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, UK
b
Laboratory of Bioarchaeology, Department of Archaeology, Faculty of Philosophy, University of Belgrade, Serbia
c
Natural History Museum, Belgrade, Serbia
d
Department of Geology, Faculty of Mining and Geology, University of Belgrade, Serbia
e
Natural 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
a b s t r a c t
The Kostolac mammoth was discovered in 2009 in Pleistocene deposits adjacent to the Drmno open-cast
lignite mine in the Serbian Danube Basin. On the basis of cranial and dental features, the individual is
identied as the so-called steppe mammoth, Mammuthus trogontherii. The remains are those of an old
male of estimated age around 62 years, and comprise one of the most complete and best-preserved
known skeletons of this species, and the rst from the region. Skeletal height is estimated as around
four metres, and body mass 9.5 t. The excellent preservation of the skeleton provides new information
about the osteology of M. trogontherii, an evolutionary intermediate between the better-known ancestral
mammoth Mammuthus meridionalis and woolly mammoth Mammuthus primigenius. The nd is also
remarkable for the articulated condition of the skeleton, the animal occupying a crouching posture which
is probably little-altered from its original death position. This and the depositional environment of the
skeleton, a broad, fast-owing river, suggest that the animal died in relatively shallow water and was
very 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 1.0
and 0.4 Ma.
2012 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
The Kostolac mammoth was discovered on May 27, 2009, in an
open lignite mine adjacent to the famous Roman site of Vimina-
cium, approximately 90 km east of Belgrade, Serbia (Korac et al.,
2010; Fig. 1). The area, within the Danube basin, lies on the
boundary between the Balkan Peninsula and the Pannonian plain.
The Kostolac coal basin, due east from the Velika Morava river, has
been exploited since 1870. The mammoth skeleton was discovered
on the north-east rim of the Drmno pit, the youngest and largest of
the open pits, near the mouth of the Mlava River, on low hilly land
that rises over the at landscape of the alluvial plains. On the west
side, above the left bank of the Mlava, the high ground of Pozar-
evacka greda spreads in a SSEeNNW direction. The mammoth was
found at 44

43
0
50
00
N, 21

14
0
21
00
E, Gauss-Krueger grid reference
4 954 024, 7 518 937, at a height of approximately 58 mabove mean
sea level, and a depth of 27 m below the modern surface.
The skeleton, remarkable for its completeness and excellent
state of preservation, is one of very few known skeletons of the
Middle Pleistocene steppe mammoth, Mammuthus trogontherii.
This species occupies an important position in the evolution of the
genus Mammuthus, as it is considered to be an intermediate
between the widespread, and better-known, Early Pleistocene
species Mammuthus meridionalis, and the familiar woolly
mammoth, Mammuthus primigenius (Lister et al., 2005). Any
information on its anatomy and adaptations is therefore of interest.
The Kostolac mammoth is also the rst skeleton of M. trogontherii
from the Mediterranean Basin, and its taphonomic situation is
remarkable, strongly suggesting that it has been preserved in death
position.
2. Regional geological setting
The geology of the Kostolac basin comprises sediments of Late
Miocene to Quaternary age. Late Miocene sediments formed in a
former gulf of the Paratethys at the southern edge of the Pannonian
province. The Pontian of the Paratethys is the chronostratigraphic
* Corresponding author.
E-mail address: a.lister@nhm.ac.uk (A.M. Lister).
Contents lists available at SciVerse ScienceDirect
Quaternary International
j ournal homepage: www. el sevi er. com/ l ocat e/ quai nt
1040-6182/$ e see front matter 2012 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2012.03.021
Quaternary International 276-277 (2012) 129e144
equivalent of the Messinian. Its mollusc fauna includes freshwater
forms such as unionids and Viviparus, indicating local freshening of
environment due to inowof river water fromthe land. This type of
facies is known in the literature as the Kostolac brackish Pontian
(Stevanovic, 1951, 1977, 1990). Here are found deposits of soft
brown coal (lignite) and layers of silt, marly clays, sands and
gravelly sands. Coal is exploited in quarries near Kostolac town and
the villages of

Cirikovac, Klenovnik and Drmno. To the south, near
the village of Poljane on the heights of Pozarevacka greda, Late
Upper Miocene deposits of Pannonian age are found, with lesser
deposits of coal (Milosevic and Miletic, 1975).
Quaternary deposits cover all the land in a wide area around
Drmno. They include Pleistocene and Holocene sediments. Pleis-
tocene features are represented by pre-loess proluvialedeluvial
deposits of the Klicevac Formation, loess, and aeolian sand (Rakic,
1978).
The Klicevac Formation, named after the settlement of Klicevac,
is located on the right bank of the Danube ENE of Drmno. It lies
uncomformably over Neogene sediments and below Pleistocene
loess and other Quaternary features. This formation consists of
loessoid sandy-clay silt and silty sands, gravels, sandy limestones,
sandstones and tufa. The most widespread are sediments deposited
on 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 Recice and Klicevac are formed from brown iron
sands with gravel and silt lenses, in which ostracods and a mixed
fauna of freshwater and terrestrial molluscs (Pisidium amnicum,
Planorbis sp., Unio sp., Clausilia sp., etc) are found (Rakic, 1978).
Aeolian sediments are the most widespread Pleistocene features
in the broad area of Drmno. Loess covers the whole land surface and
is represented by terrestrial and wetland loess. Along the right bank
of the Danube near Klicevac there are deposits of aeolian sand.
Holocene sediments are various. Alluvial deposits in river valleys
of theDanube andits tributary theMlava have the greatest thickness.
They comprise different facies including the sediments of oods, still
waters and marshes. Elsewhere, contemporary deluvial and
deluvialeproluvial deposits occur, as well as anthropogenic deposits
such 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 and
Pleistocene deposits are seen (Fig. 2). The Neogene is represented
by coal deposits of Pontian age, developed as the brackish Kostolac
type. These layers, below the mammoth, are folded in a low
syncline, different in structure from the upper, horizontal Pleisto-
cene sediments.
The coal deposits, around 50 m in depth, consist of gray ne-
grained mica and silt sands with thin intercalations of somewhat
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) 129e144 130
harder grayeyellow sands. These are deltaic sediments formed in
a coastal area of the Paratethys gulf where a river emptied fromthe
south.
Younger deposits, lying directly beneath the mammoth layer,
are formed of orange and brown medium-grained and coarse-
grained sands with cross-bedded stratication. These are sedi-
ments of a deltaic front formed in a phase of strong river ow and
withdrawal of the Paratethys coastal line. Samples from these
sediments have yielded rodent and insectivore remains including
Occitanomys sp., Kowalskia sp., Desmanodon sp. and Sorex sp.
The overlying Pleistocene sediments are divided into a lower
complex of river sediments, and an upper complex of loess. The
lower complex comprises two horizons, with a total thickness in
the studied section of about 10 m.
The skeleton of the mammoth was found in the lower horizon of
the lower complex, which lies unconformably over the Pontian
palaeorelief (Fig. 2). It is formed fromyellow, brownish-yellow and
gray gravels and sandy gravels, and brownish-reddish, yellow and
gray dusty sands, silt lenses and clays. The gravels are slightly
rounded and contain quartz, hornstone and metamorphite, as well
as smaller fragments of Mesozoic sandstones, limestones and
igneous rocks. Sands and gravels are cross-bedded in places. Based
on lithostratigraphic characteristics it can be concluded that these
sediments were deposited by a river trending SSEeNNW, with
varying but often strong ow. The river valley had a substantial
breadth of approximately 600 m. The mineral composition of the
clasts suggests that the Pleistocene river owed from the zone of
Serbian-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 ood deposit. This
comprises sandy clays and gray-brownish silts, with lenses of
oxbow-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 include
a fragmentary elephantid molar and a radius and ulna of the giant
deer Megaloceros giganteus.
The upper complex of Pleistocene sediments comprises loess,
and is also divided into two horizons. The older horizon is a wetland
loessoid sediment with a terrestrial and freshwater mollusc fauna.
The younger horizon includes sediments of land-slope loess. The
thickness of the loess complex varies from 10 to 15 m.
Anthropogenic Holocene deposits are found at the surface of the
section (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 of
the 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 skull
remains whole and in place, although its forehead is cracked and
sunken (Fig. 6a). The middle part of the right tusk, up to the alve-
olus, was detached and broken into pieces. The distal part of the
right tusk remains in situ (Fig. 3b). The left front leg bones were
detached from their position and damaged.
When the digger stopped, many fragments of the tusks and skull
that had been detached were collected but it was not possible to
assemble 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 carefully
uncovered in situ. The bones of the legs are folded beneath the
backbone, scapula and pelvic girdles (Fig. 7). The excavation was
performed by the archaeological team of the Roman site of Vim-
inacium and experts fromthe Natural History Museum in Belgrade.
Immediately after excavation, thorough conservation of the skel-
eton was undertaken and a protective structure constructed over it.
Due to these precautions the skeleton survived, as it was on
unstable 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 the
skeleton completed and the skull and tusk reconstructed.
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 silty
sand with intercalation of grayeyellow sand; 3. Orange and brown sand with cross-
bedded stratication. 4e5: Quaternary, Pleistocene. 4. Alluvial sediments. 4a:
Riverbed sediments: gravel, yellow sand, gray sandy gravel, and clays, with mammoth
remains 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
The skull preserves, on the right side, the zygoma, auditory
opening, orbital region, and the lateral wall of the braincase (Figs. 6
and 8; Table 1). The right side of the nare (trunk attachment) and
frontal are crushed and buckled downwards. Only parts of the right
tusk are preserved: a fragment of the proximal end remaining in
the alveolus, and a separate distal portion, some 85 cm in length
and ending very close to the original tip (Fig. 9). This portion
appears to be in life position relative to the skull (Fig. 3b); if so, the
original length of the tusk, fromthe alveolus to the distal tip, can be
measured as approximately 260 cm. Other pieces of tusk were
found loose. Of the mandible only one articular condyle and the
symphysis region are preserved.
Apart from the tail, the vertebral column is complete and, with
some disarticulation and movement, in anatomical order (Figs. 7
and 10). The atlas is pressed around the occipital condyles of the
skull. The axis and following ve cervical vertebrae are in line
behind it. They are inclined downward at some 45

to the line in
which the skull and the distal end of the right tusk lay. The rst and
second thoracic vertebrae continue the row. The left and right rst
ribs are articulated between the last cervical and rst thoracic
vertebrae. The proximal part of the left second rib is articulated
between the rst and second thoracics. This rib lacks part of its
body, but its middle part is in situ. The following four thoracic
vertebrae, from the third up to the sixth, are displaced, as a group,
to the right (Figs. 7 and 10). The remaining thoracics are all in
articulation. The middle thoracics (seventh up to tenth) lie in
a straight line that is parallel to the line in which the skull and the
fragment of the tusk lay. The posterior thoracics, lumbar and sacral
parts of the backbone are inclined from that line and bent to the
right. The caudal vertebrae were displaced during excavation
(Fig. 11). A total of 13 right and 11 left ribs were uncovered during
the excavation. Two ribs were found several metres apart to the
Fig. 3. a: skeleton in situ, view from SW (photo courtesy of Viminacium archaeological team). Stratied river sediments are seen in the section behind the skeleton. In the
background to the left, a prole of loessic deposits is seen, with Holocene cover on top. This includes anthropogenic deposits with brick graves of the Roman necropolis of
Viminacium. b: Close-up of the mammoth skeleton. The isolated right tusk-tip is seen to the left.
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 132
right side of the skeleton, and four ribs several metres away fromits
rear side (Figs. 4 and 12). The sternum is preserved, showing the
presternum and mesosternum fused together. The sacrum has
fallen beneath the pelvis.
The right scapula is signicantly rotated in such a way that it lies
next to the skull (Fig. 8). The right humerus is perfectly preserved,
with the articulated ulna and radius bent beneath it (Fig. 13). The
bones of the right front foot are probably beneath the distal end of
the ulna and radius, buried deep in the sediment. It was not
possible to excavate deeper beneath the skeleton and uncover the
carpals and further bones of the distal leg, since this would most
probably cause the collapse of the skeleton.
The bones of both hind legs are well preserved. There is
remarkable symmetry between the position of the left and right
limb bones (Fig. 7). The caputs of the femora had fallen out of
the acetabula, but remained just beneath them. The femora are
directed forwards and diverge somewhat laterally from the overall
midline of the skeleton. Beneath them, the tibiae and bulae are
exed backwards and lie approximately parallel to the main axis of
the skeleton. The right patella is still in position in the knee joint;
the left one fell out in the course of the excavation. The calcanea
and astragali of both feet are visible (Fig. 7). In the right foot some of
the smaller tarsal bones are also visible, together with at least eight
sesamoid bones, each 40e45 mm in diameter. The metatarsal and
rst phalange of the fth digit are visible, as well as a further
metatarsal and a rst phalange. In the left foot, the navicular is
visible, together with two metatarsals and at least six sesamoids.
Since these elements are well preserved and perfectly articulated it
is expected that further metatarsals and phalanges remain buried
in the sediment.
5. Taphonomy of the skeleton
The skeleton lies in a coarse, yellow river sand with variably-
shaded bands of ner sand, coarser sand and ne gravel, some-
times cross-bedded (Figs. 2 and 3). It is largely complete, with the
majority of bones articulated and positioned as in the live animal.
The bones of the hind legs, in particular, impressively recall the
posture of a living animal. It is likely that the animal was kneeling at
the nd-spot, and eventually died in that posture. This indicates
that the skeleton is in primary position and that the body was
buried under the sandy sediments shortly after the animal died.
The conclusion of rapid burial is strengthened by the observation
Fig. 4. Field drawing of the mammoth skeleton in situ (drawing by Dragana Petrovic).
Fig. 5. Skeleton in situ, view from NW during excavation, with cranium and right
scapula in the foreground. Photo courtesy of Viminacium archaeological team.
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 133
that the skeleton was not disturbed or damaged by predators,
which would undoubtedly have been attracted by the dead body if
it had been exposed for any period of time. The skeleton was
probably complete before its discovery and excavation.
Although the animal was of an advanced age, a survey of the
skeleton revealed no evident signs of pathology. No signicant
arthritic growth or exostosis was seen on any bone, although these
are common in skeletons of old elephantid individuals. This
suggests that the animal was in good health. The posterior articular
facets of the fourth thoracic vertebra (T4) (Fig. 14) showstrong left-
right asymmetry, although in T3 and T6 they are roughly
symmetrical, 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. The
vertebral spines are straight, and there are no perforations at their
base as seen in some Mammuthus skeletons (e.g. Lister, 2009,
Fig. 17A). Of course, the animal might have suffered from a disease
that does not leave pathological traces on bones, so illness cannot
be completely excluded as possible cause of death.
A second possibility to be considered is that the animal died of
starvation due to the advanced state of wear of its molars. Each
lower molar, in particular, has only about half of its enamel plates
left, and they are obliquely worn and narrowed because of their
proximity to the crown base (Fig. 15a). That elephants do die from
inability to feed with worn-out teeth is well-known (Shoshani,
1991). An animal the size of the Kostolac mammoth would have
needed well in excess of the 200 kg of fresh food per day typical of
African elephants today, which may chew for 15 or more hours per
day (Sukumar, 2003, p.197). The lower molars are at the 29th of 30
eruption and wear stages observed by Laws (1966) for Loxodonta
africana e implying that few African elephants survive beyond this
stage. However, the upper molars of the Kostolac mammoth still
retained a signicant proportion of their crown, and the occlusal
surfaces of the upper and lower molars are of similar dimensions. In
the absence of quantitative published data on the dental wear stage
at which elephants are threatened with starvation, it is difcult to
reach a rm conclusion on this possibility.
Modern elephants that are severely injured, diseased or starving
frequently go to water, where they commonly die. In either of the
above scenarios, the mammoth may have stood in shallow water in
the river, and then collapsed to its knees before expiring. The
illustrations of numerous elephant carcasses that died in Africa due
to drought (Beard, 2000) show in many cases that the animal died
in a crouching position with its legs folded under it, very similar to
the situation in the Kostolac mammoth.
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 lateral
close-up of cheek region of the Kostolac mammoth cranium, showing jugal bone (j) and narrow infraorbital process (i).
Fig. 7. Skeleton in situ, view from NE. Pelvis and hind legs in the foreground. Note the
symmetrical position of the left and right hind limb bones. Below the distal tibiae
proximal tarsal bones are seen, while further tarsal bones disappear into the sediment.
Photo courtesy of Viminacium archaeological team.
Fig. 8. Close-up of the cranium and right scapula, viewed anteriorly and to the right.
Photo courtesy of Viminacium archaeological team.
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 134
A nal possibility is that the animal was in good condition but
died due to a mishap in the river itself. Living elephants commonly
frequent water and are good swimmers. It is likely that M. trogon-
therii were similarly capable of walking or swimming through the
current of a river. This could have endangered the mammoth in
several ways, for example if it became embroiled in a whirlpool, or
if the mammoth fell through the ice in winter. The animal might
have struggled against the water owand/or soft substratum until,
exhausted, it gave up the struggle, knelt down and eventually died.
Sukumar (2003, p. 266) lists drowning in swiftly owing water as
one of the causes of death in living elephants.
Whatever the reason for the mammoths demise, the posture,
articulation and excellent condition of the preserved remains
indicate that it was buried rapidly, while the skeleton was still held
together by esh. The strong river ow indicated by the enclosing
lithology suggests that the river was carrying substantial quantities
of sand and gravel, and it is likely that the carcass itself acted as
a sediment trap, hastening its own burial.
Movement and slight dispersal of the bones probably happened
during the process of burial while the esh was disintegrating,
inuenced by the water ow around the massive dead body. Some
movement may also have occurred post-depositionally, i.e. after it
had been covered in wet sand. The depression and cracking of the
front of the skull may have happened due to trampling by other
mammoths, or post-depositional sediment movement. The post-
cranial skeleton mainly shows disturbance of the anterior elements
of the skeleton, namely the scapula, ribs and vertebrae. The right
scapula is rotated and displaced anteriorly, presumbly due to water
and sediment ow after sufcient soft-tissue deterioration. The
third to sixth thoracic vertebrae are displaced sideways from the
main line of the backbone, probably as a consequence of pressure
on this region of the back during the process of bodily decompo-
sition. The hind part of the backbone is inclined to the right from
the line in which lay the skull and the front part of the skeleton, at
an angle of approximately 60

. Displacement of bones mainly

affected the ribs, probably loosened by the collapse and subsequent
inclination of the backbone, and then moved by river ow. The hind
limbs are relatively undisturbed, but the caputs of the femora had
fallen out of the acetabula, a consequence of the weight of the
bones pulling them down after decomposition of the esh and
deterioration of the ligaments that held the joint. These processes
would have been aided by movement of the soft substratum.
Beside the right side of the skeleton, close to its front leg bones,
two more animal bones were found: a deer ulna and a horse radius.
Table 1
Measurements of cranium and mandible from Kostolac (mm). Because of breakage to the left side of the cranium, width measures were estimated by doubling the dimension
from 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 of
cranium from
back of occiput
to front of
preorbital rim
Length from
front of auditory
opening to
lowest point
inside orbit (Ab)
Height of orbit,
measured on
its inside (Ac)
Internal
height of
nasal
opening (Ld)
Internal width
of nasal
opening (Ih)
Width of
cranium
around
zygoma (Iq)
Width of
cranium
around
occiput (Ia)
Minimal
width
of braincase
Minimum
depth (d-v)
of zygomatic
arch
Minimum
width (m-l)
of zygomatic
arch
Diameters
of mandibular
condyle
850 417 152 197 600 900 1080 440 64 30 118 89
Fig. 9. The terminal portion of the right tusk, viewed from the direction of the skel-
eton. Photo courtesy of Viminacium archaeological team.
Fig. 10. The anterior part of the vertebral column, viewed from the left. The cervical
vertebrae are seen to the left of the image, followed by the thoracic vertebrae in three
sharply disjointed and differently-oriented groups. Photo: Hans Wildschut.
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 135
By its size and morphology the deer ulna is identied as red deer,
Cervus elaphus, while for the horse radius only a generic identi-
cation (Equus sp.) is possible. To check whether these bones indi-
cated further skeletal remains at the site, the excavation area was
enlarged several metres to the north-east, i.e. along the right side of
the mammoth skeleton. No other bones were found. These isolated
bones must represent parts of disarticulated carcasses that had
been transported during the same sedimentary event that
preserved the mammoth. Conceivably their entrapment in that
position was related to the current ow and/or sedimentation
accumulating around the mammoth carcass.
A year before the skeleton was discovered, on August 4th 2008,
a mammoth tibia was found at Kostolac, at a depth of 26 mfromthe
surface, 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 the
bone, indicating that it was buried in sand. The tibia belonged to an
immature animal, judging by its unfused epiphyses, yet it is of large
size e total length 850 mm, identical to the tibiae of the skeleton.
This would be conformable with M. trogontherii and suggests that
the discovered skeleton might not be the only one buried and
preserved in the sand, and that future excavations in the Kostolac
open mine might reveal more nds.
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 temporal
lines (Fig. 6a), are typical of Mammuthus and exclude Palaeoloxodon.
The cranial vertex, although damaged, appears to show a single
peak characteristic of Mammuthus, rather than the double peak of
Palaeoloxodon. Further taxonomic characters are provided by the
molar teeth.
The crowns of the left and right upper, and left lower, molars are
preserved almost in their entirety (Fig. 15), as well as the anterior
and posterior portions of the right lower molar with a break
inbetween. Several features indicate clearly that these are the third
(last) molars. First, the tapering form at the posterior end of the
crown. Second, in the lower molars, a signicant length of crown is
Fig. 11. Four caudal (tail) vertebrae found in articulation. Photo: Hans Wildschut.
Fig. 12. The skeleton during excavation, viewed from behind. The displacement of
several 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 precisely
beneath the humerus. Photo: Hans Wildschut.
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 136
preserved despite the tooth being in an advanced state of wear; this
is possible only in M
3
because forward movement of the tooth had
ceased. Finally, the presence of a single, curtain-like longitudinal
root is typical of M
3
in advanced wear.
The two candidate genera in the European Pleistocene are
Palaeoloxodon and Mammuthus. The Kostolac molars, although
their advanced wear state precludes clear observation of all rele-
vant features, shownone of the characteristics of Palaeoloxodon and
are fully conformable with Mammuthus (Fig. 15). First, the massivity
of the molars, with M
3
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 narrow
crowns. The individual lamellae are of simple form, narrow and
with their two enamel bands parallel, as in Mammuthus, and lack
the cigar shape and blunt medial and lateral ends common in
Palaeoloxodon. The subdivision of the plates in early wear, typically
into subequal rings in Mammuthus but with a long central ring
anked by much shorter lateral and medial ones in Palaeoloxodon,
cannot be assessed in the Kostolac individual as all lamellae are in
full wear. However, there is no sign of the characteristic, pointed
midline sinus, and tight folding, common in the enamel of Palae-
oloxodon. The moderate enamel folding in M
3
and M
3
, and midline
swelling seen especially in some lamellae of the M
3
, are normal for
elephantid molars (including Mammuthus) worn nearly to the
crown base, and do not have taxonomic signicance.
The species of Mammuthus represented can be assessed
primarily through the metric features of the molars (Table 2).
Upper and lower molars have both lost their anterior parts
through normal wear in life. In both M
3
s, 16 plates plus a small
posterior platelet are preserved, but the anterior two plates are
worn down to a dentine platform. Paired roots (sensu Sher and
Garutt, 1987) extend to the front of the preserved crown, indi-
cating that the unpaired anterior root has been lost by wear, and
suggesting that four (or possibly more) plates have been lost to
Fig. 14. Fourth thoracic vertebra (T4), in posterior view, showing the vertebral canal
and base of the thoracic spine. The strong lefteright asymmetry between the posterior
articular facets is clear.
Fig. 15. Molar teeth of the Kostolac mammoth. a, c: Left lower third molar in occlusal and medial view. Anterior is to the right. b, d: Left upper third molar in occlusal and lateral
view. Anterior is to the left. Scales 5 cm.
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 137
wear, giving a likely total of around 20 plates, or possibly more. In
the lowers, the left M
3
preserves only the posterior 11 plates and it
is not possible to reconstruct the original count.
A lamellar frequency of ca. 6.8 for each M
3
is recorded as an
average between the base and top of the crown, medial and lateral
sides, according to the method of Maglio (1973). In the M
3
s, LF can
be measured only at the crown base and therefore gives a lowvalue
(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 is
2.7e2.8 mm for the M
3
s and 2.9e3.1 mm for the M
3
s.
A nal metric of taxonomic signicance is the crown height of
unworn plates. Due to wear, in the M
3
s (Fig. 15c) this cannot be
estimated at all, and in the M
3
s is measurable only in the posterior
part of the crown. The maximum crown height in the right M
3
is
measured on the third plate from the posterior end (platelet
excluded), which is just coming into wear and measures 136 mm,
giving a hypsodonty index 1.24 relative to the maximum crown
width of 110 mm. This value, however, is clearly far below the full
crown height of the molar which would have been maximal in the
anterior 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 at
least 1.6.
These parameters strongly indicate referral of the Kostolac
mammoth to the species M. trogontherii. Identity with the ancestral
mammoth M. meridionalis is excluded by the plate count of more
than 16 and the reconstructed high crown, as well as by the
elevated lamellar frequency (cf. Table 2). Plate counts of 20 can
occur at the low end of the range of variation within the woolly
mammoth M. primigenius, but the lamellar frequency at Kostolac is
too low, and the enamel too thick, for referral to that species. Body
size is a malleable feature which should be used with care in
taxonomic assignment, but the very large size of the Kostolac
skeleton (see below) is more conformable with M. trogontherii
(especially earlier European populations) than M. primigenius
which is generally smaller.
6.2. Body size and mass
Measurement data for the postcranial bones are given in
Tables 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 other male
skeletons 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 on
a series of mounted skeletons. The three most reliable estimators,
humerus, radius and femur lengths, give skeletal shoulder heights
for the Kostolac skeleton of 369, 365 and 386 cm, respectively, with
an average of 374 cm (Table 13). Different authors allow 5e7% extra
for esh; at 6%, the Kostolac skeletal height of 374 cm gives a value
of ca. 396 cm. We suggest four metres as an approximate estimate
for the live shoulder height of the Kostolac Mammoth.
The body mass of the Kostolac mammoth can be estimated in
two ways. The rst method uses regression equations of body mass
against shoulder height for living elephants. Using the 396 cm
shoulder height estimated above, Roths (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 Kostolac
mammoth (Table 14). A second method is the use of individual
bone lengths for estimating body mass. Using a combined sample
of modern Elephas maximus and L. africana, Christiansen (2004)
derived prediction equations of body mass. Table 15 shows the T
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5
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 138
results of applying these equations to limb bone lengths of the
Kostolac mammoth. Estimates range from8 to 11 t, with an average
across bones of 9.4 t. The concordance between the mass estimate
based on shoulder height, and those based on limb bone lengths, is
excellent, and indicates a live weight for the Kostolac mammoth of
ca. 9.5 t. It should be remembered, however, that in living
elephants, an animals body mass can vary by up to 10% in the short
term, depending on daily cycles of feeding, drinking, and defe-
cating, and even more on a seasonal basis, due to food availability
and reproductive cycles such as musth (males) and pregnancy
(females). The estimated gure should be regarded as a central
estimate 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), based
on mandibular teeth. The M
3
s of the Kostolac mammoth
(Fig. 15a,c) have only the posterior 11 plates remaining, all of
which 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 plate
number of around 20 (see above), this indicates that ca. 11/20 or
55% of the plate count remains, equivalent to ca. 6 plates of
L. africana with a typical full plate number of 11 in M
3
(Laws, 1966).
By reference to Laws scheme, this indicates age class XXIX, or an
age of around 57 in African elephant years. A correction can be
made for the larger body size in M. trogontherii, which scales with
positive allometry to longevity among mammals. By applying the
formula of Blueweiss et al. (1978) (longevity in days a body mass
in grams
0.17
), and using a longevity of 60 years and a mass of 6 t for
living bull elephants, and the body mass of 9.5 t for the Kostolac
mammoth as above, a longevity of 65 years is obtained. An esti-
mated absolute age for the Kostolac mammoth is therefore
57 65/60 62 years.
Consistent with this estimate of advanced age, all limb-bone
epiphyses on the Kostolac mammoth are fused to their diaphysis
shafts. This includes the scapular margin, the proximal femur and
the distal radius/ulna, which are the last to fuse in both living
elephants and mammoths (Roth, 1984; Lister, 1999). In the skull, the
zygomatic 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 from
several lines of evidence. First, the large size of the skeleton,
comparable to known male M. trogontherii individuals such as
West Runton and Steinheim (Fig. 16), suggests male gender, since
there is clear sexual dimorphism in body size among both living
elephants and mammoths (e.g. Averianov, 1996). Skull robustness
is also markedly stronger in male individuals, but the breakage of
the Kostolac cranium makes this difcult to assess. The size of the
tusks, however, provides a further clue. Two loose chunks of tusk
from the Kostolac excavation indicate an original diameter,
Table 3
Measurements 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 spine
from top of neural canal
Maximum
width
Posterior
articular width
Posterior
centrum width (m-l)
Posterior
centrum height (d-v)
Centrum
length (a-p)
Atlas (1st cervical) 514
Axis (2nd cervical) 182
Third thoracic 560 149 163 98
Fourth thoracic 570 154 163 100
Fifth thoracic 580 363 133 93
Sacrum (5 fused vertebrae) 500
Table 4
Measurements 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 dorsal
epiphysis) TOT
Total length
(excluding dorsal
epiphysis) DIA
Length from
dorsal edge to
closest edge of
glenoid cavity
Glenoid to
dorso-posterior
corner of scapula,
parallel to
posterior edge
Maximum width
of glenoid epiphysis
Articular width
of glenoid
(a-p) LGA
Articular depth
of glenoid
(m-l) TSA
L scapula 321 219 152
R scapula 1110 900 1075 685 334 230 151
Table 5
Measurements of humeri of the Kostolac mammoth (mm). Equivalent measurement codes from Roth (1982) are given. m-l, medio-lateral.
Element Total length LONG Articular length ARTS Width of proximal
epiphysis (m-l)
Minimum diaphysis
width TSM
Width of distal
epiphysis TSD
Width of distal
condyle TTC
L humerus 352 277
R humerus 1215 1170 343 160 364 273
Table 6
Measurements of ulnae of the Kostolac mammoth (mm). Equivalent measurement codes from Roth (1982) are given. m-l, medio-lateral.
Description Total length Articular length Width of proximal
epiphysis (m-l)
Proximal articular
width (m-l)
L ulna
R ulna 1080 875 ca. 330 279
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 139
presumably close to the base, of around 200 mm. In a large sample
of adult M. primigenius tusks from Siberia, Vereshchagin and
Tikhonov (1986) give a range of 47e90 mm in females and
89e180 mm in males for tusk diameter at the base. The male
M. trogontherii from West Runton (UK), Azov I (Russia) and
Steinheim (Germany) have diameters of 210, 215 and 217 mm,
respectively (Dietrich, 1912; Baigusheva and Garutt, 1987; Lister
and Stuart, 2010). Conversely in the female M. trogontherii from
Edersleben (Germany) and Novogeorgievsk (Ukraine), the gures
are 138 and ca. 149 mm, respectively (Zakrevska, 1935; Garutt and
Nikolskaya, 1988). These data support the male gender of the
Kostolac M. trogontherii.
Most diagnostic is the form of the pelvic girdle. In Mammuthus,
the pelvic aperture is relatively smaller, and the neck of the ilium
relatively wider, in males than in females, and the sexes can be
separated by the ratio between these two measurements. The
result holds true for M. primigenius, M. meridionalis and Mammu-
thus columbi, and is conformable in available individuals of
M. trogontherii (Lister and Agenbroad, 1994; Lister, 1996a; Lister and
Stuart, 2010). In the Kostolac pelvis (Fig. 17), the ratio between
transverse aperture width and neck width is 540/244 2.21, while
between vertical aperture width (measured from the sacral scar to
the ventral midline) and neck width it is 450/244 1.84. These
values clearly fall within the male range of other Mammuthus
Table 8
Measurements of pelvic bones of the Kostolac mammoth (mm). L are codes from Lister (1996a).
Element Maximum
transverse width
of pelvic girdle L1
Diagonal height
of aperture from
sacral scar to
ventral midline L2
Maximum
transverse diameter
of pelvic aperture L3
Width of ilium
from lateral
maximum to
nearest point
of aperture L4
Minimum width
of ilium L5
Maximum width
of ilium
Antero-posterior
diameter of
acetabulum
Pelvis 1670 450 540 630 244 1040 192
Table 9
Measurements of femora of the Kostolac mammoth (mm). Equivalent measurement codes from Roth (1982) are given. m-l, medio-lateral.
Element Total length articular length Minimum (along shaft)
maximum (around shaft)
diaphysis width APM
Proximal width
(m-l) TSW
Width of femoral
head (m-l) HDD
Width of distal
epiphysis (m-l) TSI
Width of distal
articulation
L femur 1470 ca. 196 437 187 301 261
R femur 1470 186 450 298 271
Table 10
Measurements 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 of
medial proximal condyle
Minimum (along shaft)
maximum (around shaft)
diameter of diaphysis
Width of proximal
epiphysis (m-l) TSM
L tibia 850 755 140 273
R tibia 855 775 130
Table 11
Measurements of bulae 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 distal
epiphysis, perpendicular to
long axis of distal end (m-l)
Depth of distal epiphysis,
parallel to long axis
of distal end (a-p)
L bula 805 49 130 ca. 68
R bula 790 49 130 ca. 98
Table 12
Measurements of foot bones and patella of the Kostolac mammoth (mm). Astragalus
measured parallel to ventral surface; calcaneum measured parallel or perpendicular
to main vertical medial facets. d-v, dorso-ventral; a-p, antero-posterior; m-l, medio-
lateral.
Description Maximum
length (d-v)
Maximum
depth (a-p)
Maximum
width (m-l)
L patella 137
R patella 164
R astragalus ca. 186
L calcaneum 266 195 165
R calcaneum ca. 260 194 176
Table 7
Measurements 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 154 138 108
R radius 925
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 140
Limb size and proportion of Mammuthus species
scapula humerus radius femur tibia fibula
60
70
80
90
100
110
120
Nogaisk
Odessa
Azov I Georgievsk
Steinheim
Scoppito
KOSTOLAC
West Runton
Farneta
Durfort
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) for
raw 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 gure legend, the reader is referred to the web version of this article.)
Table 13
Estimation of Kostolac mammoth shoulder height fromlengths of long bones, based on linear regression equations derived frommounted skeletons of Mammuthus (Lister and
Stuart, 2010). a, b slope and intercept of regression equation.
Element Equation R-square 95% condence interval/
prediction interval
Kostolac skeletal shoulder
height estimate / 95%
condence/prediction interval (cm)
Kostolac live shoulder
height (cm), adding
6% for esh
All humerus a 2.9701, b 85.4976 0.8613 3.29/11.49 369 12.1/42.4 391
Best radius a 4.0195, b 67.6722 0.9018 3.19/10.57 365 11.6/38.6 387
All femur a 2.9458, b 438.9187 0.8861 3.09/10.27 386 11.9/39.6 409
Average 396
Table 14
Estimation of body mass (in kg) of the Kostolac mammoth based on predictive equations fromshoulder height of modern African elephant populations. Roths (1990) data from
Laws et al. (1975), Johnson and Buss (1965), Tumwasorn et al. (1980) and others; Christiansens (2004) data from Benedict (1936) and others. m mass (kg), sh shoulder
height (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 5.07*10
4
sh
2.803
9691
L. africana males, Uganda e m 3.06*10
4
sh
2.890
9841
L. africana males, Uganda f m 1.81*10
4
sh
2.97
9394
Average 9642
Table 15
Estimation 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) a b (log X), where X is bone variable (mm). %SEE and %PE are, respectively, the standard error and percent
prediction 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 9613
Radius 3.838 2.634 10.11/6.64 925 9437
Ulna 4.135 2.674 8.41/5.34 1080 9471
Femur 5.568 3.036 14.54/6.15 1460 10,939
Tibia 3.064 2.378 12.47/6.93 852 8028
Fibula 3.086 2.422 18.68/11.4 798 8763
Average 9375
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 141
(Fig. 18), and corroborate the value of this feature for gender
determination in M. trogontherii.
6.4. Further morphological features
Damage to the cranium precludes a thorough consideration of
its anatomy, but one visible feature is of interest. The zygomatic
process of the maxillary forms, in its lower part, a bony bridge
bounding the infraorbital canal. This infraorbital process is prim-
itively robust among elephantids, but is narrower and thinner in
Mammuthus, 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
point), but quite thin e this apparently mosaic condition between
primitive and derived conditions within Mammuthus (Lister and
Stuart, 2010) might reect the relatively early age of W. Runton
(ca. 700 ka) within European M. trogontherii. Alternatively the
difference of form between the Kostolac and W. Runton specimens
might simply reect individual variation; further material is
required to test between these alternatives.
Only parts of the mandible are preserved at Kostolac, but the
anterior portion shows a relatively short symphysis and modest
rostrum, conforming to the advanced or derived condition within
Mammuthus, similar to M. primigenius and most M. trogontherii, and
contrasting with the longer symphysis of most M. meridionalis (cf.
Lister and Stuart, 2010). Ferretti and Debruyne (2011) discuss the
taxonomic distribution of the anterior mandibular foramina. For
the lateral mental foramina, 86.4% of M. meridionalis specimens
(n 22) possessed 1 or 2 foramina on each hemi-mandible, and
only 13.6% possessed 3 or more. In M. primigenius, by contrast
(n 54), the gures are 44.4% and 55.6%, respectively. Average
frequency therefore increased between these taxa. The Kostolac
mandible shows two foramina on the lateral side, as does the West
Runton M. trogontherii mandible, a condition commonly also found
in the other two species; a larger sample of M. trogontherii would be
required to test if the species is statistically intermediate.
The presence of a mental foramen on the medial side of the
ramus 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 of
Mammuthus rumanus, but present in 73% of M. meridionalis
(n 30), 69% of M. trogontherii (n 18), and 100% of M. primigenius
(n 173). The frequency of the foramen therefore increases
through the sequence. In the Kostolac mandible, the right ramus
shows a single pit (14 20 mm) leading to two foramina. On the
left ramus, there is a single wide foramen leading into the
mandibular canal that can be seen due to breakage of the
mandibular wall. The presence of the foramen at Kostolac and also
Fig. 17. Pelvic girdle of the Kostolac mammoth in posterior view. The relatively small
pelvic canal and wide ilium signify male gender.
1.4 1.6 1.8 2.0 2.2 2.4 2.6 2.8 3.0 3.2
pelvic aperture ratio
E
d
e
r
s
l
e
b
e
n
N
o
v
o
g
e
o
r
g
i
e
v
s
k
K
O
S
T
O
L
A
C
S
i
e
g
s
d
o
r
f
L
i
a
k
h
o
v
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r
a
z

R
o
d
e
t
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a
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r
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e
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a
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e
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e
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o
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e
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i
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y
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R
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o
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n
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e
i
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h
e
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a
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h
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n
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a
l
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a
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a
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d
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r
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a
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a
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M. meridionalis
MALES FEMALES
M. trogontherii
M. primigenius
Fig. 18. Ratio between pelvic aperture width and ilium shaft width. Open symbols: males; lled symbols: females. For the majority of specimens, independent evidence of gender is
provided by cranial morphology, tusk size or body size (see Lister, 1996a). The identity of the female Edersleben (Germany) skeleton as M. trogontherii or M. meridionalis is debated
(Lister, 1996b; van Essen, in litt.). Based on original measurements; the ratio of the female Novogeorgievsk specimen may be slightly exaggerated as measurements taken from
a photograph (Zakrevska, 1935).
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 142
at West Runton (Lister and Stuart, 2010) increases the proportion in
M. trogontherii to 73% (n 20), identical to that of M. meridionalis.
The vertebral column of the Kostolac skeleton is the most
complete known in M. trogontherii. Another, in the cf.
M. trogontherii skeleton from Novogeorgievsk (Ukraine), is stated
by Zakrevska (1935) to possess 19 preserved thoracic vertebrae and
four sacrals. At Kostolac, there are 7 cervical vertebrae, 19 thoracics,
four lumbars, and a sacrum comprising ve fused vertebrae. Eigh-
teen thoracic vertebrae have been found in some skeletons of
M. primigenius (e.g. Berezovka: Zalensky, 1903), living elephants
typically have 20 (Mol and van Essen, 1992), and variability in
M. trogontherii is uncertain.
The two proximal caudal vertebrae are preserved, and six others,
incomplete and separated from the skeleton. The combined length
of these eight vertebrae is approximately 80 cm, indicating
a complete tail substantially longer than this. This suggests
a signicantly longer tail than in M. primigenius, where the total
length of the tail vertebrae in the Beresovka carcass was measured
as 60 cm, with the tail in the esh 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 possible
to assess the structure of the carpus, which shows signicant
variation within the genus Mammuthus (Lister, 1996b). An inter-
esting feature, however, is seen in the left hind foot, where the
metatarsal of the rst digit is complete and well preserved. The
distal end of mtI shows no articular surface for the attachment of
a rst phalange, indicating that there are only four toes on the hind
foot in this specimen, as observed frequently in other probosci-
deans. On the caudal side of the distal end a small articular surface
can be noted for the attachment of a sesamoid.
7. Geological age of the remains
The geological context, and limited other fauna found in the
mammoth layer, do not, at the present state our regional under-
standing, provide evidence on the precise age of the skeleton.
The identication of the Kostolac skeleton as M. trogontherii
does, however, place some constraints on the age of the deposit. In
Europe this species is characteristic of the early Middle Pleistocene
(Cromerian Complex), where it is well-known from localities
including West Runton (UK), Sssenborn and Mosbach (Germany)
and Tiraspol (Moldova), spanning approximately MIS 19e13 (ca.
780e500 ka). At some localities it has been identied earlier,
together with the more primitive M. meridionalis, such as at Dorn-
Durkheim, Germany (just belowthe Brunhes/Matuyama boundary)
and at Sinyaya Balka, Russia, in deposits ca. 1.0 Ma in age (Lister
et al., 2005).
After the Elsterian/Anglian glaciation (MIS 12), mammoths in
Europe showa transition fromM. trogontherii to M. primigenius. The
remains from Steinheim, Germany, considered to be MIS 11e10 in
age (ca. 400e350 ka) (Schreve and Bridgland, 2002), are dentally
similar to earlier M. trogontherii (and to Kostolac). They are also of
similarly large body size: at 370 cm, the reconstructed shoulder
height of the famous Steinheimmammoth skeleton (Dietrich, 1912)
is only slightly smaller than that of Kostolac (Fig. 16). Later European
M. trogontherii populations, while dentally still more primitive than
typical M. primigenius, were considerably reduced in size. A likely
age 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 signicantly to our
knowledge of the osteology of this species, important as an
evolutionary intermediate between the better-known
M. meridionalis and M. primigenius, between which there was
substantial evolutionary change (Lister et al., 2005). The Kostolac
skeleton provides newinformation in features such as the vertebral
count and in details of the skull and feet.
The skeleton is also the only one of M. trogontherii preserved in
a largely articulated condition, and is very unusual for any fossil
elephantid skeleton in being preserved in the animals likely death
posture, a remarkable taphonomical situation that invites further
study.
Finally, as the rst complete skeleton of this species found in the
Mediterranean basin, and with further isolated remains found in
proximity, the nds highlight the potential importance of the Kos-
tolac area for our understanding of the regional history of the
Quaternary mammal fauna. Further investigations are likely to yield
associated fauna of biostratigraphic and ecological signicance.
Acknowledgements
We are grateful to the excavators of the Roman site of Vimina-
cium for their fast response and sincere delight with an animal
much older than their period of interest. We especially thank Dr
Miomir Korac, director of Viminacium site, for organizing the
excavation, supporting the conservation project, and building
a protective cover over the skeleton. Special thanks go to Milos
Milivojevic, 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 for
photography. Thanks are also due to Marco Ferretti and Ralf Kahlke
for their rapid and valuable comments on the manuscript, and to
Frdric Lacombat and Norm Catto for accommodating our paper
within this special issue of Quaternary International. V.D.s contri-
bution forms part of the project Bioarchaeology of Ancient
Europe e humans, animals and plants in the prehistory of Serbia
(III47001) funded by the Ministry of Education and Science of the
Republic of Serbia.
References
Averianov, A.O., 1996. Sexual dimorphism in the mammoth skull, teeth, and long
bones. In: Shoshani, J., Tassy, P. (Eds.), The Proboscidea: Trends in Evolution and
Paleoecology. OUP, Oxford, pp. 260e267.
Baigusheva, V.S., Garutt, V.E., 1987. A skeleton of the steppe elephant Archidiskodon
trogontherii (Pohlig, 1885) from the north-east Azov Sea region. Proceedings of
the Zoological Institute (Leningrad) 168, 21e36.
Beard, P., 2000. The End of the Game. Chronicle Books.
Beden, M., 1979. Les lphants (Loxodonta et Elephas) dAfrique orientale: systm-
atique, phylognie, intret biochronologique. Ph.D. Thesis, Poitiers.
Benedict, F.G., 1936. The Physiology of the Elephant. Publication no. 474. Carnegie
Institute of Washington, Washington, D.C.
Blueweiss, L., Fox, H., Kudzma, V., Nakashima, D., Peters, R., Sams, S., 1978. Rela-
tionships between body size and some life history parameters. Oecologia 37,
57e272.
Christiansen, P., 2004. Body size in proboscideans, with notes on elephant metab-
olism. Zoological Journal of the Linnean Society 140, 523e549.
Dietrich, W.O., 1912. Elephas primigenius Fraasi, eine schwbische Mammutrasse.
Jahreshefte des Vereins fur Vaterlndische Naturkunde in Wrttemberg 68,
42e106.
Ferretti, M.P., Debruyne, R., 2011. Anatomy and phylogenetic value of the mandib-
ular and coronoid canals and their associated foramina in proboscideans
(Mammalia). Zoological Journal of the Linnean Society 161, 391e413.
Garutt, V.E., Nikolskaya, V.N., 1988. Uber das Skelett vom Steppenelefant aus Eder-
sleben. Beitrge zur Heimatforschung, Sangerhausen, Spengler Museum9, 3e13.
Herz, O., 1902. Berichte des Leiters der von der Kaiserlichen Akademie der Wis-
senschaften zur Ausgrabung eines Mammuthkadavers an der Kolyma-
Beresowka ausgesandten Expedition. Imperial Academy of Science, St. Peters-
burg, 1e38 pp.
Jachmann, H., 1988. Estimating age in African elephants: a revision of Laws molar
evaluation technique. African Journal of Ecology 26, 51e56.
Johnson, O.W., Buss, I.O., 1965. Molariform teeth of male African elephants in
relation to age, body dimensions and growth. Journal of Mammalogy 46,
373e384.
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 143
Korac, M., Markovic, Z., Milivojevic, M., Mol, D., 2010. The Kostolac mammoth,
Serbia: an almost entirely complete skeleton of the steppe mammoth, Mam-
muthus trogontherii. Quaternaire, Hors Srie 3, 79e80.
Laws, R.M., Parker, I.S.C., Johnstone, R.C.B., 1975. Elephants and Their Habitats: The
Ecology of Elephants in North Bunyoro, Uganda. Clarendon Press, Oxford.
Laws, R.M., 1966. Age criteria for the African elephant, Loxodonta a. africana. East
African Wildlife Journal 4, 1e37.
Lister, A.M., 1996a. Sexual dimorphism in the mammoth pelvis: an aid to gender
determination. In: Shoshani, J., Tassy, P. (Eds.), The Proboscidea: Trends in
Evolution and Paleoecology. OUP, Oxford, pp. 254e259.
Lister, A.M., 1996b. Evolution and taxonomy of Eurasian mammoths. In: Shoshani, J.,
Tassy, P. (Eds.), The Proboscidea: Trends in Evolution and Paleoecology. OUP,
Oxford, pp. 203e213.
Lister, A.M., 1999. Epiphyseal fusion and postcranial age determination in the
woolly mammoth, Mammuthus primigenius (Blum). Deinsea 6, 79e88.
Lister, A.M., 2009. Late-glacial mammoth skeletons (Mammuthus primigenius) from
Condover (Shropshire, UK): anatomy, pathology, taphonomy and chronological
signicance. Geological Journal 44, 447e479.
Lister, A.M., Agenbroad, L.D., 1994. Gender determination of the hot springs
mammoths. In: Agenbroad, L.D., Mead, J.I. (Eds.), The Hot Springs Mammoth
Site: A Decade of Field and Laboratory Research in Paleontology, Geology and
Paleoecology. Mammoth Site Inc., Hot Springs, SD, pp. 208e221.
Lister, A., Bahn, P., 2007. Mammoths: Giants of the Ice Age. Frances Lincoln, London.
Lister, A.M., Sher, A.V., van Essen, H., Wei, Guangbiao, 2005. The pattern and process
of mammoth evolution in Eurasia. Quaternary International 126-128, 49e64.
Lister, A.M., Stuart, A.J., 2010. The West Runton mammoth (Mammuthus trogon-
therii) and its evolutionary signicance. Quaternary International 228,
180e209.
Maglio, V.J., 1973. Origin and evolution of the Elephantidae. Transactions of the
American Philosophical Society, New Series 63, 1e149.
Milosevic, S., Miletic, R., 1975. Kostolacki basen. In: Petkovic, K. (Ed.), Geologija Srbije,
VII, Kaustobioliti, 144e149. Zavod za regionalnu geologiju i paleontologiju
Rudarsko-geoloskog fakulteta, Univerzitet u Beogradu, Beograd, pp. 144e149.
Mol, D., van Essen, H., 1992. De Mammoet: Sporen uit de IJstijd. BZZTH, s-
Gravenhage.
Rakic, M., 1978. Tumac Za OGK SFRJ, list Bela Crkva 1: 100 000. Savezni geoloski
zavod, Beograd.
Roth, V.L., 1982. Dwarf Mammoths from the santa Barbara, California Channel
Islands: Size, Shape, Development and Evolution. D.Phil. thesis, Yale University,
University Microlms, Ann Arbor.
Roth, V.L., 1984. How elephants grow: heterochrony and the calibration of devel-
opmental stages in some living and fossil species. Journal of Vertebrate Palae-
ontology 4, 126e145.
Roth, V.L., 1990. Insular dwarf elephants: a case study in body mass estimation and
ecological inference. In: Damuth, J., MacFadden, B. (Eds.), Body Size in
Mammalian Paleobiology: Estimation and Biological Implications. Cambridge
University Press, Cambridge, pp. 151e179.
Schreve, D.C., Bridgland, D.R., 2002. Correlation of English and German Middle
Pleistocene uvial sequences based on mammalian biostratigraphy.
Netherlands Journal of Geosciences/Geologie en Mijnbouw 81, 357e373.
Sher, A.V., Garutt, V.E., 1987. New data on the morphology of elephant molars.
Transactions of the academy of sciences of the USSR. Earth Science Section 285,
195e199.
Shoshani, J., 1991. Anatomy and physiology. In: Rogers, G., Watkinson, S. (Eds.), The
Illustrated Encyclopaedia of Elephants. Crescent Books, New York, pp. 30e47.
Stevanovic, P., 1951. Donji pliocen Srbije i susednih oblasti e Posebna izdanja SAN.
CLXXXVII, Geoloski institut, 2, Beograd.
Stevanovic, P., 1977. Oblast zapadno do Karpatobalkanida. In: Petkovic, K. (Ed.),
Geologija Srbije, II-3 (Stratigraja, kenozoik). Zavod za regionalnu geologiju i
paleontologiju Rudarsko-geoloskog fakulteta. Univerzitet u Beogradu, Beograd,
pp. 72e74.
Stevanovic, P., 1990. Pontien sudlich von der Sava und Donau in Serbien und Bos-
nien. Cronostratigraphie und Neostratotypen JAZU and SANU, Zagreb-Beograd,
119e153.
Sukumar, R., 2003. The Living Elephants: Evolutionary Ecology, Behaviour, and
Conservation. OUP, Oxford.
Tumwasorn, S., Udsanakornkul, S., Leenanuruksa, D., Kaeophrommarn, C.,
Pichaicharnnarong, A., 1980. Some weight and body measure estimates of
Asiatic elephants (Elephas maximus). Thai Journal of Agricultural Science 13,
179e185.
Vereshchagin, N.K., Tikhonov, A., 1986. A Study on Mammoth Tusks. Proceedings of
the Zoological Institute. USSR Academy of Sciences. 149, 3e14. (in Russian).
Zakrevska, G., 1935. LElephas trogontherii Pohl. de la rive droite du Dnipro moyen,
Memoirs of the Institute of Geology. Ukranian Academy of Sciences, 5. Kiev, pp.
1e138. (in Ukrainian, French summary).
Zalensky, V.V., 1982. Osteological and Odontographic Studies on the Mammoth
(Elephas primigenius Blum.), and the Elephants (El. Indicus L. and El. Africanus
Blum). Scientic Results of the Imperial Academy of Sciences Expedition to
Excavate the Mammoth Carcass Found by the Beresovka River in 1901, vol. 1.
Imperial Academy of Sciences, St. Petersburg, 1e124 pp.
A.M. Lister et al. / Quaternary International 276-277 (2012) 129e144 144