Athanassiou, 2012 - M. Trogontherii Peloponnese

A skeleton of Mammuthus trogontherii (Proboscidea, Elephantidae) from NW
Peloponnese, Greece
Athanassios Athanassiou
Hellenic Ministry of Culture, Ephorate of PalaeoanthropologyeSpeleology, Ardittou 34B, 11636 Athens, Greece
a r t i c l e i n f o
Article history:
Available online 2 April 2011
a b s t r a c t
Fossil elephant remains were identied in Loussik, NW Peloponnese, Southern Greece, when tusk
fragments were recognized in a bulldozer backll. An excavation carried out in 2001 and 2003 by the
Hellenic Ministry of Culture revealed the partial skeleton of an adult male mammoth, referred to the
Middle Pleistocene species Mammuthus trogontherii. The recovered material includes part of the skull,
the complete mandible, several vertebrae and ribs, both scapulae, ulna, tibia, most carpal and tarsal
bones, metapodials and phalanges. The metrical and anatomical study of the skeleton shows that the
living individual was about 45 years old, stood about 3.80 m high and weighed about 8 t. M. trogontherii
is a very rare species in Southern Europe. The Loussik skeleton represents the rst solid evidence of the
species presence in Southern Greece and considerably expands to the south its palaeobiogeographic
range in the Balkan area.
2011 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
Elephant fossils are common in palaeontological sites, as their
robust bones and their massive teeth have relatively good preserva-
tion potential in the fossil record, even in relatively higher energy
environments. This resulted in a fairly good fossil record, recovered
from numerous localities, that contributed to the current under-
standing of evolution and phylogeny within the family Elephantidae.
In mainland Greece, the recorded fossil elephant-bearing local-
ities are close to forty (Doukas and Athanassiou, 2003), commonly
situated in uvial and lacustrine basins. The straight-tusked
elephant Elephas antiquus is the dominant species, present in more
thantwenty sites datedto the MiddleeLate Pleistocene (Doukas and
Athanassiou, 2003; Tsoukala et al., 2010). Older, Lower Pleistocene
localities have yielded numerous remains of the primitive
mammoth Mammuthus meridionalis, often in association with the
gomphothere Anancus arvernensis. More advanced mammoth nds
are, however, very rare and they are geographically restricted to the
northern part of Greece. The insular proboscidean faunas, known
from more than thirty sites, are characterised by the presence of
dwarf endemic forms, derived mainly from continental E. antiquus.
The new site Loussik, described in the present paper, is situ-
ated in western Achaia, NW Peloponnese (Fig. 1). It was discovered
in 1988 by the archaeologist Andreas Darlas, while prospecting the
area around the village of Loussik for prehistoric artefacts. There,
in the southern slope of the Serdin valley, he noticed the presence
of tusk and bone fragments in a bulldozer backll. The bulldozer,
while levelling the ground for agricultural use, had cut across the
proximal part of the skull, destroying the sole preserved right tusk,
as well as an unknown number of bones. The site was excavated
several years later, in 2001 and 2003, by the Ministry of Culture
(Ephorate of PalaeoanthropologyeSpeleology, Athens, and ST
Ephorate of Prehistoric and Classical Antiquities, Patras) under the
direction of Dr. A. Darlas, with the participation of the present
author. The excavation revealed a partial proboscidean skeleton
that included the skull, the mandible, and part of the axial and
appendicular skeleton. The recovered specimens were prepared in
the laboratory of the Ephorate of PalaeoanthropologyeSpeleology
in Athens and are currently stored in the collections of the
Archaeological Museum of Patras. The skull still remains in Athens
because of its fragility and demanding preparation, and it will be
also transported to Patras when the preparation and consolidation
are nished.
The elephant skeleton of Loussik is referred here to the Middle
Pleistocene steppe mammoth Mammuthus trogontherii (Pohlig,
1885). However, the nding was preliminarily referredto E. antiquus
Falconer and Cautley, 1847 (Doukas and Athanassiou, 2003;
Athanassiou, 2010), a quite common Pleistocene elephant species,
mainly because of a misinterpretation of the skull morphology
during the excavation (see Section 7). After the plaster jacket
removal, the preliminary preparation and re-examination of the E-mail address: aathanas@geol.uoa.gr.
Contents lists available at ScienceDirect
Quaternary International
j ournal homepage: www. el sevi er. com/ l ocat e/ quai nt
1040-6182/$ e see front matter 2011 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2011.03.030
Quaternary International 255 (2012) 9e28
skull and dental remains, it became evident that the Loussik skel-
eton belongs to M. trogontherii, a very rare species in Greece, known
until nowonly from scanty dental material.
M. trogontherii is usually considered as an intermediate form in
the Mammuthus lineage, temporally and evolutionary placed
between the species M. meridionalis (Nesti, 1825) and Mammuthus
primigenius (Blumenbach, 1799) (Garutt, 1964; Maglio, 1973;
Dubrovo, 1977; Lister, 1996a). An anteroposterior shortening of
the skull and mandible, and an increase of hypsodonty and molar
plate number are the main trends along this evolutionary lineage,
which in a short time interval (less than 2.5 million years) is
thought to have more or less gradually transformed a woodland
dwelling generalist (M. meridionalis) to a highly specialised, cold-
adapted grazer (M. primigenius). However, evidence published
fairly recently (Lister and Sher, 2001; Lister et al., 2005; Wei et al.,
2006, 2010) indicate that this simple process is less plausible, at
least for Europe, as the Mammuthus species appear much earlier in
the East Asian fossil record. This implies repeated migration of new
taxa from Asia to Europe, where they replaced the existing species,
possibly after a short period of coexistence. This theory, though
implying a complicated evolutionary pattern, is in accordance with
some older published opinions (e.g. Depret et al., 1923; Osborn,
1942; Azzaroli, 1977) that favoured repeated immigrations of new
forms into Europe as the cause of Mammuthus species turnover.
Apart fromthe mammoth skeletal elements, the locality yielded
also two tortoise plastron fragments, as well as a cervid antler
fragment and a ruminant distal phalanx; the latter is attributable to
a small deer, the size of Capreolus. Moreover, a coprolite was found
among the elephant bones, suggesting the presence of a carnivore
(see Section 4.1).
2. Material and methods
The excavated material was compared to recent Elephas maximus
osteological material belonging to the Museumof Palaeontology and
Geology, University of Athens, in order to determine the anatomical
position of each specimen. The osteological description of Loxodonta
africana, published in a series of papers by Smuts and Bezuidenhout
(1993, 1994), van der Merwe et al. (1995), Bezuidenhout and
Seegers (1996), as well as illustrated fossil skeletons descriptions
(e.g. Trevisan, 1954; Kroll, 1991), were also of great help.
The broken or fragile specimens were glued together and/or
consolidated using Paraloid B72, a highly reversible acrylic polymer
(resin), which is commonly used in fossil bone preparation and
conservation.
All measurements were carried out using callipers or measuring
tape (for the larger specimens). Inaccurate measurements, because
of incomplete preservation, distortion or inaccessibility, that can be
reliably estimated are given in brackets. For molar measurements
and calculation of indices I followed Maglio (1973); the lamellar
frequency parameter is, however, measured only labially on the
upper molars, and labially and lingually on the lower molars,
because all studied teeth are in situ, not allowing measurements at
their base. The lingual side of the upper molars is also not acces-
sible, because of the presence of supporting material. The bone
measurements are anatomically oriented, i.e. parallel or perpen-
dicular to the sagittal plane. DAP stands for anteroposterior
diameter; DT stands for transverse diameter; the height is
measured dorsoventrally, except for the scapula, the metapodials
and the phalanges, were it is meant proximodistally (as the
anatomical orientation of these bones is not vertical). Any
measurement technique that deviates from this general scheme is
described in the relevant table. The upper molars are indicated by
M, the lower ones by m.
3. Geological setting
The new locality Loussik is situated in the valley of the small
stream Serdin, a tributary of the river Peros, which is the major
stream of the drainage basin. The fossil-bearing deposits are of
uvial origin. They consist of clayey sand, sand and coarse sand,
often exhibiting cross bedding (Fig. 2), and belong to a depositional
terrace of the Peros River system, occurring at an altitude of about
50 m above sea level. The rather ne-grained sediment suggests
a low-energy uvial environment. At the top of the section there is
a calcrete-rich palaeosol and the recent soil. The terrace deposits
overlie unconformably a Pliocene shallow-marine sequence, which
covers the wider area of western Achaia, and consists of sandstones,
sandy clays and marls, rich in invertebrate fossils, mainly bivalves
and gastropods (Tsoias and Fleury, 1980). The basement consists
of Late EoceneeOligocene ysch that belongs to the Tripoli
geotectonic zone of the Alpine orogene. South of the studied area
there are also some outcrops of Cretaceous limestones of the same
geotectonic zone.
4. Taphonomy
The excavation extended in an area 6 m long (EeW) to 3 mwide
(NeS), that is 18 m
2
. It yielded most of the cranial and axial skel-
eton, except for the sacrum, the pelvis and all but one caudal
Fig. 1. Topographical map of W. Achaia (SW Greece) showing the geographic position of the new locality Loussik (asterisk; 38
5
0
53.8
00
N, 21
35
0
32.1
00
E, altitude 45 m, WGS84
datum) south of the homonymous village. Contour interval: 100 m. Graphical scale: 3 km.
A. Athanassiou / Quaternary International 255 (2012) 9e28 10
vertebrae. Most of the long bones are, though, missing, except for
an ulna and a tibia. The preservation of the Loussik skeleton is
generally good, with the exception of the skull, which suffers from
erosion and root weathering.
The recovered specimens were not in anatomical association but
totally disarticulated and dispersed; they were scattered mainly in
an NEeSW direction, in a layer of ne clayey uvial sand (Fig. 3).
Elements that are anatomically remote fromeach other were found
lying close together: e.g. the right tibia was lying to the left of the
skull, the left ulna to the right of the skull, while the left calcaneus
was found under the right tusk sheath. Other associated elements,
as the mandible and the skull, were, though, not widely separated.
The recovering of delicate hyoid bones from the ventral side of the
skull indicates minimal, if any, translocation of the skull (the hyoid
apparatus attaches to the skull through a cartilage and it is easily
lost or destroyed during decomposition of the carcass and burial).
No specimen shows signs of edge rounding because of rolling, again
strongly suggesting minimal or no transportation.
The skull was found in upright position, sitting on its ventral
side (which may imply an original sudden death posture, i.e. the
animal lying on its belly; Haynes, 1988). The mandible, broken at
the symphysis level, was found considerably lower (40e70 cm)
than the skull, with its rostrum facing SSW, at the opposite direc-
tion than the skull (Figs. 3 and 4). The depositional level difference
indicates rapid sediment deposition, covering most of the skeleton,
except perhaps for the apex of the skull, which is eroded. Alter-
natively, the mandible might be pushed in the sediment by
a passing elephant that stepped on it. Most of the skull is pene-
trated by roots (Figs. 5 and 8c) that have caused extensive damage,
especially at the less compact and sinuous parts of it. The left tusk
was removed from its alveolus after death and before burial, as the
alveolus was found empty. Some bones exhibit post-depositional
breakage, as the tibia, which was broken by a fault near its distal
end (Fig. 5). No recovered specimen shows signs of severe distor-
tion or compression.
It is quite possible that several of the remaining bones are more
broadly scattered, beyond the excavated area. Actually, several bone
and tusk fragments were recovered from the disturbed sediment
NE of the excavated area, but many others may have been missed,
as the disturbed sediment was widely dispersed. The original
position of these fragments and their taphonomic relation to the
skeletal elements that were excavated in situ are unknown.
The overall pattern of the skeletal elements position implies that
the original orientation of the mammoth corpse before burial was
also NEeSW, with the head to the NE. The area SWof the excavated
quarry may preserve the distal part of the axial skeleton, as well as
some long bones. However, the excavation was not extended to the
0
1 m
Pliocene marls
clayey sand
cross bedd sand ed
sand, coarse sand
palaeosols
pebbles
soil
Fig. 2. Lithostratigraphy of the uvial sediments at Loussik locality. The mammoth
skeleton was found in the clayey sand layer.
Fig. 3. The distribution of the mammoth skeletal elements at Loussik (based on
original sketches by L. Stavropoulou, redesigned and completed with own data). Some
minor fragments are omitted for clarity reasons. Smaller bones that lie under larger
ones are also not shown (e.g. metapodials under the skull). Graphical scale: 1 m.
Fig. 4. Loussik mammoth skeletal elements in situ: 1, skull (rostral part is on the
right); 2, right hemimandible; 3, left hemimandible (still unexcavated; number is
placed on m2); 4, left ulna; 5, left calcaneus. The vertical white line indicates the
direction of the bulldozer cut (EeW). Note the level difference among the skull, the
right and the left hemimandible, indicating rapid sediment deposition and burial.
Graphical scale: 30 cm.
SW mainly for nancial and technical reasons. The observed
NEeSW dispersion does not correspond to a hypothetical palae-
ocurrent ow direction, as the excavated skeletal elements, with
the exception of the tibia, two thoracic vertebrae with long spinal
processes and some costae, are not arranged parallel to this direc-
tion (Fig. 3). Nonetheless, there is also another bone concentration,
in which the elongated specimens (costae, left scapula fragment)
are arranged in an NWeSE direction (Figs. 3 and 6). No sorting
pattern is observed.
4.1. Biological agencies in skeleton disassociation
A possible explanation for the observed dispersal pattern is that
the mammoth bones were scattered because of trampling and/or
carnivore activity. Extant elephants are known to be attracted to,
examine and manipulate bones of their dead conspecics using
their trunk and feet; they usually kick, drag, toss or carry them
around, often to long distances (Haynes, 1991, p. 157e158; Moss,
2000, p. 270e271, photo plates after p. 128). The excavated speci-
mens exhibit, however, no trampling marks and no long bone
fractures that can be attributed to trampling proboscideans.
Carnivore or rodent gnawing marks are also absent. At least a minor
carnivore activity is, though, indicated by the presence of a bilobe,
subspherical coprolite, with one concave and one conical end. Its
morphology and dimensions (44 mm long, with a diameter of
37e43 mm) suggest that it can be referred to a large spotted hyena
(Crocuta crocuta), although its attribution to a larger hyaenid
species as Pachycrocuta brevirostris cannot be excluded (Keiler,
2001; Partt and Larkin in Lewis et al., 2010).
4.2. Possible human activity
A single bone, the left fourth metatarsal, bears a cluster of linear
marks on its plantar surface that exhibit the following characters:
(a) they are subparallel to each other and run perpendicular to the
long axis of the bone; (b) they are sharp with well-dened borders;
(c) they occur in a recessed area, not exposed to abrasion; (d) they
are concentrated in an area suitable for cutting the plantar muscles.
Though these observations are very preliminary, they are consis-
tent with human induced cut marks (Shipman and Rose, 1983;
Fisher, 1984; Olsen and Shipman, 1988; Blumenschine et al.,
1996). However, since no similar marks have been observed on
other skeletal elements, this observation is considered a mere
indication, not evidence of human activity around the mammoth
corpse.
5. Systematics
Order: Proboscidea Illiger, 1811
Family: Elephantidae Gray, 1821
Genus: Mammuthus Burnett, 1830
Mammuthus trogontherii (Pohlig, 1885)
Synonymy:
Elephas antiquus Falconer and Cautley, 1847 (Doukas and
Athanassiou, 2003, p. 100, Table 3)
Elephas antiquus Falconer and Cautley, 1847 (Athanassiou,
2010)
Elephas antiquus Falconer and Cautley, 1847 (Tsoukala et al.,
2010, Table 1, p. 10)
5.1. Material
The Loussik partial skeleton consists of the skull and mandible,
the anterior part of the vertebral column, several costae, both
scapulae, left ulna, right tibia, most carpals, tarsals and meta-
podials, and several phalanges. A detailed list of the recovered
anatomical parts is given in Table 1. Their distribution in the
mammoth skeleton is presented graphically in Fig. 7.
5.2. Description
5.2.1. Skull
The skull is partially preserved, lacking most of the sinuous
upper part, above the level of the zygomatic arches (vertex,
braincase, nasal cavity, upper part of the occipital d Fig. 8). The
preserved parts are in rather bad condition as they are much
fragmented and suffer from recent root weathering. The tusk
alveoli are also damaged, lacking rostral parts, but their general
morphology is recognizable: they are subcircular in cross section
and diverge slightly from each other, as they bend laterally in their
rostral regions, remaining, though, very close together. In their
anteriormost preserved part, the tusks appear to have been only
about 5 cm apart. Only the basal part of the right tusk remains in
situ. In lateral view, the skull appears high and anteroposteriorly
shortened. Both zygomatic arches are present, though badly frac-
tured, forming with the tusk alveoli an angle of about 110
. The
latter are directed downwards, forming with the molar alveolar
plane an angle of about 120
. The maxilla is particularly high to

Fig. 5. Loussik mammoth skeletal elements in situ: 1, the occipital region of the skull
(partly covered with consolidant-impregnated gauze); 2, the right tibia. Note the
presence of roots that run through the skull and the post-depositional fracture and
displacement in tibia. Graphical scale: 30 cm.
Fig. 6. Loussik mammoth skeletal elements in situ: 1, left scapula; 2, right scapula;
note the spine fragment a that was broken off from its original position b during
deposition; 3, an accumulation of costae. Graphical scale: 30 cm.
accommodate the hypsodont molars. In caudal view, only the basal
part of the occipital is preserved, with its elliptical foramen
magnum and the massive condyles. The highly pneumatised
occipital is very much developed laterally and posteriorly (Fig. 8c).
Its central part, dorsally to the foramen magnum is denser and
distinctly recessed in relation to the lateral parts. The available
cranial measurements are given in Table 2. The skull remains
partially unprepared, particularly ventrally.
5.2.1.1. Hyoid apparatus. Among the recovered cranial parts are the
basihyoid and a fragment of the right stylohyoid (Fig. 9), two of the
ve ossicles (one basihyoid, two thyrohyoids, two stylohyoids) that
constitute the hyoid apparatus. Another fragment may belong to
a thyrohyoid. The hyoid apparatus is situated in the gular region
and it is articulated to the cranial base via the tympanohyal carti-
lages (Shoshani and Marchant, 2001; Shoshani et al., 2007). The
basihyoid is a dorsoventrally attened, straight bone that becomes
thicker laterally and bents slightly posterodorsally to articulate
with the left and right thyrohyoids. The Loussik specimen is
80 mmwide; the maximumand minimumdiameters at the middle
are 19.0 and 11.5 mm respectively. The stylohyoid is a Y-shaped
bone consisting of a superior, a posterior and an inferior ramus. It is
very rare as a fossil nd, but it is potentially signicant for taxo-
nomic and phylogenetic studies (Shoshani and Tassy, 2005;
Shoshani et al., 2007). The Loussik specimen has broken rami, so
their relative size cannot be assessed. The identication of the rami
Table 1
List of the recovered anatomical parts that belong to the partial skeleton of
Mammuthus trogontherii from the locality of Loussik. The anatomical
determination of the specimens marked with a question mark is not
absolutely positive, because of incomplete preservation. A graphical
presentation of this list is given in Fig. 7.
Anatomical part Side
Skull with M3s
Tusk fragments dex.?
Second upper molar dex.
Mandible with m2s, m3s
Basihyoid
Stylohyoid dex.
Atlas
Cervical vertebra 3
Cervical vertebra 5
Cervical vertebra 7
Thoracic vertebra 1
Thoracic vertebra 2
Thoracic vertebra 3
Thoracic vertebra 4
Thoracic vertebra 5
Thoracic vertebra 6
Thoracic vertebra 7
Thoracic vertebra 9
Thoracic vertebra 10
Thoracic vertebra part
Caudal vertebra?
More than 15 costae
Scapula sin.
Scapula dex.
Ulna sin.
Scaphoid dex.
Lunar sin.
Lunar dex.
Triquetrum sin.
Triquetrum dex.
Pisiform dex.
Trapezium dex.
Trapezoid sin.
Trapezoid dex.
Magnum dex.
Hamatum sin.
Hamatum dex.
Metacarpal II dex.
Metacarpal III sin.
Metacarpal III dex.
Metacarpal IV sin.
Metacarpal IV dex.
Metacarpal V sin.
Metacarpal V dex.
Phalanx prox. manus IV sin.
Phalanx prox. manus IV dex.
Phalanx media manus III sin.
Phalanx media manus III dex.
Phalanx media manus IV dex.
Ischium part sin.
Ischium part dex.
Femoral greater trochanter epiphysis dex.
Tibia proximal part sin.
Tibia dex.
Calcaneus sin.
Astragalus sin.
Navicular dex.
Cuboid dex.
Metatarsal I?
Metatarsal IV sin.
Metatarsal IV dex.
Phalanx prox. pedis IV sin.
Phalanx media pedis II?
Sesamoid
a
b
Fig. 7. Schematic drawing of a mammoth skeleton, showing the anatomical position of
the excavated skeletal elements at Loussik: a, left side; b, right side. Based on
a drawing of Palaeoloxodon antiquus by C. Beauval (available at www.archeozoo.org),
adapted to the mammoth morphology.
was based on their relative robustness, the superior ramus being
the most robust and the inferior the least. All three rami are
elliptical in cross section, more attened medially. Due to incom-
plete preservation none of the characters described by Shoshani
et al. (2007) can be scored on this specimen. A peculiarity is that
the superior and posterior rami are not aligned but form an open
bent (an angle of 132
between them). A similar morphology is

observed in Mammuthus columbi (Shoshani et al., 2007, Fig. 3), but
not in M. trogontherii from West Runton (Lister and Stuart, 2010,
Fig. 24).
5.2.2. Mandible
The mandible (Fig. 10) is very massive. The symphysis is badly
damaged, particularly at its right side. The base of the symphyseal
process is, though, preserved, indicating that the rostral process
was weak. The mandibular corpus is very robust, especially near
the mandibular angle. Rostrally it is very deep, but it becomes
gradually shallower caudally, as its ventral border curves dorsally,
forming a rounded mandibular angle. The mandibular ramus is
much developed anteroposteriorly. It has a thick caudal border that
ends to the condylar process. Rostrally the ramus is much thinner
and forms the coronoid process, which is connected to the condylar
one by an oblique ridge. The masseteric fossa is wide but rather
shallow. Measurements are given in Table 3.
5.2.3. Upper dentition
The tusk morphology is known fromthe right tusk base, retained
inthe alveolus, andfromseveral small fragments recoveredfromthe
bulldozer debris, whichpresumably belongtothesame right tusk. In
Fig. 8. The skull of the Loussik mammoth: a, anterior view; b, lateral view (right
side); c, caudodorsal view in an early preparation stage (note the presence of two
metapodials under the skull). Graphical scale: 20 cm.
Table 2
Cranial measurements of the Loussik mammoth (in mm).
Maximal length (parallel to the alveolar level) >1200
Maximal width (at the zygomatic arches) (840)
Width of the praemaxillaries (tusk alveoli) (500)
Minimal distance between tusks 30
Width at the level of the molar alveoli 310
Occipital width >870
Occipital condyles width (at their lateral borders) 220
Foramen magnum height 70
Foramen magnum width 91
Fig. 9. Bones of the Loussik mammoth hyoid apparatus: a, right stylohyoid, lateral
view; b, basihyoid, dorsal view. Graphical scale: 20 mm.
cross section the tusk is subcircular, with a maximum diameter of
199 mm and a minimum of 180 mm at its base, in the alveolus. The
largest recoveredtuskfragment is 440mmlong, withmaximumand
minimum cross section diameters of 203 mm and 183 mm respec-
tively. Although it is very short, it exhibits a noticeable curvature.
In the natural (due to breakage) cross sections of two other
small fragments the Schreger pattern can be observed. The
Schreger pattern is a special character of the proboscidean ivory
and has been used in genus-level taxonomy (Espinoza and Mann,
1993; Palombo and Villa, 2001; Trapani and Fisher, 2003). It
consists of two sets of radiating spiral lines that intersect with each
other, forming the Schreger angles, and reects macroscopically the
dentine internal structure. The Schreger lines in both examined
specimens form acute angles of 87e89
near the cement/dentine

junction (Fig. 11).
The skull has both M3s in situ, while open alveoli in both sides
indicate the presence of M2s in life. A much worn molar, found
isolated, may well represent the right M2. It consists of 8 plates and
a distal talon. Its length is 153 mm and its width 95 mm. The M3s
are little worn, so the distal parts of them are still in the alveoli.
Nine plates plus the anterior talon are in use (a tenth plate is hardly
worn), forming broad occlusal surfaces that are 150 mm long and
102 mm wide (Fig. 12). As both third molars are in situ their
maximal dimensions and plate number cannot be measured or
counted; e.g. the width is expected to be larger towards the crown
base. A metrical estimation based on the morphology of the maxilla
is given in Table 4. It is not possible to estimate the molar height,
but the very high maxillas could accommodate third molars as high
as 200 mm. The enamel is rather thick and moderately folded and
wrinkled, particularly at the anterior and median region of the
occlusal surface, without, though, forming loxodont sinuses. The
enamel at the occlusal surface of the distal, slightly worn lamellae is
almost unwrinkled. At the distalmost part of the occlusal surface
the incipient wear produces a transverse series of subcircular
enamel islets; the median islets merge together with advancing
wear, resulting in a transversely elongated islet.
5.2.4. Lower dentition
The two hemimandibles bear both second and third molars
(Figs. 10a and 13). The former have triangular occlusal surfaces and
are totally worn. They consist of three plates that are worn down to
the roots. The left m3 has the anterior talonid and nine plates in
Fig. 10. Left hemimandible of the Loussik mammoth: a, occlusal view; b, lateral view.
Graphical scale: 10 cm.
Table 3
Mandibular measurements of the Loussik mammoth (in mm).
Left Right
Maximal length (parallel to the labial wall of the corpus) 690 >610
Maximal width of the corpus 185 205
Maximal height (at the articulation) 480 (480)
Height at the coronoid process
(parallel to its anterior border)
e (360)
DAP of the ramus, at the level of the coronoid process
(parallel to the labial wall of the ramus)
315 320
Corpus height, at the base of the ramus 250 235
Corpus height, at the level of m2 305 e
Fig. 11. Cross section of a tusk fragment of the Loussik mammoth exhibiting the
Schreger pattern. The Schreger angles are slightly acute. Graphical scale: 10 mm.
Fig. 12. Loussik mammoth upper left M3, occlusal view. Anterior side is on the left.
Graphical scale: 30 mm. The molar is in situ; its non-occlusal part and the maxilla are
not shown, because they are covered by supporting material.
use, while 16 plates are totally visible. It is estimated that another
4e6 plates are still inside the alveolus at the posterior part of the
mandibular corpus, raising the total plate number to 20e22. Its
total length could be greater than 400 mm. Based on the consid-
erable depth of the mandibular corpus the molar may be as high as
200 mm. The occlusal surface is rather broad and oval in shape. The
enamel is fairly thick (thicker than that of the upper molars) and
moderately wrinkled at the medial part of the occlusal surface. The
enamel of the slightly worn distal lamellae (seventh and eighth)
forms a transversally elongated islet and two elliptical ones
lingually and labially of it. The right m3 is similar to the left, but has
10 plates in use and 17 visible. Metrical data are given in Table 4.
5.2.5. Vertebral column and ribs
Only the anterior part of the vertebral column is available. The
vertebrae have the typical elephantid morphology (Bezuidenhout
and Seegers, 1996). The atlas is stoutly built and has much devel-
oped lateral wings (Fig. 14a). Apart from atlas, the cervicals were
identied as third, fth and seventh, mainly because they do not t
to each other when trying to re-articulate them. The seventh
(Fig. 14b) is also morphologically distinct, as it has articular facets
for the rst pair of ribs ventrolaterally in its caudal side and it lacks
foramens in its transverse processes. The articular surfaces of all
available cervical vertebrae are fused to the bodies. Measurements
of the cervical vertebrae are given in Table 5.
The preserved thoracic vertebrae are characterised by their long
spinal processes, particularly the anterior ones. The spinal
processes are strongly inclined backwards, forming with the
vertebral body plane an angle of 50
e60
(the angle increases

caudally), quite unlike certain E. antiquus specimens, as that from
Upnor that has almost vertical spinal processes (Andrews and
Forster Cooper, 1928). (The Upnor vertebral column may be,
though, morphologically aberrant, as in other E. antiquus skeletons
this character is less accentuated de.g. Trevisan, 1954; Maccagno,
1962; Melentis, 1963; Kroll, 1991). The exact anatomical position of
the at least 13 thoracic vertebrae is generally not known, as they
were not found in anatomical association and it is not possible to
Table 4
Upper and lower third molar measurements (in mm) of the Loussik mammoth.
Next to the width is given the plate number at the level of which it was measured.
Plate number excludes any talons.
Left M3 Right M3 Left m3 Right m3
Length (350) (350) >300
a
>290
a
Width 102 e 100 (at 5th) 98 (at 4th)
Plate number >14 >14 >16
b
>17
b
Plate frequency 6.3 6.3 6.1 6.2
Enamel thickness 2.6e2.9 2.6e3.0 2.7e3.2 2.7e3.2
a
Estimated 370e410.
b
Estimated 20e22.
Fig. 13. Loussik mammoth mandibular dentition, occlusal view: a, left m2 and m3; b,
right m2 and m3. Graphical scale: 10 cm.
Fig. 14. Loussik mammoth cervical vertebrae: a, atlas, cranial view; b, seventh
cervical vertebra, caudal view. Graphical scale: 10 cm.
Table 5
Cervical vertebral measurements of the Loussik mammoth (in mm).
Atlas Third Fifth Seventh
Maximal height 213 >240 >290 362
Maximal width 431 305 e 337
Cranial articular width 228 154 144 150
Cranial articular height 112 164 168 (140)
Caudal articular width 195 171 181 155
a
Caudal articular height 101 167 166 159
DAP of the corpus
(measured ventrally, at the sagittal plane)
53 57 87
DAP of dorsal arc 90
a
Excludes the articular surfaces for the rst ribs.
re-articulate them, because most of them have incompletely
preserved bodies or lack one of their body epiphyses (not yet
fused). An exception is the second (Fig. 15a) and third, which were
found associated to each other and were identied mainly on their
very long and robust spinal processes. The best preserved thoracic
vertebrae were positioned relative to each other using the char-
acters that change gradually in craniocaudal direction: raising of
the transverse processes; increased inclination of the spinal
process; raising of the articulations for the costae; changing of the
body shape to more triangular (in cranial or caudal view); changing
of the vertebral foramen shape from triangular to elliptical (in
cranial or caudal view) (Kulczyski, 1955; Bezuidenhout and Seegers,
1996). The more caudally placed thoracic vertebrae (starting from
the ninth) (Fig. 15b) have bodies with unfused anterior articula-
tions. The posterior articulation is fused in all available thoracic
vertebrae. None of the studied thoracic vertebrae exhibits any
marked bilateral asymmetry in the development of the transverse
processes or the articular facets, as it is often found in individuals of
the genus Mammuthus (Lister, 2009; Lister and Stuart, 2010).
Measurements of the thoracic vertebrae are given in Table 6. No
lumbar or sacral vertebrae were found. A much eroded specimen
possibly represents one of the rst caudals.
The costae are fragmentary. The anterior ones are morphologi-
cally distinct, as their head forms an almost right angle with their
body. This angle becomes more obtuse caudally, as the costal head
articulates at a gradually more dorsal area on the vertebral bodies.
Due to their incomplete preservation the costae were not identied
anatomically; their relative position was just characterized as
anterior, middle or posterior. A tentative anatomical position of the
available costal fragments is given in Fig. 7.
5.2.6. Thoracic limb
The thoracic limbs are represented by both scapulae, left ulna,
most of carpal and metacarpal bones and ve phalanges. Their
dimensions are given in the Tables 7e11. The scapulae are incom-
pletely preserved. The left one preserves only the anterior and
central part along the spine, lacking most of the posterior blade part
(fossa infraspinata). The spine is almost straight, presenting a weak
concavity cranially. It preserves both hamate and suprahamate
processes, which form an angle of 35
between them. The right

scapula is more complete, lacking the cranial part of the blade (fossa
supraspinata). Both have fusing proximal epiphyses, the fusion lines
remaining well visible. Their dimensions are quite similar to the
mammoth specimens from Steinheim (Dietrich, 1912) and West
Runton (Lister and Stuart, 2010). The latter has, however, quite
larger glenoid cavity. Compared to the Greven E. antiquus speci-
mens (Tsoukala and Lister, 1998), the Loussik scapulae are pro-
portionally wider anteroposteriorly (though equally high) and have
Fig. 15. Loussik mammoth thoracic vertebrae: a, second, caudal view; b, about
twelfth, caudal view. Graphical scale: 10 cm.
Table 6
Thoracic vertebral measurements (in mm) of the Loussik mammoth. The numbering of the 4th to 12th is mostly tentative, used to indicate the relative position of the
vertebrae, and corresponds to their approximate anatomical position in life. The remaining thoracic vertebrae were fragmentary and not measurable. The articular
measurements exclude the articulations for the costae.
1st 2nd 3rd 4th 5th 7th 9th 11th 12th
Maximal height 595 (535) >580 e >550 e e e 360
Maximal width 344 e (320) (306) 316 e 320 221 243
Cranial articular width 157 e 139 130 e 122 e e e
Cranial articular height 149 e 123 141 e 139 e e e
Caudal articular width 156 e 142 139 e e 142 128 127
Caudal articular height 130 e (125) (141) e (139) 140 137 123
Spinal process length
(from the top of the vertebral foramen to the apex of the process)
380 510 >450 e >440 e e e 315
DAP of the corpus
(measured ventrally, at the sagittal plane)
82 e e 77 (83) 77 (78) (90) (80)
Table 7
Scapula measurements of the Loussik mammoth (in mm).
Left Right
Maximal height (parallel to the spine) 1070 1045
Articular height (parallel to the spine) 1010 990
Length of the dorso-caudal border
(from the dorsal end of the spine to the caudal angle)
e (990)
Length of the caudal border
(from the distal articulation to the caudal angle)
e 855
DAP of the distal end 290 e
DT of the distal end (140) e
DAP of the distal articulation 216 e
DT of the distal articulation (135) 145
distinctly smaller glenoid cavity (articular) dimensions. The same
observation on the glenoid cavity dimensions applies also when
comparing the Loussik scapulae to E. antiquus specimens from
Italy (Trevisan, 1954; Maccagno, 1962). Compared to E. antiquus
from Upnor (Andrews and Forster Cooper, 1928) the Loussik
specimens are smaller in all dimensions.
The ulna (Fig. 16a) is robust with a massive olecranon. In lateral
aspect it is rather straight and not bent as in Loxodonta (as guredby
Smuts and Bezuidenhout, 1993, Figs. 12 and 13). The diaphysis is
generally triangular in cross section, though it gradually becomes
almost square-shaped towards the distal extremity of the bone. The
distal epiphysis, while rmly attached to the shaft, is not completely
fused, exhibiting a deep cleft caudally, along the fusing line. The
Loussik ulna is metrically smaller and slenderer than most speci-
mens attributed to male M. trogontherii (Steinheim, Untermafeld,
Table 8
Ulna measurements of the Loussik mammoth (in mm).
Maximal height 975
Articular height (from the medial coronoid process
to the distal articulation)
825
DAP of the proximal end 300
Maximal DT of the proximal end (articular level) 243
DT of the proximal articulation 235
DAP of the olecranon 230
Minimum DAP of the diaphysis 127
Minimum DT of the diaphysis 110
DAP of the distal end 186
DT of the distal end 213
DAP of the distal articulation 102
DT of the distal articulation 152
Table 9
Carpal measurements of the Loussik mammoth (in mm).
Scaphoid Right
Maximal height 142
Maximal DAP 119
Lunar Left Right
Maximal height 83 85
Maximal DAP 145 141
DAP of the proximal articulation e 114
DT of the proximal articulation e 135
DAP of the distal articulation 126 121
Triquetrum Left Right
Maximal DAP e 138
Maximal DT e 146
DAP of the proximal articulation 100 e
DT of the distal articulation e 142
Pisiform Right
Maximal DAP 144
Maximal height 57
Minimal DT (at the middle of the bone) 75
Trapezium right
Maximal height 97
Maximal DT 95
Trapezoid Left Right
Maximal DAP 126 126
Maximal DT 88 85
DAP of the proximal articulation 101 100
DT of the proximal articulation 66 66
DT of the distal articulation 71 63
Magnum Right
Maximal height 123
Maximal DAP 155
DAP of the proximal articulation >120
DAP of the distal articulation e
Hamatum Left Right
Maximal height 123 (125)
DT of the proximal articulation 114 (119)
DAP of the distal articulation 136 e
DT of the distal articulation 122 e
Table 10
Metacarpal measurements of the Loussik mammoth (in mm).
Metacarpal II Right
Maximal height 192
Minimal DT of the shaft 73
Metacarpal III Left Right
DAP of the proximal end 126 e
DT of the proximal end 82 84
Minimal DT of the shaft 72 71
DAP of the distal end 95 93
DT of the distal end 95 99
Metacarpal IV Left Right
DAP of the proximal end 117 119
DT of the proximal end e 99
Minimal DT of the shaft 82 82
Metacarpal V Left Right
DAP of the proximal articulation e 95
Table 11
Phalanges manus measurements of the Loussik mammoth (in mm).
Phalanx prox. IV Left Right
Height 93 100
DT of the proximal end 94 (97)
DT of the proximal articulation 78 (82)
Phalanx media III Right
Height 61
DAP of the proximal end 47
DT of the proximal end 70
DAP of the proximal articulation 43
Phalanx media IV Right
Height 43
Maximal DAP 38
Maximal DT 50
Odessa, West Runton, Kostolac d Dietrich, 1912; Dubrovo, 2001;
Lister and Stuart, 2010) and similar or smaller than specimens
attributed to E. antiquus (Andrews and Forster Cooper, 1928;
Trevisan, 1954; Maccagno, 1962; Melentis, 1963).
The right carpus is complete, represented by all eight bones; the
left lacks the scaphoid, the pisiform, the trapeziumand the magnum.
The bones of the right carpus can be re-articulated very well to each
other, despite damage at the anterior part of the magnum, revealing
Fig. 16. Elements of the Loussik mammoth thoracic limb: a, left ulna, lateral view; b, right pisiform, medial view; c, right scaphoid, lateral view; d, right triquetrum, proximal view;
e, right triquetrum, distal view; f, right lunar, proximal view; g, right lunar, distal view; h, right trapezium, lateral view; i, right trapezium, medial view; j, right trapezoid, proximal
view; k, right trapezoid, distal view; l, left hamatum, proximal view; m, left hamatum, distal view; n, right magnum, proximal view; o, right magnum, distal view. In proximal and
distal views the anterior side is facing towards the bottom of the gure. Graphical scales: 20 cm (ulna), 5 cm (carpals).
an aserial carpus structure (the lunar extends well over both
magnumandtrapezoid). The scaphoid(Fig. 16c) is triangular inshape
andlaterally attened. It is stouter thanthat of Loxodonta (Smuts and
Bezuidenhout, 1993) and it is very close to the morphology of
M. primigenius, as depicted by Andrews and Forster Cooper (1928, pl.
III). Its radial facet is high and ends proximally to a pointed top. The
distal facet (for the trapezoid) extends proximolaterally till about the
middle of the bone. The triangular-shaped lunar (Fig. 16f and g) is
wider than the magnum(situatedjust belowit), also overlapping the
trapezoid. This disposition has been considered characteristic for the
mammoths (e.g. Trevisan, 1954, Figs. 22and47), thoughlater authors
oftenattribute it toindividual variation(e.g. BedenandGurin, 1975).
The triquetrum(Fig. 16d and e) is also of triangular shape with a long
apophysis that projects lateropalmarly. Both, left and right, are
damaged, particularly the former, which is broken palmarly. The
pisiform(Fig. 16b) is the lateralmost bone of the proximal carpal row.
It is elongated, projecting palmarly. The trapezium (Fig. 16h and i),
situated medially in the distal carpal row, is mediolaterally attened
and pentagonal in medial aspect, due to a tuberosity at its palmar
side. The trapezoid (Fig. 16j and k) has two main triangular articular
surfaces proximally and distally. The distal articular surface extends
laterocaudally to form a facet for the magnum, unlike certain
E. antiquus specimens (Andrews and Forster Cooper, 1928, pl. IV;
Trevisan, 1954, Fig. 21). Cranially there is only one facet for the
magnum, which corresponds to a similar unied facet in the
magnum. This is considered as a character of Mammuthus, not
observedinE. antiquus (Dubrovo, 2001). Themagnum(Fig. 16nando)
is a massive, cuboid bone with a prominent palmar tuberosity. Its
proximal articular facet has almost parallel lateral and medial
borders. The distal facet for the second metacarpal is well developed,
extending to the whole length of the distal articulation. This facet is
usually reduced dorsally in E. antiquus (Beden and Gurin, 1975),
thoughthe presence of non-reducedfacets is alsoreported(Trevisan,
1954, p. 36e37; Maccagno, 1962, p. 112). The hamatum (Fig. 16l and
m) bears two large articular surfaces for the triquetrumand the fth
metacarpal that meet eachother alongthelateral marginof the bone.
The contact of these surfaces is very short in E. antiquus (Beden and
Gurin, 1975).
The metacarpals (Fig. 17) are elongated, robust, dorsopalmarly
attened bones, with wide proximal articular facets for the distal
carpal row and a distal trochlea for the articulation of the proximal
phalanges. The proximal articulation of the third metacarpal is
noticeable, because it is very prominent, the facets for the magnum
andthe hamatumbeingstrongly inclinedandforminganacute angle
between them. Metrically the Loussik metacarpals are smaller and
rather less stout than certain E. antiquus ones (Andrews and Forster
Cooper, 1928; Trevisan, 1954, Figs. 25 and 26; Tsoukala and Lister,
1998). Compared to the metacarpals of M. trogontherii from Stein-
heimtheyare alsosmaller, particularly the fthmetacarpal (Dietrich,
1912).
5.2.7. Pelvic limb
The pelvis fragment comes from the symphyseal area of the
ischial bones at the caudal part of the pelvis. The symphysis is
fused, but the fusion line is clearly demarcated. The ischium height
at its caudal area (at the ischial tuberosity) is about 33 cm. No other
measurement can be taken on this specimen. The only femoral
remain is the epiphysis of the greater trochanter of the right femur.
It has no trace of any incipient fusion with the femoral body. The
right tibia (Fig. 18a) is well preserved, though it is somewhat
deformed at its proximal end. It is robust and fully grown, both
epiphyses being already fused. Of the left tibia only the proximal
end is available. Measurements are given in Table 12.
The calcaneus (Fig. 18b) is heavily built with very rugose non-
articular surfaces, particularly at the tuber. The facet for the
bula, as well as the lateral part of the ectal facet, are missing. The
sustenactular facet is markedly triangular. The general morphology
of the bone is very similar to that of M. primigenius calcaneus as
depicted by Andrews and Forster Cooper (1928, pl. VII). Metrically
(Table 13) it is smaller than the M. trogontherii specimens from
Steinheim and Untermafeld (Dietrich, 1912; Dubrovo, 2001), as
well as most of those attributed to E. antiquus (Andrews and Forster
Cooper, 1928; Trevisan, 1954, Figs. 25 and 26; Tsoukala and Lister,
1998), except for Riano (Maccagno, 1962, p. 116).
The astragalus (Fig. 18e and f) is low and broad, attened in
a dorsoplantar direction. The morphology and the disposition of its
articular surfaces are verysimilar tothoseof M. primigenius according
to Andrews and Forster Cooper (1928, pl. VI) and to the West Runton
specimen (Lister and Stuart, 2010, Fig. 7). Metrically (Table 13) the
astragalus from Loussik is smaller than the specimens from Stein-
heim, Untermafeld and West Runton (Dietrich, 1912; Dubrovo,
2001; Lister and Stuart, 2010) and larger that the specimens from
La Fage, attributed to M. aff. trogontherii (Beden and Gurin, 1975).
The other two recovered tarsal bones, navicular and cuboid,
belong to the right foot. The navicular (Fig. 18c and d) is at with
a concave proximal and a convex distal articular surface, with four
Fig. 17. Loussik mammoth right metacarpal IIeV series, dorsal view. The MC II is on the right. The gured MC V is the left one (pictured mirrored). Graphical scale: 5 cm.
facets; the facet for the cuboid is the largest. At its caudal surface
the facet for the calcaneus is small. The posteromedial part of the
bone has eroded surfaces. It is smaller than the navicular from
Untermafeld(Dubrovo, 2001) andvery similar insize to the smaller
navicular fromStreinheim(Dietrich, 1912). The cuboid(Fig. 18handi)
is triangular in shape. Unlike the cuboid from Upnor (E. antiquus d
Andrews and Forster Cooper, 1928, pl. VIII) it does not have a prom-
inent tubercle in its caudal surface. Measurements are given in
Table 13.
The fourth metatarsal (Fig. 18g) is a short and robust bone with
a very faintly divided triangular proximal articulation for the
ectocuneiform and the cuboid. The corpus is trapezoid in cross
section, wider dorsally. It is shorter than the metatarsal IV from
Steinheim (Dietrich, 1912) and metrically similar to the same bone
of E. antiquus from Riano (Maccagno, 1962). Measurements are
given in Table 14.
The available phalanges manus and pedis are not differ-
entiated morphologically from the typical of the family
Elephantidae. Their measurements are given in Tables 11 and 15
respectively.
Fig. 18. Elements of the Loussik mammoth pelvic limb: a, right tibia, dorsal (anterior) view; b, left calcaneus, cranial view; c, right navicular, proximal view; d, right navicular, distal
view; e, left astragalus, proximal view; f, left astragalus, distal view; g, right metatarsal IV, dorsal view; h, right cuboid, proximal view; i, right cuboid, distal view. Graphical scales:
20 cm (tibia), 5 cm (tarsals and metatarsal).
Table 12
Tibia measurements of the Loussik mammoth (in mm).
Left Right
Maximal height e 750
Maximal DT of the proximal end 257 264
Minimum DT of the diaphysis e 129
DAP of the distal end e 160
DT of the distal end e >190
DAP of the distal articulation e 126
DT of the distal articulation e 172
Table 13
Tarsal measurements of the Loussik mammoth (in mm).
Calcaneus Left
Maximal height (normal to the cuboid articulation) 222
Maximal diameter of the head (tuber calcaneus) 143
Maximal DAP (normal to the astragalus articulations) 141
Total DT of the articular surfaces for the astragalus 197
Maximal diameter of the lateral articular surface for the astragalus (112)
Maximal diameter of the medial articular surface for the astragalus 102
Astragalus Left
Maximal height (parallel to the calcaneus articulations) (150)
Maximal DAP (normal to the calcaneus articulations) 90
Maximal DT (parallel to the calcaneus articulations) 177
Navicular Right
Maximal DAP (100)
Maximal DT 153
DT of the distal articulation e
Cuboid Right
Height 58
Maximal DAP 123
6. Inferred physical and ontogenetic characters
6.1. Stature
A simple method for estimating the shoulder height in life of the
Loussik mammothwouldbe toaddtogether the articular heights of
the thoracic limb bones, and then correct for the non-preserved soft
tissues and cartilage (e.g. Osborn, 1942, p. 1022). This is because the
longbones of most Proboscidea (withthe exceptionof archaic small-
sized forms) are anatomically placed one under another, due to the
columnar stance of the limb. However, a humerus has not been
found among the excavated bones, so this method cannot be fol-
lowed here. A less accurate estimation can, though, be based in
comparisons of the available limb bone (scapula, ulna and tibia)
dimensions with those of other mammoth skeletons of known
shoulder height. The tibia is rather problematic for stature estima-
tion, as it stops growing early in ontogeny, not reecting the
subsequent body growth. The distal epiphysis of ulna remains
unfused until late in life (Roth, 1984; Lister, 1999), constituting
a better predictor. Christiansen (2004) gives correlated ulna
dimensions with shoulder height for the M. meridionalis skeleton
fromDurfort, mounted at the MusumNational dHistoire Naturelle
in Paris. This skeleton is 383 cm high and its ulna has a maximal
height of 1083 mm. Similarly, the M. trogontherii skeleton from
Steinheim is 370 cm high and its ulna maximal height is 1075 mm
(Dietrich, 1912). The Loussik ulna is 10%and9%smaller respectively
and d hypothesizing an isometric limb bone scaling d we get
equally shorter statures of 345 and 336 cm respectively. Nonethe-
less, this simplistic approachmayunderestimate the actual height at
withers, as the Loussik mammoth has relatively large scapulas and
not fully fused ulna. Comparing the combined left scapula and ulna
articular length with corresponding measurements on other
M. trogontherii skeletons taken from Lister and Stuart (2010, Suppl.
Table B) a higher shoulder height is predicted. The available data are
given in Table 16 and presented graphically in Fig. 19. The trend line
illustrated in the graph predicts a shoulder height of 363 cm for the
Loussik skeleton. If the female individuals are not taken into
account, then the predicted Loussik skeleton height increases by
5e6 cm. It should be noted however, that Lister and Stuart (2010)
consider scapula and ulna as problematic bones for skeletal height
estimation (mainly because of inconsistent measurement methods
among authors) and, consequently, the estimated stature of 363 cm
should be considered as highly approximate. To estimate the live
shoulder height fromskeletal height another 6.3% (Osborn, 1942) or
15 cm (Christiansen, 2004) has to be added, resulting in 386 or
378 cm respectively.
6.2. Body mass
The body mass is typically estimated using humerus or femur
dimensions (maximum height, diaphyseal circumference), or
shoulder height. In the absence of humerus or femur at Loussik the
estimated shoulder height has to be used. This is already prob-
lematic at the beginning, as the estimated skeletal height is very
approximate, and because extant elephants are smaller and
anatomically different frommammoths, as well as from each other.
Moreover, two elephants of identical shoulder height can differ in
body mass by a factor of 2 (Roth, 1990) rendering any estimation
even less accurate.
Two approaches were used for the estimation of the Loussik
individual body mass from the shoulder height. The rst is a linear
regression based on a data sample of male E. maximus (Christiansen,
Table 14
Metatarsal measurements of the Loussik mammoth (in mm).
Metatarsal IV Left Right
DAP of the proximal end 86 e
DAP of the proximal articulation 82 e
Table 15
Phalanges pedis measurements of the Loussik mammoth (in mm).
Phalanx prox. IV Left
Height 96
Table 16
Correlation between combined scapula and ulna articular height (in cm) and the
measured or estimated skeletal shoulder height (in cm) of Mammuthus skeletons.
Data taken from Lister and Stuart (2010, Suppl. Table B).
Skeleton Species Gender Scapula ulna
articular length
Skeletal height
Condover M. primigenius _ 155 296
Praz Rodet M. primigenius _ 151 280
Olyosh M. primigenius \ 120 215
Loussik M. trogontherii _ 184 363
a
West Runton M. trogontherii _ 190 369
Edersleben M. cf. trogontherii \ 170 345
Nogaisk M. meridionalis _ 220 410
Georgievsk M. cf. meridionalis _ 193 396
a
Estimated from the graphical presentation of the data (Fig. 19).
190
220
250
280
310
340
370
400
430
100 120 140 160 180 200 220 240
s
h
o
u
l
d
e
r
h
e
i
g
h
t
(
c
m
)
scapula + ulna height (cm)
M. primigenius
M. trogontherii
M. meridionalis
Fig. 19. Graphical presentation of the correlation between combined scapula and ulna
articular height and the measured or estimated skeletal shoulder height of Mammu-
thus skeletons (see also Table 16). The two female skeletons are marked as such.
Comparative data (open circles) taken from Lister and Stuart (2010, Suppl. Table B). The
estimated position of the Loussik skeleton is marked with a black circle.
2004, Table 2) and gured in Fig. 20. Using a live shoulder height of
380 cm, a body mass of 7850 kg is estimated for the Loussik
mammoth. The second approach estimates the body mass using the
equation BM5.07 10
4
SH
2.803
, where BMis the body mass in
kgandSHthe shoulder height incm(Christiansen, 2004, p. 527). The
equation is based on a sample of male L. africana from Uganda. For
a shoulder height of 380 cm the equation yields a body mass of
8633 kg. In conclusion, based on the available metrical data and
having in mind the rough approximation of the results, a body mass
of about 8 t is estimated for the Loussik mammoth.
6.3. Gender determination
The extant elephants exhibit evident sexual dimorphism in
several characters, the most obvious of which is that the males are
considerably larger than females. In fact elephants count among the
most sexually dimorphic mammals: the males are about 20e40%
larger in linear dimensions and can be almost twice as heavy as the
females (Haynes, 1991; Sukumar, 2003). Moreover, the tusks of
female individuals are much shorter, thinner and less curved. Other
sexual dimorphic characters include the shape of pelvis and
mandible (relativewidth, relative corpus height, development of the
symphyseal process), as well as the more rapid and continuous body
size growth of males till late age. Similar dimorphism patterns have
also been observed in fossil proboscideans, as mammoths
(M. primigenius, M. columbi) andmastodonts (Mammut americanum)
(Haynes, 1991; Averianov, 1996; Lister, 1996b). Inference about the
Loussik mammoth gender can be drawn from skeletal size,
symphyseal process development, presence of unfused epiphyses
and tusk size (as a reasonably complete pelvis is not available).
The estimatedskeletal shoulder height of the Loussik individual
is placed in the range of male skeletons of M. trogontherii (Lister and
Stuart, 2010, Table 1), being 10 cm higher than the Azov II skeleton,
which is referred to a very large female. The limb bone dimensions
are generally smaller than corresponding male M. trogontherii
dimensions published in the literature (see Sections 5.2.6 and 5.2.7).
This would allow the attribution of this skeleton to a rather small
male. However, an assignment to an exceptionally large female,
though improbable, cannot be literally rejected; no clear-cut
boundary can be drawn between male and female dimensional
ranges, particularly when temporal and/or geographical variation is
possible among compared fossil samples.
The inferred small symphyseal process of the Loussik mandible
has been considered as a female character, nonetheless not unam-
biguously, as there are several recorded exceptions: Averianov
(1996, Table 26.1) reports female woolly mammoth specimens
with developed process, as well as male specimens with small
symphyseal process. Thus this character cannot be conclusive.
Another sexually dimorphic character is the presence of not fully
fused bones in male individuals of advanced age, as they continue to
growas adults (Roth, 1984; Haynes, 1991). The occurrence of fusing
or unfused limb bone epiphyses in a mammoth in its forties (see
Section6.4), as the Loussik individual, is a clear indicationof a male.
The maximumbase diameter of the preserved tusk fragments at
Loussik (199e203 mm) is placed among the high values reported
for Mammuthus samples. A large sample of M. primigenius tusks
gave maximum base diameters well smaller than 200 mm for
males (a value comparable to that of Loussik) and 90 mm for
females (less than half the diameter of the Loussik tusk)
(Averianov, 1996, based on Vereshchagin and Tikhonov, 1986). The
available published measurements on M. trogontherii specimens
are rather few. The tusk of the West Runton mammoth, which is
assigned to a male individual, has a base diameter of 217 mm(Lister
and Stuart, 2010, Table 2), slightly larger than Loussik. Other male
M. trogontherii specimens measure 210/175 mm (max/min base
diameter) (Steinheim skeleton, Dietrich, 1912) and 215 mm (Azov I
skeleton, Baigusheva and Garutt, 1987, cited by Lister and Stuart,
2010). Specimens assigned to female M. trogontherii from Edersle-
ben and Novogeorgievsk, have base diameters of 138 and 149 mm
respectively (Lister and Stuart, 2010). The maximum base diameter
of specimens attributed to M. meridionalis can be larger (e.g.
238 mm in LAquila, Maccagno, 1962, Fig. 17) 230e250 mm in
Chilhac (Boeuf, 1983, p. 179). The large tusk diameter of the Lous-
sik individual is a clear indication of a male animal, as no female
would possess such a robust tusk. This character, together with the
evidence of continuing growth mentioned above, are considered
here as the most reliable of the available anatomical evidence for
gender determination and thereby the Loussik mammoth is
referred to a male.
6.4. Ontogenetic age
The ontogenetic age of living and fossil elephantid individuals is
commonly estimated by identifying the cheek teeth in use and
observing their wear state. The method is based on the almost
horizontal eruption and progression of upper and lower molari-
form teeth, which gradually bring the more distal molar lamellae
and molars at the occlusal area. This peculiar type of molar eruption
is characteristic of the Elephantidae. Laws (1966) established an
ontogenetic scheme of thirty age groups (IeXXX) for Loxodonta,
dened by mandibular dental progression stages, and assigned
a corresponding age in years for each group. Though widely used,
Laws method has received some criticism, as the resulting ages less
than 30 years have been considered as overestimated (e.g. Lark,
1984; Jachmann, 1985). An amendment of the method for the age
groups of 10e30 years was published by Jachmann (1988). Despite
the possibility of introduced errors (which can simply result from
variations in diet among studied populations that may produce
slightly different occlusal wear patterns in individuals of the same
age) Laws method remains a useful tool in estimating the onto-
genetic age of an African elephant.
4000
4500
5000
5500
6000
6500
7000
7500
8000
8500
260 280 300 320 340 360 380 400
b
o
d
y

m
a
s
s
(
k
g
)
shoulder height (cm)
Fig. 20. Linear regression of male Elephas maximus shoulder height to body mass,
based on Christiansen (2004, Table 2). The estimated position of the Loussik skeleton
is marked with a black circle.
The Laws method has been applied in fossil elephantid species,
as they also exhibit the same horizontal molar progression. As fossil
elephantids do not necessarily have the same plate number in each
molar as Loxodonta, a proper adjustment has to be done where the
method refers to the number of plates in use. A comparison of the
occlusal surface of the Loussik mandible (Figs. 10a and 13) to Laws
(1966) gures leads to the conclusion that the studied individual is
placed to the age group XXII. In this group the m2 has only 2e3
remaining enamel lamellae or it is shed, leaving an open alveolus,
while the m3 has 6e7 lamellae in use. Given a mean African
elephant m3 plate number of 13 (Maglio, 1973, Table 11) and an
estimated plate number of 20e22 for the Loussik m3s, the 9e10
lamellae in use in the Loussik mandible are equivalent to six
lamellae in Loxodonta. The almost totally worn m2s of the Loussik
mandible are also consistent with the m2 state in this age group.
Laws (1966) assigns an age of 39 2 years to group XXII.
Assigning this age to the Loussik individual would, however, be an
underestimation of its real age at death, as a species longevity is
known to scale positively to body size (Eisenberg, 1990; Maiorana,
1990). M. trogontherii was a much larger animal than L. africana, so
it is expected to have a longer lifespan. A 40 years old male African
elephant weights about 5 t (Laws, 1966, Fig. 14), while the body
mass of the Loussik individual is estimated to 8 t (see Section 6.2).
Using the formula given by Maiorana (1990) (longevity scales as
a 0.25 power of body mass) an age of death of about 45 years can be
estimated for the Loussik mammoth.
Studying Asian elephant samples in a similar way to Laws
(1966), Roth and Shoshani (1988, Table 2) cite an age of 37 years
for a specimen, which has more than half of m2 in wear as well as
a slightly worn m3. This information that refers to another extant
species (E. maximus) corroborates the estimated age of the some-
what older Loussik individual.
An estimation of the individual ontogenetic age can also be
based on the epiphyseal synostosis state. This is possible because
the ontogenetic growth of elephants does not cease when the adult
age is reached d as it is common among mammals d but they
continue to grow as adults (Roth, 1984; Haynes, 1991). As the bone
growth principally takes place between the body and the epiph-
yses, several epiphyseal sutures remain open until late in life. The
fusion sequence, as well as the ontogenetic stage during which each
fusion occurs, are generally consistent among individuals and
among extant and fossil species (though minor differences do
occur) (Roth, 1984; Haynes, 1991; Lister, 1999). However, they differ
considerably between the genders, as the males have much more
prolonged growth period in order to acquire their much greater
body size. For any given age class based on dental eruption
sequence, as those of Laws (1966) mentioned above, females
generally have more already fused epiphyses than males.
The Loussik partial skeleton does comprise limb bones, the
epiphyseal synostosis state of which can contribute to the onto-
genetic age estimation. These are the scapulae, the ulna, the
trochanteric epiphysis of the femur and the tibia. The latter is the
only fully grown, with fused epiphyses, the former two have fusing
epiphyses, while the trochanteric epiphysis is completely detached
(found isolated). Studies in extant elephant samples (Roth, 1984;
Haynes, 1991, Appendix) have documented the fusing sequences
of the limb bone epiphyses: Both tibial ends stop growing in Lox-
odonta males during the age classes XVIIIeXX and distal ulna fuses
during or after XXII. The ossied scapular cartilage also fuses late in
life (after class XXI) in Loxodonta (Roth, 1984; Smuts and
Bezuidenhout, 1993) and during or after the XXIIeXXIII classes in
M. primigenius males (Lister, 1999). The trochanteric epiphysis fuses
during the classes XXIIeXXV in extant female Elephas, during XXI in
a single male Loxodonta specimen and during XXIIIeXXV in
M. primigenius (Roth, 1984; Lister, 1999).
The vertebral body epiphyses, several of which remain unfused
in the Loussik skeleton, do not completely fuse to the bodies until
very late in life (sixth decade in both males and females) in extant
African elephant populations (Haynes, 1991). This allows the
vertebrae to continue to grow anteroposteriorly, even when the
individuals have stopped growing in terms of shoulder height.
In conclusion, the synostosis state observed in the Loussik
skeleton is concordant with the fusing sequences of E. maximus, L.
africana and M. primigenius and also corroborates the ontogenetic
classication in the age group XXII, as suggested by the dental
progression state.
7. Discussion
Elephants of the mammoth lineage appeared in Eurasia during
the Late Pliocene, about 3.5 million years ago. The taxonomy of the
early forms is still ambiguous, as they are only known from scanty
and incomplete samples, but they are recently grouped under the
specic name Mammuthus rumanus (S tef anescu, 1924) (Markov and
Spassov, 2003; Lister et al., 2005). The Pleistocene mammoths d
M. meridionalis, M. trogontherii and M. primigenius dconstitute an
evolutionary line exhibiting anatomical trends as increasing molar
hypsodonty and lamellar frequency, enamel thinning, cranial
anteroposterior shortening and heightening and changes in body
size. The main anatomical characters of the mammoths include
a high, domed skull, narrowpraemaxillary bones, twisted tusks and
wide molars with weak enamel folding (Osborn, 1942; Maglio,
1973; Lister, 1996a). Another Pleistocene elephant is the straight-
tusked E. (Palaeoloxodon) antiquus, which coexisted in Europe
with M. trogontherii and M. primigenius. E. antiquus is distinguished
from the mammoths by the presence of a two-domed frontopar-
ietal crest in its skull, the weakly curved, untwisted tusks, the
presence of median loxodont enamel sinuses in the molars, and the
rather thicker molar enamel. In the case of isolated molars,
however, the distinction between E. antiquus and M. trogontherii
can be difcult, due to their similar hypsodonty and lamellar
frequency. This difculty is accentuated by the great morphological
and metrical variation that characterises the elephants in general,
a fact that led Soergel (1913) to establish varieties of intermediate
morphology in these two species: Elephas trogontherii var. anti-
quus and Elephas antiquus var. trogontherii.
The postcranial elements are difcult or impossible to distin-
guish, as they are similar in size and morphology. Several authors
have tried to nd distinguishing criteria based on the shape and the
relative development of the autopodial (mainly astragalus and
calcaneus) facets (e.g. Andrews and Forster Cooper, 1928; Neuville,
1946; Trevisan, 1954; Melentis, 1963; Beden and Gurin, 1975).
Using these criteria, particularly those reported by Andrews and
Forster Cooper (1928), the Loussik postcranial elements are
generally similar to those of Mammuthus (but not without excep-
tions dsee Sections 5.2.6 and 5.2.7). However, there is accumulating
evidence against the validity of such anatomical distinctions: Beden
and Gurin (1975) have observed that the facet morphology changes
during ontogenetic development of individuals that belong to
a single species, so they attribute these morphological differences to
intraspecic variation. Lister and Stuart (2010) also found that no
published character can by reliably used to separate Mammuthus
from Elephas (Palaeoloxodon). Moreover, a recent re-examination of
the dental material from Megalpolis (unpublished data), a fossil
fauna where the criteria of Melentis were based on, as well as
geochemical data (Iliopoulos et al., 2010), have shown that only one
species, E. antiquus, exists in the sample studied by Melentis.
The proboscidean skeleton excavated at Loussik was prelimi-
narily attributed to E. antiquus (Doukas and Athanassiou, 2003;
Athanassiou, 2010), so far the only known MiddleeLate Pleistocene
elephant species in Southern Greece, because of the following
reasons:
(a) The skull vault is damaged (eroded) dorsally in a way that
reveals a horizontal section of the caudally expanded occipital
bone (Fig. 8). During the excavation it was thought that this
section is positioned much higher in the skull and the caudally
(at both sides of the external occipital crest) expanded occipital
bone was erroneously interpreted as the two-bulged fronto-
parietal crest of E. antiquus.
(b) The lower m3 morphology is characterised by high hypsodonty,
fairly folded, rather thick enamel and medium lamellar
frequency, all seen in E. antiquus. Also, the incipiently worn
lamellaeformawidemedianenamel loopandsmaller labial and
lingual ones, as it is usually the case in this species. The
appearance of the occlusal surface is broad, but very close to the
maximumwidth cited for E. antiquus (Maglio, 1973, Table 18).
During the recent preparation of the skull, it was observed that
the whole part above the nasal cavity was destroyed before nal
burial (a usual damage in fossil elephant skulls), so a frontoparietal
crest or vertex is not preserved. Moreover, the tusk sheaths were
found to run almost parallel to each other and are situated very
close together. This morphology, in spite of the absence of the
vertex and tusks, points towards the genus Mammuthus and
excludes and attribution to the straight-tusked elephant, which has
markedly diverging tusk alveoli (Osborn, 1942; Maglio, 1973).
Subsequent observations on two tusk fragments, exhibiting the
Schreger pattern came to corroborate the unexpected cranial
evidence. The Schreger pattern has been used in the proboscidean
genus-level taxonomy, as well as for the forensic discrimination of
ivories that come from fossil or extant elephants. The acute angles
measured perpendicular to the tusk axis and close to the cement/
dentine junction (Fig. 11) are consistent with an attribution to
Mammuthus, in contrast to Elephas (Palaeoloxodon), which exhibits
obtuse outer Schreger angles (over 93
), similar to the extant

genera (Palombo and Villa, 2001).
The attribution to Mammuthus is also compatible with the molar
morphology, as described in Sections 5.2.3 and 5.2.4, particularly the
wide crown, as well as the absence of loxodont sinuses in the enamel
loops. The molars, mainly the third ones, have been used extensively
in mammoth species-level taxonomy, as their morphology directly
reects the species dietary adaptations. Unfortunately, the preser-
vation in situ of the third molars at Loussik does not allow certain
parameters to be measured or counted accurately. The high lamellae
number (about 20e22) and their high hypsodonty are advanced
characters that preclude the attribution of the skeleton to a less
derived species as M. meridionalis. The latter has a maximum plate
number of 14, and a hypsodonty index less than 165 (Maglio, 1973,
Table 30), or even lower (99e146) in specimens from Valdarno, its
type locality (Lister and Stuart, 2010, p. 190). Late forms of
M. meridionalis ddatedat thelatest EarlyPleistoceneor thetransition
to the Middle Pleistocene and often placed in the separate subspecies
Archidiskodon meridionalis tamanensis Dubrovo, 1963 d show
advanced dental characters (higher hypsodonty, higher lamellar
frequency, thinner enamel d Dubrovo, 1977; Baigusheva and Titov,
2010), but again do not reach the high plate number and level of
hypsodonty seen at Loussik.
On the other hand, the Loussik dentition is not so progressive
as in the most advanced species of the Mammuthus lineage,
M. primigenius. The lamellar frequency of 6.1e6.3 plates per 10 cm
of molar length (which may be better attributed to the large molar
size, not to low plate number) is outside the variation of
M. primigenius (more than 6.5 in Maglio, 1973, Table 32; more than
7.4 in Lister, 1996a, Fig. 19.4), a species well known for its tightly
packed enamel plates. A related metrical parameter is the enamel
thickness, which is much thinner (less than 2.0 mm according to
Maglio, 1973 and Lister and Stuart, 2010, p. 191) in M. primigenius,
but as high as 3.2 mm in Loussik m3s. Another distinguishing
feature between the Loussik mammoth and M. primigenius is the
larger size of the former. Although there are large-sized woolly
mammoths, their maximal dental size (upper and lower molar
length less than 285 and 320 mm respectively; Maglio, 1973,
Table 32) is clearly smaller than the estimated for Loussik.
Comparing the Loussik dental specimens to the sample from
Senborn, Germany, the type locality of M. trogontherii, it is
observedthat theyfall well intothe range of variationof this species.
Guenther (1969) counts 14e21.5plates inbothM3s andm3s, placing
the Loussik molars among the maximal values of Senborn. The
same is true for crown height (maximal values 209 mmand 172 mm
for M3 and m3 respectively), though these parameters are roughly
estimated on the Loussik molars (especially the lowers). The same
author reports M3 length and width ranges of 235e375 and 75e120
respectively, and m3 length and width ranges of 260e390 and
75e115 respectively. Again the studied specimens are closer to the
maximum values. Maglio (1973, Table 31) reports similar ranges
(though with lower maxima, especially for the parameter of length)
and also gives a lamellar frequency range of 5.0e8.2 and 5.0e7.2 for
M3 and m3 respectively, with Loussik molars placed very close to
the mean values. Other important dental samples of typical
M. trogontherii are those from Mosbach, Germany, and Tiraspol,
Moldova; their metrical characters do not differentiate essentially
from Senborn (Guenther, 1969; Dubrovo, 1975; Table 2). In
comparison to the recently studied West Runton mammoth (Lister
and Stuart, 2010), the Loussik dental material is slightly dimen-
sionally smaller (as it is also the case with the postcranial skeleton)
and has similar plate number and frequency.
The late Middle Pleistocene mammoths are often more advanced
in dental characters (higher plate number, higher lamellar
frequency, thinner enamel etc.) and tend to be smaller in body size.
They have been placed in a distinct subspecies, M. trogontherii cho-
saricus Dubrovo, 1966 (often given specic rank), in a distinct
species, M. intermedius (Jourdan, 1861) (Labe andGurin, 2005), or in
M. primigenius; their taxonomic status still remains ambiguous
(Lister et al., 2005; PalomboandFerretti, 2005). Goodsamples of this
morphotype have been described from the former Soviet Union,
Germany and Italy (Siegfried, 1956; Ambrosetti, 1964; Palombo,
1972; Dubrovo, 1977; Kotsakis et al., 1978). The Loussik
mammoth differs from these samples by its generally larger dental
and body size, lower lamellar frequency and thicker enamel.
The metrical comparison shows, in conclusion, that the partial
skeleton from Loussik is dentally more advanced than
M. meridionalis, less derived and larger than M. primigenius, and
corresponds well to the upper range of the dental sample from
Senborn, type locality of M. trogontherii.
8. BiostratigraphyePalaeoecology
The site of Loussik did not yield any biochronologic evidence,
as the accompanying fauna is very poor and atypical taxonomically.
The biochronology of the locality has to be based exclusively on the
mammothnd. Theearliest remains of M. trogontherii inEuropedate
from the Early to Middle Pleistocene transition, probably over-
lapping chronologically with the latest M. meridionalis. The typical
M. trogontherii morphotype persisted through the early Middle
Pleistocene, with smaller, chosaricus-type forms appearing during
the later Middle Pleistocene. The good metrical and morphologic
correspondence of the Loussik mammoth with the type material of
M. trogontherii and other samples referred to the typical formof the
species can justify the stratigraphic placing of the site in the lower
Middle Pleistocene, despite its reduced body size in comparison
withother skeletons datedtothis time span. The rather small stature
difference can be well attributed to intraspecic variation, particu-
larly when the geographic distance between the compared samples
is evidently long. Moreover, the Loussik skeleton comprises several
non-fused epiphyses, clearly indicating that this individual was still
growing at the time of death. Also the retention of an ancestral
character, as the thick enamel, if not of local trophic signicance, can
be considered as indicative of an older geochronologic age. There-
fore, an early Middle Pleistocene age is more probable for Loussik.
The typical M. trogontherii from Central-East Europe is consid-
ered as a steppe dweller d thus its vernacular name steppe
mammoth das it is well adapted to open landscapes and aridity. It
initiates an evolutionary line that ended to the extremely speci-
alised, cold-adapted grazer, the woolly mammoth. Recent palae-
oecological data referring to the environment of the West Runton
mammoth (Stuart and Larkin, 2010) have shown that M. trogontherii
may be associated with a temperate forest, but this is still a sole
indication referring in a northern region with anyway cool climate.
The site of Loussik has not produced any palaeoecological
evidence, which could be used for the reconstruction of the palae-
oenvironment. Nonetheless, the nearby lacustrine basin of Mega-
lpolis, situated in central Peloponnese, has yielded a good
palaeoenvironmental record for the Middle Pleistocene, mainly
based on pollen (Okuda et al., 2002). The basin is lled with cyclic
beds of lignite, depositedduringinterglacials, anddetritus, deposited
during the glacials. All the lignite seams yieldpollenof temperate oak
forest (as well as an interglacial mammal fauna rich in E. antiquus,
Hippopotamus sp. and deer), whereas the prevailing plant genus in
the detrital beds is the shrub Artemisia, a strong indication of semi-
arid steppe. The Megalpolis palaeoenvironmental record suggests
that cold and arid steppic conditions did prevail during glacial
periods in a quite southern region as central Peloponnese. This piece
of evidence ts well with the unexpected discovery of a cold-
adapted animal at Loussik. The presence of M. trogontherii in Pelo-
ponnesecan, therefore, begeochronologicallyspottedmoreprecisely
to a cold episode of the early Middle Pleistocene, as those of the
Marine Isotope Stages (MIS) 14, 16 or 18, the climatic conditions of
which would allowthe southern expansion of the steppe mammoth
population.
9. Palaeobiogeographic implications
Despite its apparent expansion during the glacials, M. trogontherii
seems to have beenmuchless abundant inregions of less continental
climatic conditions, as the Western and Southern Europe, away from
its core region in Eastern and Central Europe. In Mediterranean
Europe inparticular (Iberian, ItalianandBalkanpeninsulas) it is quite
uncommon, usually identied on the presence of isolated molars
with trogontherioid morphology. In Italy, the species is totally
unknown southern of Rome (nonetheless the more derived late
Middle Pleistocene mammoths expanded to the South till Capri,
40.5
N) (Palombo and Ferretti, 2005). In Spain, though also rare, it is

reported to have reached as south as the GuadixeBaza basin (about
37.5
N) (van der Made and Mazo, 2001). The Balkan Peninsula is also
poor in M. trogontherii. Radulesco et al. (1965) describe dental
material from the Bras ov area in Romania, while Lenardic (1994)
studied an advanced chosaricus-like skull from Croatia. A recent
important discovery in Eastern Serbia (at Kostolac) refers to
a complete M. trogontherii skeleton (Korac et al., 2010).
The presence of the species in Greece was already reported by
Psarianos (1958) and Marinos (1964) in the north of the country
(localities Sotr in Western Macedonia and Phlippi in Eastern
Macedonia) based on a total of three isolated molars. The poor
photographic documentation (without a metrical scale) and the
absence of any metrical data do not allow a re-evaluation of this
taxonomic attribution. The Philippi specimens are morphologically
very similar to M. trogontherii, though they could also be referred to
an early M. primigenius (a species also reported from Phlippi). The
specimen from Sotr, presumably a left m3, exhibits weak enamel
plication and virtually absent loxodont sinus, but the crown is
rather narrow for a mammoth. More recent material coming from
the same locality (a sand pit) is also referred to Mammuthus
(Velitzelos and Schneider, 1973). However, the provided photo-
graphic and metrical data (narrow crowns, wrinkled enamel etc.)
point clearly to E. antiquus, as is also correctly observed by Tsoukala
et al. (2010). In conclusion, the only until nowplausible evidence on
the presence of M. trogontherii in Greece comes fromthe peat basin
of Phlippi, situated in the NE of the country. The studied skeleton
from Loussik expands considerably to the south the species
biogeographical distribution in Greece.
Fig. 21. Map of the Mediterranean region showing the geographical distribution of the Mammuthus trogontherii-bearing localities according to available data. Loussik is among the
southernmost sites. The question mark denotes samples of problematic taxonomic determination. Data according to Marinos (1964), Radulesco et al. (1965), Horowitz (1977),
Lenardic (1994), van der Made and Mazo (2001), Palombo and Ferretti (2005), Albayrak (2009), Mol and Lacombat (2009), Korac et al. (2010). Composite satellite image source:
NASA World Wind. The mammoth gure in taken from Osborn (1942, p. 1052).
Dental nds attributable to M. trogontherii are also reported from
other countries of the EasternMediterraneanregion. Albayrak(2009)
cites four sites inTurkey, one of which is the typical locality of Elephas
armeniacus Falconer, 1857, a synonymof M. trogontherii. Another site,
Dursunlu, located in SW Turkey near Konya, is almost as far south as
Loussik. A few other dental specimens from Syria and Israel (local-
ities Latamne, Evron, Benot Yaakov) are referred to M. trogontherii
(Horowitz, 1977, based mainly on the studies of Hooijer). Maglio
(1973) considered these specimens as Elephas namadicus
(E. antiquus), but Hooijers specic determinations have been
accepted by other scholars (Adam, 1988, pp. 15e16; Gurin et al.,
1993). In a later review of the Levantine proboscidean remains
Tchernov and Shoshani (1996) consider these samples as taxonomi-
cally problematic, but they do assign some dental fragments from
Latamne to M. trogontherii (Tchernov and Shoshani, 1996, Table 21.1).
Considering these controversial opinions and the generally frag-
mentary state of the available material, it seems that the presence of
M. trogontherii in the Levant is rather ambiguous. Consequently the
biogeographic distribution of the species can be securely extended to
the south to Southern Turkey and Southern Greece in Eastern
Mediterranean (about 38
N), and Southern Spain (Granada) in the

Western Mediterranean (about 37.5
N) (Fig. 21).
10. Conclusions
Given the rarity of associated skeletons referred to M. trogon-
therii, the Loussik skeleton is an important nd, preserving most of
the cranial and postcranial elements of this mammoth. It is the
most complete and well-preserved nd of this species in the
Eastern Mediterranean region and one of the few known skeletons
in Europe. The study of its anatomical and metrical characters
showed that the skeleton belongs to a male, about 45 years old, that
weighted about 8 t and stood in life about 3.80 mhigh. Dentally, it is
very similar to the type sample of the species.
M. trogontherii is extremely rare in Greece and was previously
unknown in the southern part of the country. Its presence extends
considerably to the South the species known geographical distri-
bution in the Balkan Peninsula.
Based on the specic determination of the nd, and taking into
account its particular anatomical characters, the fossiliferous
continental deposits of Loussik can be dated to the early Middle
Pleistocene, quite probably to a cold stage.
Acknowledgements
The Loussik excavation was nanced by the Hellenic Ministry
of Culture (Ephorate of PalaeoanthropologyeSpeleology, Athens,
and ST Ephorate of Prehistoric and Classical Antiquities, Patras);
the Municipality of Olena contributed to the excavation team with
three workers. The team consisted of A. Darlas (archaeologist,
director), A, Athanassiou (geologistepalaeontologist), P. Poly-
doropoulos (preparator), D. Bouzas (geologist), L. Stavropoulou
(designer), S. Stamatopoulos, G. Martzaklis, A. Chrysikopoulos, N.
Spiliopoulos and G. Tsironis (workers). The nds were prepared in
the laboratory of the Ephorate of PalaeoanthropologyeSpeleology
by P. Polydoropoulos, P. Ghioni, V. Papamikou and V. Klaridi. The
geologist E. Ypsilanti helped with the measurements. A recent re-
examination of the Loussik partial skeleton (currently in the
collection of the Archaeological Museum of Patras) was facilitated
by E. Kollia and A. Paraskevopoulos, archaeologists of the ST
Ephorate of Prehistoric and Classical Antiquities. M. Golnopoulou
(ST Ephorate of Prehistoric and Classical Antiquities) provided
a copy of one of the excavation plans. G. Iliopoulos (University of
Patras) offered hospitality during my stay in Patras and provided
useful comments and discussions on the elephant osteology and
taphonomy. An anonymous (Bavarian State Collection for Palae-
ontology and Geology) and Yasemin Tulu (Calvert Marine Museum)
greatly helped in the search of hard to nd publications. Two
anonymous reviewers are especially thanked for critically reading
the manuscript and providing useful comments.
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A skeleton of Mammuthus trogontherii (Proboscidea, Elephantidae) from NW

. The maxilla is particularly high to

between them). A similar morphology is

near the cement/dentine

(the angle increases

between them. The right

), similar to the extant

N) (Palombo and Ferretti, 2005). In Spain, though also rare, it is

N), and Southern Spain (Granada) in the

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