After the Elsterian/Anglian glaciation (MIS 12), mammoths inEurope showa transition fromM. trogontherii toM. primigenius. Theremains from Steinheim, Germany, considered to be MIS 11e10 inage (ca. 400e350 ka) (Schreve and Bridgland, 2002), are dentallysimilar to earlier M. trogontherii (and to Kostolac). They are also ofsimilarly large body size: at 370 cm, the reconstructed shoulderheight of the famous Steinheimmammoth skeleton (Dietrich,1912)is only slightly smaller than that of Kostolac (Fig.16). Later EuropeanM. trogontherii populations, while dentally still more primitive thantypical M. primigenius, were considerably reduced in size. A likelyage bracket for the Kostolac skeleton is therefore ca. 1.0e0.4 Ma.

8. Conclusions

The Kostolac skeleton is one of very few largely complete skel-etons of M. trogontherii. It therefore adds significantly to ourknowledge of the osteology of this species, important as an

evolutionary intermediate between the better-knownM. meridionalis and M. primigenius, between which there wassubstantial evolutionary change (Lister et al., 2005). The Kostolacskeleton provides new information in features such as the vertebralcount and in details of the skull and feet.

The skeleton is also the only one of M. trogontherii preserved ina largely articulated condition, and is very unusual for any fossilelephantid skeleton in being preserved in the animal’s likely deathposture, a remarkable taphonomical situation that invites furtherstudy.

Finally, as the first complete skeleton of this species found in theMediterranean basin, and with further isolated remains found inproximity, the finds highlight the potential importance of the Kos-tolac area for our understanding of the regional history of theQuaternary mammal fauna. Further investigations are likely to yieldassociated fauna of biostratigraphic and ecological significance.

Acknowledgements

We are grateful to the excavators of the Roman site of Vimina-cium for their fast response and sincere delight with an animalmuch older than their period of interest. We especially thank DrMiomir Kora�c, director of Viminacium site, for organizing theexcavation, supporting the conservation project, and buildinga protective cover over the skeleton. Special thanks go to Milo�sMilivojevi�c, senior geological preparator, Natural History Museum,Belgrade, for his effort in conservation of the skeleton and resto-ration of the skull and tusks. We are grateful to Hans Wildschut forphotography. Thanks are also due to Marco Ferretti and Ralf Kahlkefor their rapid and valuable comments on the manuscript, and toFrédéric Lacombat and Norm Catto for accommodating our paperwithin this special issue of Quaternary International. V.D.’s contri-bution forms part of the project “Bioarchaeology of AncientEurope e humans, animals and plants in the prehistory of Serbia”(III47001) funded by the Ministry of Education and Science of theRepublic of Serbia.

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Lister Et Al., 2012 - M. Trogontherii Drmno, Serbia