5

GEOLOGICA BALCANICA, 35. 34. Sofia, Dec. 2006, p. 540.

Introduction
The fossil materials presented herewith to the scien-
tific community were found by Dimitar Kovachev in
1965. The late Ivan Nikolov has worked for a long
time on their restoration, with minor participation of
D. Kovachev. In 1972 Nikolov reported to the De-
partment of Palaeontology at the Geological Insti-
tute, Bulgarian Academy of Sciences, that the resto-
ration process was finished. The assembled skeleton
has been mounted and permanently exhibited in the
Museum of Geology and Palaeontology, Sofia Uni-
versity !St. Kliment Ohridski". A diminished (3/4 of
normal size) copy has been prepared and is now on
exhibit at the Asenovgrad Palaeontological Branch
of the National Natural History Museum. Both Ivan
Deinotherium thraceiensis sp. nov.
from the Miocene near Ezerovo, Plovdiv District
Dimitar Kovachev
1
, Ivan Nikolov
2
1
Asenovgrad Palaeontological Branch, National Natural History Museum, Asenovgrad
2
formerly at the Geological Institute, Bulgarian Academy of Sciences, 1113 Sofia
(Submitted: 07.08.2004; accepted for publication: 18.12.2006)
. , . Deinotherium thraceiensis sp. nov.
, . # -
, -
. 1965 ,
, . -
. , - -
Trilophodon angustidens Cuvier, Hipparion microtaton
Nicolov Mastodon sp.
, , , ,
. -
.
. -
. ,
Deinotherium thraceiensis sp. n.
Abstract. The paper presents to the international palaeontological community a description of
the full skeleton of a deinothere found and excavated near the village of Ezerovo by D. Ko-
vachev in 1965, and described, restored and mounted by the late Ivan Nikolov. The fossil remains
are found in sediments of Maeotian age. Former excavations have yielded fossil bones of Trilo-
phodon angustidens Cuvier, Hipparion microtaton Nicolov and parts of mandible of Mastodon sp.
When the whole skeleton was excavated only a few bones were missing, and namely, the whole
rear left thighbone, the right rear fibula, most of the ribs as well as some caudal vertebrae. They
are restored on the basis of symmetrical analogues existing. The measurements on the bones
found and the comparisons with other deinotheres described in the literature showed certain
differences. Some of the differences relative to the skull have certainly a taxonomic value. There-
fore, the deinothere described is referred to a new species, Deinotherium thraceiensis sp. n.
Kovachev, D., Nikolov, I. 2006. Deinotherium thraceiensis sp. nov. from the Miocene near
Ezerovo, Plovdiv District. # Geologica Balc., 35, 34; 540.
Key words: Deinotherium thraceiensis, Miocene, Maeotian, Bulgaria.
6
Nikolov and Dimitar Kovachev were fully aware of
the importance of this very rare case of finding a
whole skeleton of such a huge extinct animal, and
they estimated it to be a representative of a new spe-
cies of genus Deinotherium. Ivan Nikolov, well-known
for his thoroughness, wanted to make an exemplary
publication but his early death stopped for a long
time the work on it. The results of his enormous la-
bour put into the description of the bones, the resto-
ration and mounting of this unique skeleton, re-
mained in the archives of the Geological Institute.
After the sudden untimely death of Nikolov in 1984,
the work on the publication remained unfinished.
Now, D. Kovachev decided to prepare the materials
for publication with the clear idea about the respon-
sibility undertaken, and that the whole blame for even-
tual flaws and errors should be addressed to himself.
Such publication would be of importance for the
future studies on genus Deinotherium.
Many scientists have helped the authors in dif-
ferent time. Prof. Dr. R. Dehm (Director of Univer-
sitt-Institut f#r Palontologie und historische Geolo-
gie # M#nchen) and Prof. Dr. H. Tobien (Director
of Palontologisches Institut der Johanes-Gutenberg
Universitt # Meinz) helped I. Nikolov during the
first studies and restoration. Academician T. Nikolov
(Geological Institute, Bulgarian Academy of Scienc-
es) gave valuable advice with the material. D. Ko-
vachev wishes to express his gratitude to Dr. Marin
Ivanov from the Sofia University for his responsive-
ness to the problems of this research. Dr. N. Spassov
(National Natural History Museum, Sofia) kindly
helped with the literature I. Nikolov had used at the
library of the Museum.
Studies on the genus Deinotherium
a historical review
As early as the 17 century a place near Lyon, France,
was known as !the field of the giants" because of the
large animal bones often found there. Some of those
bones came to Matsorier # a surgeon, who used to
show them for years in France and Germany as the
bones of king T%teboch. Much later, the real tomb of
the king was found, and the deceit was exposed. The
huge bones were transferred to the Natural History
Museum in Paris. Probably these are the first Deinothe-
rium remains found.
One century later, in 1715, R'aumure admitted
he could not refer the bones to any known animal.
Kenedy supposed in 1775 that the materials were re-
lated to mammoths. G. Cuvier thought in 17791836
that the animal had been a large tapir coexisting with
mastodons. He called it Tapir gigantesque and
thought that its tusks were curved upwards.
Kaup created in 1829 the genus Deinotherium with
the species D. giganteum upon the skull and mandi-
ble found near Eppelsheim, Germany. He thought
this was an intermediary form between the sloth and
the mastodons and referred them to the larger taxo-
nomic unit Curtognati. In 1841 and 1857 he de-
scribed further remains of that genus found near
Eppelsheim and Westhofen. According to him, the
adult animals had five teeth in their upper and low-
er jaws. In 1833 he described the new species D. cu-
vieri, but because it was often found in Bavaria, H. v.
Meyer called it D. bavaricum. This name is still used
today. However, arguments about the exact place and
the life environment of deinotheres still continued.
P. Gervais accepted in 1848 the opinion of Kenedy
and Koch that the genera Deinotherium and Masto-
don should be referred to Proboscidea. He recognized
the existence of three species: D. giganteum Kaup =
Tapir gigantesque Cuvier; D. intermedium and D. cu-
vieri Kaup. S. Pictet reported in 1853 about the finding
near the village of Absdorf of a whole skeleton of D.
giganteum Kaup. Only the head, the first two vertebrae,
the corpus of a thoracic and a caudal vertebra and
fragments of the limbs were preserved. These remains
showed that it is close to the mastodon and belongs to
Pachydermae. Four deinothere species were recognized
in 18581859 by M. Lartet, and namely, Deinotherium
bavaricum H. v. Meyer, D. giganteum Kaup, D. sp. (an
intermediary form between the first two), and D. cu-
vieri Kaup. The first species is known from the Mi-
ocene of France and Bavaria, the second from the Late
Miocene of France, Germany and Greece, the third
and fourth from the Miocene of France. After Lartet,
the dental formula of genus Deinotherium is: I 0-0/1-1;
Pm 2-2/2-2 and M 3-3/3-3 for the permanent teeth,
and for the deciduous # I 0-0/1-1; deciduous molars
3-3/3-3. New fossil finds in Greece (Pikermi), India (Si-
walik), Europe (France, Switzerland, Austria), etc. have
been reported during the last quarter of the 19
th
centu-
ry by A. Gaudry, H. Hensel, B. A. Lydekker, E. Chantre,
M. Vacek, V. Biber, C. Deperet, and other authors. Gr.
Stefanescu first reported in 1891, and then described
(1895#1910) a deinothere skeleton found near Man-
zati, Romania. Because of the large dimensions of all
its bones and teeth, he named it D. gigantissimum. Ehik
(1930) published upper and lower molars and some
finger bones, found in the Pliocene sediments of Hun-
gary, as a new species # Prodinotherium hungaricum.
Osborn (1936) changed the name of the genus (the
Latinized form Dinotherium) to the ancient Greek
form Deinotherium. He revised all then known
deinothere materials, discussing in details each spe-
cies, and illustrating skulls/mandibles and teeth from
D. giganteum Kaup, D. bavaricum H. v. Meyer, D.
cuvieri Kaup, D. pentapotamiae Lydekker, D. indi-
cum Falconer, etc. (Fig. 1).
Numerous new finds of deinotheres have been re-
ported during the whole 20
th
century from Austria,
Bohemia, Bulgaria, France, Germany, Greece, Hun-
gary, Macedonia, Moldova, Pakistan, Romania, Rus-
sia, Serbia, Ukraina, etc.
Studies on the genus Deinotherium
in Bulgaria
The first Deinotherium remains in Bulgaria have been
found by G. Bonchev (, 1897) who made ex-
7
Fig. 1. Distribution of genus Deinotherium in Southeastern Europe and Asia Minor
cavations in the Middle Sarmatian limestones near
Nessebar, studying the hipparion fauna. Three years
later he reported new finds from the same locality.
(1911/1913) published the two deinothere
mandibles excavated by G. Bonchev, together with
other single teeth from the village Sovolyano, near
Pleven, the village Archar and the village Mazgosh.
They were all brought to the Institute of Geology by
different people. From these materials Bakalov de-
termined two species # D. giganteum Kaup race mi-
8
nor and D. giganteum Kaup race major. To the first
he referred the smaller teeth, and to the second #
the larger ones. Later on (, 1949/1950)
Bakalov described Deinotherium teeth found at the
!Meander of the Konska river" near the village No-
evtsi; the village (now # a town; for some time called
Batanovtsi) Temelkovo (Pernik region) and Aksako-
vo, near Varna, as D. giganteum Kaup race major
(from the first two localities, Pliocene) and D. gigan-
teum Kaup race minor (from the Sarmatian lime-
stones near Aksakovo). (1960) reported a
molar fragment of D. giganteum Kaup race major
from the lignite coals near the village Hrabarsko (So-
fia region), assuming a Pontian age. The whole fossil
material belonging to genus Deinotherium then known
from Bulgaria has been published by ,
(1962). They recognize two races # minor
and major, and refer the finds as follows: two man-
dibles, an upper jaw and three separate teeth to race
minor, and a mandible, seven complete upper and
lower teeth and three tooth fragments to race major.
The age of both races varies from Sarmatian to Le-
vantian. (1962) referred the molar and frag-
ment of a scapula found in the Levantian sediments
near the village Popitsa, Vratsa region to D. gigan-
teum Kaup. , (1966) described a
lower molar found by the latter author in the sands
in the valley of Cherkezitsa river, near the village
Ahmatovo. They assume the age to be Levantian be-
cause of the Anancus arvernensis Croizet et Jobert
teeth found there, referring the find to D. giganteum
Kaup. (1966) reported the finding of an
almost complete skeleton of Deinotherium near the
village Ezerovo, Plovdiv region. This same skeleton is
the subject of the present paper. ,
(1966) published a brief review of the proboscideans
in Bulgaria, and gave also some preliminary infor-
mation about the finding of Kovachev reporting
metrical data on some bones.
Distribution of genus Deinotherium
in Southeast Europe and parts of Asia
Minor
Until now, more than 80 localities (partly shown on
Fig. 1) within the region have yielded deinothere re-
mains, and their number is constantly growing.
There are 16 localities from former Yugoslavia:
1. Veles, Mokranje, Smederevska palanka, Resnik,
Meduhana, Vracevic, Duboko, Tsiganski potok, and
between the villages Vite*evo and Porodin # D. gi-
ganteum Kaup. Near Veles and Ravanica, together
with D. giganteum Kaup, D. bavaricum H. v. Meyer
is also present. The sediments near the villages Medu-
hana, Ravanica and those between Vite*evo and
Porodin are of Lower Sarmatian, and the rest # of
Pontian age.
2. Drenovica, Chukovica, Bresnica, Prebrezka, Zdar-
ski and Kriva reka # D. bavaricum H. v. Meyer. The
age cited is Helvetian-Tortonian, Lower Sarmatian,
Sarmatian and Middle Miocene.
In Greece D. giganteum Kaup is known from four
localities: Pikermi, Thessaloniki and Tanagra (Pon-
tian), and Samos (Middle Pontian). From the Sar-
matian of Chios D. bavaricum H. v. Meyer has been
described.
From Turkey (Asia Minor), D. giganteum Kaup
has been described from K#-#k-ekmece (Maeotian).
From the territory of Romania and part of Bessa-
rabia, two taxa are known # D. giganteum Kaup from
the Pontian and Dacian sediments near Vernesti,
Culm, Lichtental, Pelineu, Telenesti, Visterniceni,
Baimaclia, Cimislia, Comanesti, Curtea and Solcus-
tria; and D. gigantissimum Stefanescu, # from the Plio-
cene sediments near Manzati, Breaza and Cimislia.
From the territory of Moldova and Bessarabia, two
taxa are known too: 1. D. giganteum Kaup near Tara-
klia, Ciobruciu, Sofijevka, Tanovka, Chernevo, Ti-
PLATE I
Deinotherium thraciensis sp. n.
1, 2. Skull in lateral and ventral view. x 4.5% coll. SU.M. No. SU Pl 312/1; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE II
Deinotherium thraceiensis sp. n.
Skull in dorsal view. Scale ca. 4.2% coll. SU. M. No. SU Pl 312/1; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE III
Deinotherium thraceiensis sp. n.
1. Skull and mandible. Scale ca. 2.9% coll. SU.M. No. SU Pl 312/1, SU Pl 312/2
2. Skull from behind # os occipitale. Scale ca 5.6% coll. SU.M. No. SU Pl 312/1; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE IV
Deinotherium thraceiensis sp. n.
1. Upper cheek teeth # P
3
to M
3
sin et dext. Scale ca. 25% coll. SU.M. No. SU Pl 312/1; Locality: Ezerovo, near Plovdiv; Level:
Maeotian

PLATE I PLATE II
PLATE III PLATE IV
PLATE V
PLATE VI
PLATE VII
PLATE VIII PLATE IX
PLATE X PLATE XI
PLATE XII PLATE XIII
PLATE XIV PLATE XV
PLATE XVI PLATE XVII
PLATE XVIII
PLATE XX
PLATE XIX
PLATE XXI
PLATE XXII
PLATE XXIII
PLATE XXIV
9 2 Geologica Balcanica, 3-4/2006
Fig. 2. Distribution of genus Deinotherium in Bulgaria: 1. Ezerovo; 2. Nesebar; 3. Izgrev; 4. Parvomai; 5. Ahmatovo; 6. Lyubovishte;
7. Sovolyano; 8. Batanovtsi (Temelkovo); 9. Noevtsi; 10. Sofia; 11. Kremikovtsi; 12. Hrabarsko; 13. Aldomirovtsi; 14. Katina; 15.
Kula; 16. Archar; 17. Mihaylovo; 18. Glozhene; 19. Popitsa; 20. Pleven; 21. Novo selo; 22. Aksakovo; 23. Yarebichna; 24. Breznik;
25. Hadjidimovo; 26. Varna # Galata; 27. Kalimantsi; 28. Konyovo; 29. Rogozen; 30. Gabra # former open pit !Bolshevik"; 31.
Mazgosh (now on Serbian territory)
PLATE V
Deinotherium thraceiensis sp. n.
1. Mandible. Scale ca. 8.3% coll. SU.M. No. SU Pl 312/2; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE VI
Deinotherium thraceiensis sp. n.
1. Lower cheek teeth # P
3
to M
3
. Scale ca. 25% coll. SU.M. No. SU Pl 312/2; Locality: Ezerovo, near Plovdiv; Level: Maeotian

rasspol, Donau, Velikovo and Krivoy Rog. The sedi-

ments are of Maeotian, even Upper Sarmatian age.
2. D. gigantissimum Stefanescu near Volchinec, Rah-
ny-Lessovye and Pripecheni. The host sediments are
also of Maeotian and Upper Sarmatian age.
A considerable number of deinothere localities
are known from the territory of Bulgaria (Fig. 2).
Some of the first Deinotherium finds in Bulgaria
came from the locality Mazgosh (now in Serbia).
The larger number of localities in the western part
of the country is a result from the fact that this
part is better studied. Some of the more important
finds come from Nessebar (by G. Bonchev), No-
evtsi, locality !Meander of the Konska river" (by P.
Bakalov), and the village Ezerovo, !Kolnu dere"
(by D. Kovachev).
Some of the materials have been found and
brought to the specialists by common people (farm-
ers, workers, etc.). Therefore, the exact location has
been mentioned only approximately, and there is no
stratigraphic profile. In such cases, a probable age
has been assumed after the age of the sediments in
the noted areas.
The vertical distribution of deinotheres in Bul-
garia starts in the Sarmatian, includes the Maeotian
and Pontian and ends in the Pliocene. The remains
of D. bavaricum H. v. Meyer are found only in Sar-
matian sediments, while D. giganteum Kaup is present
from the Middle Sarmatian up to the Romanian.
Stratigraphic notes on the sediments
near Ezerovo, Plovdiv region
The village of Ezerovo is situated ca. 30 km east of
Plovdiv (Fig. 3). Sediments of Eocene, Miocene,
Pliocene and Pleistocene Age crop out in the area
(Fig. 4).
The Upper Eocene (Priabonian) is represented by
light organogenic limestones, rich of nummulites and
corals. South of the village the dam wall is built onto
them. The thickness exposed is about 50 m. In a bore-
10
Fig. 4. Stratigraphic profile of the locality Ezerovo
Fig. 3. Position of the locality of Ezerovo in the Upper Thracian plain
11
hole (No 16; , , 1965) situated
near the village of Ezerovo, these rocks are covered
by massive limestones, marls, and well-cemented
quartz sandstones. They are referred to the Oligocene
on the basis of rich foraminifer and ostracod fauna.
These rocks pass into the Lower Miocene dark gray
aleurolites and marls with foraminifer and ostracod
fauna similar to the Aquitanian-Burdigalian faunas
reported in France (, , 1965). The
maximum thickness of the Lower Miocene is about
180 m.
The Palaeogene and Lower Miocene marine sed-
iments are unconformably covered by fluvial sedi-
ments referred to the Ahmatovo Formation (-
, , 1979). They crop out at nu-
merous places in the valley of the river Cherkezitsa.
At the basis there are aleurolites and shales covered
by sands of light rusty colour, alternating with con-
glomerate lenses. At places the sand layers are cross-
bedded. Petrified trees occur in the sands. The thick-
ness of the sand layers varies from 1.5 m to 7.5 m;
and that of the conglomerates is up to 1.5 m. The
whole thickness of the sands has been revealed only
by drillings and is up to 515 m. Ahmatovo Forma-
tion is referred to the Maeotian, Pontian and Da-
cian regional Paratethys stages. Three macrocycles
have been distinguished (, -
, 1979). The first macrocycle is referred to the
Maeotian (possibly also upper parts of the Sarma-
tian) on the basis of Lower to Middle Turolian (!Pik-
ermian") mammal fauna. The Pontian age of the
lower parts of the second cycle is determined by Upper
Pikermian (= Upper Turolian) fauna, and the up-
per parts of the second cycle are referred to the Da-
cian Regional Stage by Ruscinian fossil remains. The
fossil locality of Deinotherium thraceiensis sp. n. be-
longs to the first cycle of the Ahmatovo Formation
(Nikolov, 1985, p. 58).
About 800 m west of the village of Ezerovo, right
from the road to the dam, at the fork with the road
for the pumping station, in light medium-grained
sands, D. Kovachev found in 1965 the skeleton of
Deinotherium thraceiensis sp. n. described in the
present paper. It was lying at a depth of ca. 40 m
relatively to the level of borehole No 16. At the same
level, but 50 m to the east, V. Tsankov and I. Vaptzarov
have found in 1956 a part of a skull, determined by
P. Bakalov as Trilophodon angustidens Cuvier. In 1962
St. Stoykov and I. Nikolov found 45 m below the
find described here, teeth of Hipparion microtaton
Nikolov and a part of a mandible of Mastodon sp.
They all indicate a Maeotian age. Above the
deinothere have been found teeth of Hipparion med-
iterraneum Gervais and Tetralophodon longirostris
Kaup. They unequivocally prove a Pontian age.
After 1965, D. Kovachev collected the following
fossils during numerous field trips:
1. A skull, a mandible and bones from the limbs of
Mastodon sp. In the structure of its molars it resem-
bles T. angustidens Cuvier, but the tusks and the skull
are different from this species.
2. A whole mandible with a preserved symphysis and
M
3
dext et sin of Choerolophodon sp.
3. Two maxillary fragments with M
3
also belonging
to Choerolophodon sp.
4. M
3
sin et M
2
dext # Choerolophodon sp.
5. Right semimandible with M
3
# Tetralophodon lon-
girostris Kaup.
6. M
3
sin # Anancus sp.
7. M
2
and M
3
dext and M
3
sin # Zygolophodon bor-
soni Hays.
8. M
3
dext # Zygolophodon borsoni Hays.
9. M
3
sin Deinotherium giganteum Kaup.
10. Right semimandible # Microstonyx major.
They will be subject of other publications.
The Neogene sands are covered by a layer of soil
1.0 to 1.5 m thick including isolated rock fragments
of different size. These soils are referred to the Pleis-
tocene.
Palaeontological part
Order Proboscidea Illiger, 1821
Suborder Deinotherioidea Osborn, 1921
Family Deinotheriidae Bonaparte, 1845
Genus Deinotherium Kaup, 1829
Type species of the genus Deinotherium giganteum
Kaup, 1829
Deinotherium thraceiensis sp. n.
(Pl. I # XXII)
Derivatio nominis: thraceiensis from Greek !Thra-
ceia" # Thrace, the region of the find.
Holotype. As such we design all the bones of the skel-
eton # No. 312/1 to No. 312/23-30, collection of the
Museum of Geology and Palaeontology of the Sofia
University !St. Kliment Ohridski".
Locality. Light, medium-grained sands (Ahmatovo
Formation, first cycle) near Ezerovo, Plovdiv region,
just at the turn for the pumping station from the
road to the dam.
Age. Maeotian.
Diagnosis. Large animals, skull short and high. Na-
sal bones short, fusing at the anterior ends, curved
and slightly flattened laterally. Large external nares.
Ear area also large and with trapezoid shape. Eye
sockets almost separated from the ear areas. Fore-
head high and, compared to the external nares #
short. The occipital bone is high and wide. Symphy-
sis of the mandible curved downwards and back-
wards. Incisors of moderate length, slightly curving
outwards. Their tips end exactly beneath processus
angularis. Neck short, with seven thin cervical verte-
brae. First ones lack processus spinosus, in the last
ones it is weakly developed. Skull and mandible, rel-
atively to the size of the whole skeleton, are much
smaller than in the other proboscideans. Fingers high,
with large phalanxes # hoofs, strongly developed on
the third finger.
12
Fig. 5. Deinotherium thraceiensis sp. n.
Description and comparison
Skull. (Pl. I, fig. 1 and 2; Pl. II, fig. 1 and 2; pl. III, fig.
2; Fig. 5 # A, B, C and D).
The skull described here is entirely preserved. It is
very short and its posterior part # high (Fig. 5).
The premaxillary bones (os premaxilarae), which are
strongly developed in Proboscidea, do not merge at
their anterior ends and are slightly curved downwards.
There are no upper tusks and alveoli for them in
these bones. They are widening in front of the eye
sockets. All their surface is smooth, without any rug-
osity.
Nasal bones (os nasale) are short, high in their ante-
rior part, rounded and laterally flattened. On the
inner side, in the region of the maxillary bones, the
nasals are slightly concave, so that together with the
premaxillary bones they have a rounded, wedge-like
shape (Fig. 5, A). They are connected by a strong su-
ture. The nasal canal has an inner width of 70.0 mm
and length to the end of the external nares 710.0 mm.
External nares (Fig. 5, ex. n., and Pl. II, figs 1 and 2)
pear-shaped and rather large. They start almost at
the beginning of os nasale, gradually becoming wid-
er and deeper. By the beginning of the eye socket
they narrow, then becoming straight and ending with
a regular, arch-shaped curve just at the beginning
of the ear area. They penetrate for 70#80 mm more
and end in the oscular area with the dorsal nasal
opening, which is small and oval, parallel to the fore-
head. On the bottom, parallel to the nasal cavity, cris-
ta conchalis dorsalis is outlined. (Measurements of
the external nares in Table 1).
The calvaria # os frontale and os parietale (fronto-
parietal part) starts against the beginning of the ear
area ends with os occipitale. In the middle it is slightly
concave, and in the posterior part # strongly convex
(measurements in Table 1).
Occipital bone (os occipitale). (Fig. 5 A, C, D ocs;
Pl. III, Fig. 2). High and wide, laterally slightly round-
ed, under the condyles and by the parietal bone al-
most straight and parallel. On both sides its wings
are curved backwards, and in the region of the
condyles and above them it is slightly convex. In re-
lation to the forehead this bone is almost perpendic-
ularly situated, at an angle of 80 (measurements in
Table 1).
Occipital condyles. (Fig. 5; Pl. I, Fig. 1 and 2).
They are situated in the middle and in the lower part
of the occipital bone. Their shape is irregular and
oval. Wider and convex in their upper part. They
surround foramen magnum # fm, which has an oval
shape and is situated perpendicularly to the occipi-
tal bone (measurements in Table 1).
Eye socket (orbita) (Fig. 5 A orb.) is preserved only
on the right side of the skull. In front it is limited by
the massive and laterally strongly protruding os lac-
rimale, and in the back # by the well pronounced
processus postorbitale, which is long and thin. From
below it is limited by os zygomaticum. Thus the eye
socket is almost entirely separated from the ear area
(measurements in Table 1).
13
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14
Ear area. (Fig. 5 A, C or. r and Pl. I, Fig. 2). It is high,
wide and deep. Begins immediately behind the eye
socket. In the posteriror part it borders os occipitale.
Maxillary bone (Maxilla) (Fig. 5 B m and Pl. I, Fig. 2).
Wide, slightly concave at the palatum. Length
400.0 mm, height at M
1
250 mm. Facial crest and
infraorbital canal not pronounced.
Palatum durum. (Fig. 5 B; Pl. I, Fig. 2). Long and
very narrow. Strongly concave in the middle (mea-
surements in Table 1).
Upper cheek teeth (Pl. I, Fig. 2 and Pl. IV, Fig. 1).
Both toothrows are entirely preserved. Only the first
molars are slightly worn out, so that part of the den-
tin is seen. There is no difference in the structure of
the left and the right ones. Their sizes differ with 1 to
3 mm (see Table 2) but this is assumed to be normal.
We shall discuss only the morphological differenc-
es, where there are such.
The third premolar (P
3
dext) is large, of almost
regular trapezoid shape. It has one large outer ridge
and two inner, situated diagonally to it. Their inner
ends touch each other, and the outer ends the
anterior and the posterior part of the outer ridge cor-
respondingly. Thus the three ridges surround a deep
triangular valley. The anterior inner ridge is wider,
with a triangular shape, slightly worn out at the tip.
Its inner side is concave and, together with the cin-
gulum on this side of the crown, surrounds a small
and shallow triangular valley. On the inner side it is
slightly more worn out. The cingulum is well devel-
oped on the whole anterior and the posterior outer
part of the crown.
The fourth premolar (P
4
dext) has an almost tet-
ragonal shape. It consists of one outer and two inner
ridges, situated perpendicularly to the outer. The
anterior inner ridge is wider than the posterior and
less worn out. It has the shape of a isosceles triangle,
the base of which touches the inner anterior part of
the outer ridge. At this place the two ridges surround
a shallow valley. The posterior ridge is narrower and
less worn out. By touching the outer ridge it is divid-
ed in two parts, and V-shaped. Thus the ridges sur-
round a deep triangular valley with steep walls. On
the inner and outer posterior side and on the anteri-
or side of the crown there is a slightly developed cin-
gulum. The left P
4
has a weaker cingulum on its an-
terior side.
The first molar (M
1
dext) is long and narrow. It
consists of three ridges. First two are more worn out.
The three ridges on the inner side are wider. They
gradually become thinner toward the outer wall of
the crown. At the very end they are slightly curved
downwards but they don!t touch each other. They
are separated by small valleys narrowing in their mid-
dles. The second and third ridges are slightly con-
cave on the posterior side, while the anterior side of
the third ridge is more convex. A tubercle at the base
of each valley, but only from the inner side, closes
them. The valleys are open outwards. In front, the
crown has a cingulum which, together with the an-
terior wall of the first ridge, surrounds a long and
shallow valley.
The second molar (M
2
dext) is large and almost
square. It is built by two thick and high ridges which
are slightly curved backwards on the outer side. Thus
on the posterior wall of the ridge there is a small
concavity, while in front they are straight. A deep
valley with steep walls separates them. Both teeth (left
and right) have a clear cingulum on the anterior
and the posterior sides. In the anterior part of the
crown the cingulum is something like an anterior
talon and has a worn-out tip.
The last molar (M
3
dext) is the largest. It has two
thick ridges, slightly convex forwardly. A deep and
wide valley separates them. The ridges are wider on
the inner side and slightly curved backwards on the
outer side. They form a small concavity on the poste-
rior side. The valley between the two ridges is free.
The cingulum is well developed both at the anterior
and the posterior part, shaping an anterior and pos-
terior talon.
Comparison
According to the literature, there are only two com-
pletely preserved skulls found up till now. The first
has been found near Eppelsheim in 1835, described
by Andrews (1921) as D. giganteum Kaup and is
stored at the British Museum. The second has been
found near the village Gussiatin, Tirasspol region,
by a palaeontological expedition from the Institute
of Zoology, Academy of Sciences of the Ukrainian
SSR in 1963, described by Svistun in 1974 and re-
ferred by him to D. levius Jourdan. In Vienna there
is a mounted skeleton of D. bavaricum H. v. Meyer
but its skull is entirely reconstructed (see Pl. XXII).
Similar is the case with D. gigantissimum Stefanescu
in Bucharest. From the skull of the deinothere skele-
ton recently mounted in Kishinev only part of the
maxilla with the upper teeth is preserved.
In conclusion, it seems that the only possible com-
parisons are with the skulls from Eppelsheim and
Gussiatin. The main differences between them and
the skull described by us (cf. Figs. 5, 6 and 7) are as
follows:
1. Relatively to the body size, the skull described here
is short, but high. Compared to the other two, the
Ezerovo skull is highest, and the one from Gussiatin
the lowest.
2. The premaxillary bones have interesting differences
(Figs. 5, 6 and 7 pmx). In the skull described by us
they are curved downwards as in the Eppelsheim skull,
but are not so strongly flattened, the part not knit
together is much smaller and the incisure between
them is narrower and shallower. In the Gussetin skull
the curving is weaker. In this species, in their pre-
orbital part the lateral processes are much stronger than
in both our and the Eppelsheim skull. Besides, in the
Gussiatin skull the premaxillary bones are coarsely rug-
ged with a tuberosity protruding forward. On it, ac-
cording to Svistun 1974, there could be horn-like struc-
tures substituting for the upper tusks. There is no such
thing in our material, neither in the Eppelsheim skull.
15
D
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18
Fig. 6. Deinotherium levius Jourdan
Fig. 7. Deinotherium giganteum Kaup
3. The external nares (Figs. 5, 6, and 7 exn) in D.
thraceiensis sp. n. is large, deep and pear-shaped. It
starts from the very beginning of the nasal bones,
gradually widening and taking two thirds of the
length of the whole skull. In the other species it is
shorter and almost of the same width in its anterior
and posterior parts.
4. The nasal bones in D. thraceiensis sp. n. are short,
narrow, frontally rounded and fused. In the other
species they are longer, wider, almost flat for most of
their length, and separated. In the Gussiatin species
they protrude above the external nares. This is not
the case with the two other skulls (Figs. 5, 6, and 7).
5. The calvaria of the skull (fronto-parietal part, Figs.
5, 6, and 7 Ic in D. thraceiensis sp. n.) is high and
wide). In the other two species it is lower and shorter.
In the Gussiatin species it is even strongly concave
in its anterior part.
6. The eye socket (Figs. 5, 6, and 7 orb) in the
Ezerovo skull is large and separated from the ear
area. In D. giganteum Kaup they are almost fused,
and in D. levius Jourdan from Gussiatin the situa-
tion is closer to our case.
7. The zygomatic arch (Figs. 5, 6, and 7 zyg) of D.
thraceiensis sp. n. with its developed proc. postorbit-
alis resembles the one of D. levius Jourdan, but in its
posterior part, behind its connection with the tem-
poral bone, it lacks the processus that Svistun notes.
He regards this processus as related to the develop-
ment of the lower tusks. No comparison with the zy-
gomatic arch of the Eppelsheim skull is possible,
because it is not preserved.
19
8. There are significant differences between the three
skulls in the occipital region. Os occipitale in D. thra-
ceiensis sp. n. is high, wide and at an angle of 80
toward the forehead, and of ca. 70 to the plane of
the teeth. In D. giganteum Kaup these angles are 70
and 50 correspondingly; in D. levius Jourdan the
declination of the occipital bone is even larger, so
the angle is only 60. As a whole, the skull of D. levius
Jourdan is very flat and low. Os occipitale is visibly
concave and very wide, forming two lateral wings
(Fig. 6 C and D OS). Those wings are almost lack-
ing in D. thraceiensis sp. n. and in D. giganteum
Kaup they are much less developed.
9. The position of the occipital condyles is rather
different (Figs. 5, 6, and 7 oc). In D. thraceiensis
sp. n. they are situated in the middle and in the low-
er part of the occipital bone, almost the same is their
position in D. giganteum Kaup, and in D. levius Jour-
dan they are on the upper half of the noted bone.
According to Svistun, this permitted the animal to
raise its head almost at right angle to its neck, thus
fully using its back curved lower tusks.
10. Worth mentioning is a characteristic peculiarity
of the Gussiatin skull lacking in all the others,
including the skull from Ezerovo. This skull has a
double articulation with the mandible. Once by an
articular surface on the zygomatic process of the
temporal bone and then by a second surface on the
petromastoideum. Between those two surfaces is the
external meatus of the auditory canal. How and
when this second articular surface was used, is
unclear.
After comparing their anatomy we can see that
deinotheres could be divided in two groups. One in-
cludes D. giganteum Kaup, D. gigantissimum Stefa-
nescu, D. thraceiensis sp. n., D. levius Jourdan and
D. indicus Falconer, which have larger teeth, and the
second D. bavaricum H. v. Meyer, D. pentapota-
miae Lydekker and others, all with smaller teeth.
As a whole, the teeth are rather similar and differ-
ences are seen mainly in the two premolars P
3
and P
4
.
In D. giganteum Kaup P
3
is more rounded on the
inner side. The two inner ridges are more perpendic-
ular to the outer one. The anterior inner ridge is not
connected with the outer and is at an acuter angle to it
than in other taxa. They almost don!t touch each other
on the inner side. If anything like that is observed in
these species, than it is down at the very basis. The
ridges merge when they are worn. The cingulum is
more clearly seen here. It consists of numerous large
and small tubercles on all sides of the crown.
No comparison with the third premolars of D. gi-
gantissimum Stefanescu is possible, because they have
been restored after D. giganteum Kaup.
In D. levius Jourdan the anterior inner ridge of P
3
is situated diagonally to the outer one and touches
it. The posterior one is wider and V-shaped in its
upper part, with a shorter anterior part. It is also di-
agonal to the outer ridge, but does not touch it.
In D. indicum Falconer the inner ridges are some
more diagonally situated, compared to D. giganteum
Kaup, but their tips do not contact.
D. bavaricum H. v. Meyer has small P
3
s. Their
inner ridges are some more perpendicular to the outer
one. This is rather clear with the anterior inner ridge.
The valley between them is wider and deeper.
In D. pentapotamiae Lydekker, D. levius Jourdan
and D. indicum Falconer the differences are almost
the same, concerning the position of the two inner
ridges and the size of the tooth and its cingulum.
The fourth premolar of D. thraceiensis sp. n. has
inner ridges perpendicular to the outer one. The
posterior is narrower and longer. The triangular val-
ley surrounded by the posterior inner wall of the out-
er ridge and the inner wall of the posterior ridge is
clearer than in all the other species.
D. gigantissimum Stefanescu has a longer P
4
. Its
transverse ridges are at a larger distance from each
other. Besides, it has a strong cingulum on the ante-
rior and the posterior side, lacking in our species.
In D. bavaricum H. v. Meyer P
4
s are smaller and
of a more regular square shape. The transverse ridg-
es are more distant at the inner side of the crown,
and the valley surrounded by them and the outer
ridge is wider and larger.
In D. levius Jourdan the posterior inner ridge is
also attached to the outer one with its longer posteri-
or branch.
In D. indicum Falconer and D. pentapotamiae
Lydekker the last premolars have a square shape and
are smaller. Their transverse ridges are parallel and
of equal size. The valley between them is larger and
everywhere equally wide.
In the M
1
s of D. giganteum Kaup, unlike D. thra-
ceiensis sp. n., the three ridges are almost of equal
size, with an equally developed cingulum on both
sides of the valleys, blocking them entirely on the
outer side. The latter is of significant taxonomical
value.
In D. gigantissimum Stefanescu has a developed
cingulum on the outer anterior and on the posterior
side of the crown, resembling a chain of large tuber-
cles, closely arranged along the end of the tooth.
There are no significant differences in the struc-
ture of the first molars of D. bavaricum H. v. Meyer,
except in their size.
The other molars (M
2
and M
3
) of D. thraceiensis
are not very different from those of the other species.
Mandible. (Pl. III, Fig. 1 and Pl. V).
Like the skull, it is entirely preserved. Compared to
the size of the body, it is not large (measurements in
Table 3). Ramus horizontalis long, high, wide and
laterally slightly flattened. Below P
3
there is a well
shaped double opening foramen mentale. The
highest point of the mandible is at the symphysis, it
becomes lower towards the posterior end (see Plate
V). Because of this, the last molar is not horizontal,
but inclined backwards. Processus angularis is strong-
ly developed, and fossa masseterica is wide but not
very deep. The most typical feature of the animals
belonging to this group is the shape of the symphys-
is. Together with the tusks, it is curved down and
back. In D. thraceiensis sp. n. the symphysis is solid,
16
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17
3 Geologica Balcanica, 3-4/2006
N
o
t
e
:

I
n

t
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e

c
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l
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m
n
s

a
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s
.
T
a
b
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e

2

B
U
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R
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D
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p
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1
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a
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8
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.

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.

E
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1
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,

P
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1

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-

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-


18
Fig. 6. Deinotherium levius Jourdan
Fig. 7. Deinotherium giganteum Kaup
3. The external nares (Figs. 5, 6, and 7 exn) in D.
thraceiensis sp. n. is large, deep and pear-shaped. It
starts from the very beginning of the nasal bones,
gradually widening and taking two thirds of the
length of the whole skull. In the other species it is
shorter and almost of the same width in its anterior
and posterior parts.
4. The nasal bones in D. thraceiensis sp. n. are short,
narrow, frontally rounded and fused. In the other
species they are longer, wider, almost flat for most of
their length, and separated. In the Gussiatin species
they protrude above the external nares. This is not
the case with the two other skulls (Figs. 5, 6, and 7).
5. The calvaria of the skull (fronto-parietal part, Figs.
5, 6, and 7 Ic in D. thraceiensis sp. n.) is high and
wide). In the other two species it is lower and shorter.
In the Gussiatin species it is even strongly concave
in its anterior part.
6. The eye socket (Figs. 5, 6, and 7 orb) in the
Ezerovo skull is large and separated from the ear
area. In D. giganteum Kaup they are almost fused,
and in D. levius Jourdan from Gussiatin the situa-
tion is closer to our case.
7. The zygomatic arch (Figs. 5, 6, and 7 zyg) of D.
thraceiensis sp. n. with its developed proc. postorbit-
alis resembles the one of D. levius Jourdan, but in its
posterior part, behind its connection with the tem-
poral bone, it lacks the processus that Svistun notes.
He regards this processus as related to the develop-
ment of the lower tusks. No comparison with the zy-
gomatic arch of the Eppelsheim skull is possible,
because it is not preserved.
19
8. There are significant differences between the three
skulls in the occipital region. Os occipitale in D. thra-
ceiensis sp. n. is high, wide and at an angle of 80
toward the forehead, and of ca. 70 to the plane of
the teeth. In D. giganteum Kaup these angles are 70
and 50 correspondingly; in D. levius Jourdan the
declination of the occipital bone is even larger, so
the angle is only 60. As a whole, the skull of D. levius
Jourdan is very flat and low. Os occipitale is visibly
concave and very wide, forming two lateral wings
(Fig. 6 C and D OS). Those wings are almost lack-
ing in D. thraceiensis sp. n. and in D. giganteum
Kaup they are much less developed.
9. The position of the occipital condyles is rather
different (Figs. 5, 6, and 7 oc). In D. thraceiensis
sp. n. they are situated in the middle and in the low-
er part of the occipital bone, almost the same is their
position in D. giganteum Kaup, and in D. levius Jour-
dan they are on the upper half of the noted bone.
According to Svistun, this permitted the animal to
raise its head almost at right angle to its neck, thus
fully using its back curved lower tusks.
10. Worth mentioning is a characteristic peculiarity
of the Gussiatin skull lacking in all the others,
including the skull from Ezerovo. This skull has a
double articulation with the mandible. Once by an
articular surface on the zygomatic process of the
temporal bone and then by a second surface on the
petromastoideum. Between those two surfaces is the
external meatus of the auditory canal. How and
when this second articular surface was used, is
unclear.
After comparing their anatomy we can see that
deinotheres could be divided in two groups. One in-
cludes D. giganteum Kaup, D. gigantissimum Stefa-
nescu, D. thraceiensis sp. n., D. levius Jourdan and
D. indicus Falconer, which have larger teeth, and the
second D. bavaricum H. v. Meyer, D. pentapota-
miae Lydekker and others, all with smaller teeth.
As a whole, the teeth are rather similar and differ-
ences are seen mainly in the two premolars P
3
and P
4
.
In D. giganteum Kaup P
3
is more rounded on the
inner side. The two inner ridges are more perpendic-
ular to the outer one. The anterior inner ridge is not
connected with the outer and is at an acuter angle to it
than in other taxa. They almost don!t touch each other
on the inner side. If anything like that is observed in
these species, than it is down at the very basis. The
ridges merge when they are worn. The cingulum is
more clearly seen here. It consists of numerous large
and small tubercles on all sides of the crown.
No comparison with the third premolars of D. gi-
gantissimum Stefanescu is possible, because they have
been restored after D. giganteum Kaup.
In D. levius Jourdan the anterior inner ridge of P
3
is situated diagonally to the outer one and touches
it. The posterior one is wider and V-shaped in its
upper part, with a shorter anterior part. It is also di-
agonal to the outer ridge, but does not touch it.
In D. indicum Falconer the inner ridges are some
more diagonally situated, compared to D. giganteum
Kaup, but their tips do not contact.
D. bavaricum H. v. Meyer has small P
3
s. Their
inner ridges are some more perpendicular to the outer
one. This is rather clear with the anterior inner ridge.
The valley between them is wider and deeper.
In D. pentapotamiae Lydekker, D. levius Jourdan
and D. indicum Falconer the differences are almost
the same, concerning the position of the two inner
ridges and the size of the tooth and its cingulum.
The fourth premolar of D. thraceiensis sp. n. has
inner ridges perpendicular to the outer one. The
posterior is narrower and longer. The triangular val-
ley surrounded by the posterior inner wall of the out-
er ridge and the inner wall of the posterior ridge is
clearer than in all the other species.
D. gigantissimum Stefanescu has a longer P
4
. Its
transverse ridges are at a larger distance from each
other. Besides, it has a strong cingulum on the ante-
rior and the posterior side, lacking in our species.
In D. bavaricum H. v. Meyer P
4
s are smaller and
of a more regular square shape. The transverse ridg-
es are more distant at the inner side of the crown,
and the valley surrounded by them and the outer
ridge is wider and larger.
In D. levius Jourdan the posterior inner ridge is
also attached to the outer one with its longer posteri-
or branch.
In D. indicum Falconer and D. pentapotamiae
Lydekker the last premolars have a square shape and
are smaller. Their transverse ridges are parallel and
of equal size. The valley between them is larger and
everywhere equally wide.
In the M
1
s of D. giganteum Kaup, unlike D. thra-
ceiensis sp. n., the three ridges are almost of equal
size, with an equally developed cingulum on both
sides of the valleys, blocking them entirely on the
outer side. The latter is of significant taxonomical
value.
In D. gigantissimum Stefanescu has a developed
cingulum on the outer anterior and on the posterior
side of the crown, resembling a chain of large tuber-
cles, closely arranged along the end of the tooth.
There are no significant differences in the struc-
ture of the first molars of D. bavaricum H. v. Meyer,
except in their size.
The other molars (M
2
and M
3
) of D. thraceiensis
are not very different from those of the other species.
Mandible. (Pl. III, Fig. 1 and Pl. V).
Like the skull, it is entirely preserved. Compared to
the size of the body, it is not large (measurements in
Table 3). Ramus horizontalis long, high, wide and
laterally slightly flattened. Below P
3
there is a well
shaped double opening foramen mentale. The
highest point of the mandible is at the symphysis, it
becomes lower towards the posterior end (see Plate
V). Because of this, the last molar is not horizontal,
but inclined backwards. Processus angularis is strong-
ly developed, and fossa masseterica is wide but not
very deep. The most typical feature of the animals
belonging to this group is the shape of the symphys-
is. Together with the tusks, it is curved down and
back. In D. thraceiensis sp. n. the symphysis is solid,
20
T
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5
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6
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7
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8
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9
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1
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1
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1
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1
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2
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2
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2
2
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21
T
a
b
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4
L
o
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c
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k

t
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s
p
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E
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B
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S
t
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,

1
9
0
9
,

R
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i
a


B
a
k
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o
v
,

N
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k
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,

1
9
6
2
,


B
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a
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a

r
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m
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r
,

B
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v
,

N
i
k
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,

1
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,

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u
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g
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r
i
a


M
e
a
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3

W
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W
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M
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W
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2

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W
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3

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22
T
a
b
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e

4

A
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B
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,

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,

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R
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M
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,

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1
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d
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d
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P
3

W
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L
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P
4

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M
1

W
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W
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24
PLATE VII
Deinotherium thraceiensis sp. n.
1. Part of vertebral column (vertebrae, ribs). Scale ca. 5%
Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE VIII
Deinotherium thraceiensis sp. n.
13. Atlas. Scale ca. 13.6%. Coll. SU.M. No. SU Pl 312/3-1
4, 5. Axis. Scale ca. 13.6%. Coll. SU.M. No. SU Pl 312/3-2
Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE IX
Deinotherium thraceiensis sp. n.
1. Third cervical vertebra. Coll. SU.M. No. SU Pl 312/3-3
2. Fourth cervical vertebra. Coll. SU.M. No. SU Pl 312/ 3-4
3. Fifth cervical vertebra. Coll. SU.M. No. SU Pl 312/3-5
4. Second thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-9
5. Sixth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-13
Scale of all figures ca. 11.9%; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE X
Deinotherium thraceiensis sp. n.
1. Ninth thoracic vertebra. Coll. SU.M. No. SU Pl 312/ 3-16
2. Tenth thoracic vertebra. Coll. SU.M. No. SU Pl 312/ 3-17
3. Eleventh thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-18
4. Twelfth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-19
5. Thirteenth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-20
6. Fourteenth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-21
Scale of all figures ca. 11.8%; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XI
Deinotherium thraceiensis sp. n.
13. Sacrum. Coll. SU.M. No. SU Pl 312/3-32
4. First lumbar vertebra. Coll. SU.M. No. SU Pl 312/3-22
Scale for all figures ca. 12.9%; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XII
Deinotherium thraceiensis sp. n.
1, 1a. Right scapula in medial and lateral view. Coll. SU. M. No. SU Pl 312/6
2, 2a. Left scapula in medial and lateral view. Coll. SU.M. No. SU Pl 312/5
Scale ca. 4.6%; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XIII
Deinotherium thraceiensis sp. n.
1, 1a. Right humerus in lateral and cranial view. Coll. SU.M. No. SU Pl 312/8
2. Left humerus in cranial view. Coll. SU.M. No. SU Pl 312/7
Scale ca. 5.4%; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XIV
Deinotherium thraceiensis sp. n.
1, 1a. Right ulna in lateral and medial view. Coll. SU.M. No. SU Pl 312/10
2, 2a. Right radius in lateral and medial view. Coll. SU.M. No. SU Pl 312/12
Scale ca. 6.2%; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XV
Deinotherium thraceiensis sp. n.
Carpals
1. Cuneiforme (ulnare) sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-1
2. Lunare sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-2
3. Magnum sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-7
4. Unciforme sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-8
5. Pisiforme sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-4
Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE VII
PLATE VIII PLATE IX
PLATE X PLATE XI
PLATE XII PLATE XIII
PLATE XIV PLATE XV
PLATE XVI PLATE XVII
PLATE XVIII
PLATE XX
PLATE XIX
PLATE XXI
PLATE XXII
PLATE XXIII
PLATE XXIV
25
PLATE XVI
Deinotherium thraceiensis sp. n.
1, 1a. Metacarpus I of the first digit in frontal and lateral view. Scale ca. 18.6%. Coll. SU.M. No. SU Pl 312/14-1
2, 2a. Metacarpus II of the second digit in frontal and lateral view. Scale ca. 18.6%. Coll. SU.M. No. SU Pl 312/14-2
3, 3a. Second phalanx of the second digit in frontal and lateral view. Scale ca. 14.1%. Coll. SU.M. No. SU Pl 312/15-2
Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XVII
Deinotherium thraceiensis sp. n.
1, 1a. Metacarpus V sin in frontal and lateral view. Scale ca. 22.2%. Coll. SU.M. No. SU Pl 312/14-5
2, 2a. Hoof of the third digit from above and from below. Scale ca. 65%. Coll. SU.M. No. SU Pl 312/15-10.
3, 3a. Second phalanx of the fourth digit in frontal and lateral view. Scale ca. 22.2%. Coll. SU.M. No. SU Pl 312/15-8
4. Patella. Scale ca. 22.2%. Coll. SU.M. No. SU Pl 312/18-2
Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XVIII
Deinotherium thraceiensis sp. n.
1. Left hand. Scale ca. 7.7%. Coll. SU.M. No. SU Pl 312/13
Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XIX
Deinotherium thraceiensis sp. n.
1. Pelvic bones with the sacrum. Scale ca. 4.0%. Coll. SU.M. No. SU Pl 312/16-1 and 2; SU Pl 312/3-32
Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XX
Deinotherium thraceiensis sp. n.
1. Pelvis in lateral view. Scale ca. 4.0%. Coll. SU.M. No. SU Pl 312/16-1
Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XXI
Deinotherium thraceiensis sp. n.
1, 1a. Right femur in dorsal and caudal view. Coll. SU.M. No. SU Pl 312/17-2 Figs. 2 and 2a. Left femur (restored) in caudal view.
coll. SU.M. No. SU Pl 312/17-1
Scale ca. 6.8%; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XXII
Deinotherium thraceiensis sp. n.
2, 2a. Left tibia (restored) in dorsal and caudal view. Coll. SU.M. No. SU Pl 312/19-1
3. Right tibia (restored) in dorsal view. Coll. SU.M. No SU Pl 312/19-2
Scale ca. 8.0%; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XXIII
Deinotherium thraceiensis sp. n.
1, 1a. Pes (restored). Scale ca. 9.0%. Coll. SU.M. No. SU Pl 312/14
2. Calcaneus sin. Scale ca 20.3%. Coll. SU.M. No SU Pl 312/21-1
Locality: Ezerovo, near Plovdiv; Level: Maeotian.
PLATE XXIV
Deinotherium thraceiensis sp. n.
Restored skeleton
Coll. SU.M. No SU Pl 312
Locality: Ezerovo, near Plovdiv; Level: Maeotian
4 Geologica Balcanica, 3-4/2006
26
wide and regularly curved. In its upper part there is
a wide deep groove, gradually becoming narrower
and shallower with the curving of the sympnysis.
Ramus ascendens wide and thick. All the area of
proc. angularis is thick at the base and strongly pro-
truding backwards. This thickness reaches as far as
proc. articularis. The anterior part of ramus ascen-
dens is significantly thinner, however, incl. proc. coro-
noideus itself. All the ramus ascendens in this part is
laterally slightly concave. Foramen mandibulae wide
and deep. Proc. coronoideus almost vertical in the
anteriror part. Tips of the processes curved back-
wards; at the posterior ends there is a moderate con-
cavity (incisure). Processus articularis thick and high.
Perpendicularly positioned, with well shaped artic-
ular surfaces. Two rami of the mandible not parallel.
At the level of M
1
a widening begins, rami coming
closer again at the level of M
3
, then going apart again.
Most distant from each other at the posterior ends.
The incisors (tusks) are a sequence of the sym-
physis, shaping the curve together with it. Their basis
starts deep inside the symphysis. There they are hollow,
their alveoli are of semi-circular shape (Fig. 8).
At the same time they are becoming thicker in-
side. At the end of the symphysis the tusks are al-
ready wholly solid. They gradually become thinner
and, curving in two directions ! outwards and back-
wards, go apart from each other. Their tips are point-
ed, slightly smoothed only at the foremost part of the
inner side, but there is no clear flat surface. Their
tips end just below the end of processus angulare.
Lower cheek-teeth. (Pl. VI). The toothrows also have
five teeth each ! two premolars and three molars.
Unlike the upper premolars, the lower are much
narrower, and their structure is very different,
especially P
3
.
P
3
has a long and narrow crown, pointed at the
anterior end. One large longitudinal ridge is situat-
ed along its axis, and another, transversal ! in the
posterior part. They form the occlusial surface of
the tooth. The longitudinal ridge is wider at its basis,
gradually narrowing towards the apex of the crown.
Its widest part is in the middle of the crown. There it
is more worn out and part of the dentine is seen. On
the inner part of the tooth, at the place where the
longitudinal ridge contacts the inner one, the first
narrows abruptly thus shaping a large triangular
valley. Outside the crown is smooth.
P
4
dext has a larger crown, elongated and wide.
The occlusal surface is moderately worn. It is formed
by two transversal ridges, contacting at the outer part
of the crown. At this place they divide the valley be-
tween them in two not quite equal parts. The anterior
wall of the first ridge is strongly concave. The poste-
rior part of the crown has a weak cingulum.
M
1
dext has three transversal ridges. First two are
more worn. Their structure is generally the same as
in M
1
. All three ridges are wider on the outer side.
The valleys dividing them become gradually narrower
towards the outer part. In this part the anterior walls
of the second and third ridge are slightly convex but
Fig. 8. Shape of the tusks of Deinotherium thraceisensis sp. n.
A ! shape of the tusks at the basis of the alveolus; B ! shape of
the tusks at 115.0 mm from their basis in the symphysis; C !
cross-section of I dext at 600.0 mm from the symphysis !
natural size
they don"t touch the ridge in front of them. On the
inner side, at the bottom of each valley there is a weak
tubercle, and on the anterior side ! a small cingulum.
Left M
1
resembles the right but is more worn.
M
2
dext is large and tetragonal. It has two thick
transversal ridges, wider on the outer side. There both
ridges form a small concavity. The valley between
them is deep and free but on the inner side there is a
small tubercle. On its anterior and posterior sides the
crown has a cingulum.
M
3
dext is large too but with an irregular tetrago-
nal shape. Built by two transversal ridges. The first is
wider. Both are widening outwards. At the ends !
inner and outer ! they are slightly curved to the front,
forming with their anterior walls shallow valleys. The
valley between them is deep and unblocked. The
posterior talon on the inner side of the crown is formed
by numerous tubercles of different size.
Comparison. The structure of the described mandi-
ble is close to the other species (Fig. 9).
The symphysis is moderately large. There are dif-
ferences in the shape of ramus horizontalis, proc.
articularis, proc. coronoideus and the tusks.
1. The symphysis of D. bavaricum H. v. Meyer is more
protruding. In D. levius Jourdan it is more rounded
and curved inwards and in D. giganteum Kaup, as
well as in D. gigantissimum Stefanescu and D. thra-
ceiensis sp. n. the curving is even stronger and more
gradual.
27
2. In D. thraceiensis sp. n. ramus horizontalis (in the
region of proc. angualris) strongly protrudes back-
wards with a pronounced curve. It almost reaches
the level of incisura mandibulae.
3. Ramus ascendens is almost perpendicular to ra-
mus horizontalis in its anterior part.
4. Processus articularis is high and abruptly separat-
ed from ramus ascendens, forming with it an obtuse
angle in its posterior part. Anteriorly it descends
steeply toward incisura mandibulae.
5. Processus coronoideus is high too and anteriorly
almost vertical, nearly at the same level as proc. ar-
ticularis.
6. Incisura mandibulae is deep and almost horizon-
tal in relation to the mandible, perpendicular to ra-
mus ascendens.
In D. gigantissimum Stefanescu the curve at proc.
angularis is weaker. Poc. articularis is not so abrupt-
ly separated from it. Proc. coronoideus posterioly
turns into incisura mandibulae. The latter is very
shallow. The symphysis is some more elongated, but
it has been restored. No P
3
is known for this species.
7. The third premolar ! P
3
in D. thraceiensis sp. n. is
built mainly by one longitudinal ridge and one weak
transversal ridge on the posterior inner side. The lon-
gitudinal crest touches the inner one. No cingulum.
In D. giganteum Kaup there is a bending tubercle
on the anterior inner side of P
3
which surrounds a
deep valley together with the longitudinal ridge. A
high cingulum encircles the crown.
In D. bavaricum H. v. Meyer P
3
is smaller and
built by two smaller transversal ridges and a large
longitudinal one, positioned on the outer side of the
crown.
8. P
4
in D. thraceiensis sp. n. has two transversal ridg-
es connected on the outer side. Cingulum is weak
and only on the posterior side.
In D. giganteum Kaup the structure of the tooth is
the same, but with well developed cingulum on the
inner side of the crown.
In D. gigantissimum Stefanescu P
4
has the same
structure as D. giganteum Kaup but without the large
cingulum.
P
4
in D. bavaricum H. v. Meyer has two ridges.
The posterior is larger, with two parts at a right angle
to each other. It is elongated and touches the poste-
rior wall of the anterior ridge. The latter is #C-
shaped$. It surrounds a tetragonal valley anteriorly.
Strong cingulum.
As a whole, the fourth premolars in the studied
taxa are rather close in their structure.
9. M
1
, M
2
and M
3
in the species described show no
significant differences when compared to the other
studied species. A small exception is the fact that
our species lacks a developed cingulum. The dis-
placement of the third ridge in M
3
is not regarded as
a distinct taxonomic peculiarity by us.
We had no opportunity to compare the material
to the species D. cuvieri Kaup, D. levius Jourdan, D.
indicum Falconer, D. pentapotamiae Lydekker.
Vertebral column (Pl. VII).
In D. thraceiensis sp. n. it consists of 7 cervical, prob-
ably 14 thoracic, probably 14 lumbar, 3 sacral and
probably 6 caudal vertebrae.
Atlas (Pl. VIII, Figs. 1, 2 and 3).
Almost entirely preserved. Processes are only partial-
ly broken, without losing the shape of the vertebra. It
consists of two arches ! dorsal and ventral, limiting
a large foramen vertebrae. The foramen widens down-
wards and is slightly constricted in the middle. There"s
no corpus vertebrae. In fact the atlas is a wide bone
ring. The transverse processes have become large bone
plates ! wings of the atlas. There is no spinous pro-
cess. The atlas articulates with the skull by wide oval
concavities, cranially positioned on the arches. Cau-
dally, on the ventral arch, there is an articular sur-
face for the dens of the axis. Foramen ovale and for.
ventrale lateralis on both sides of the vertebra and
large (measurements in Table 5).
Axis (Pl. VIII, Fig. 4 and 5). Body narrow and oval,
spinous process high, wide and long. Two strong
Fig. 9. Comparison of deinothere mandibles: a ! Deinotheri-
um bavaricum H. v. Meyer; b ! Deinotherium levius Jourdan;
c ! Deinotherium giganteum Kaup; d ! Deinotherium thra-
ciensis sp. n.
28
wards the spinous process. All other thoracic verte-
brae resemble the second. There are differences only
in the size of proc. transversus and proc. spinosus,
which are smaller in each next vertebra. From the
first to the twelfth the size of the corpus and foram-
ina decreases, then increases to the last. In the fif-
teenth and all the following there is no fovea costal-
is. So, there were no ribs attached to these vertebrae
(measurements in Table 6).
Lumbar vertebrae (Pl. XI, Fig. 4)
Resemble the last thoracic vertebrae. Here, too, fovea
costalis is missing. The diameter and thickness of
corpus vertebrae gradually decreases from the first
to the last. Spinous processes become shorter. As in
the thoracic vertebrae, they are inclined backwards.
In the other vertebrates these processes on the lum-
bars are pointing anteriorly. This peculiarity shouldn"t
be of taxonomic importance for genus Deinotherium
because it is present in the other proboscideans as
well. The transverse processes are not long as should
be expected for lumbars so we cannot speak of proc.
costarius. These peculiarities in the development of
the processes of the lumbars and the lack of fovea
costalis in the last thoracic vertebrae make it diffi-
cult for us to decide unequivocally where one group
of vertebrae ends and the other begins. This is why
we cannot say for sure how many the thoracic and
correspondingly the limbar vertebrae are. The diam-
eters of foramen vertebrae increase as in the last tho-
racic vertebrae. They are largest in the last vertebra
(Table 7). This character too creates difficulties in
the determination of the two groups of vertebrae but
crests on tip with a deep furrow between them. Dens
epistrophei connects the axis with the atlas. Fora-
men vertebrae much smaller than in atlas, tetragonal
(measurements in Table 5).
Third, fourth and fifth cervicals (Pl. IX, Figs. 1, 2
and 3). All have similar structure. Narrow bodies.
Cranially slightly convex, caudally slightly concave.
Transverse processes small and rounded in the ends.
Transverse foramina limited by the bodies and the
transverse processes. Well pronounced articular sur-
faces on the cranial and caudal processes. Foramen
vertebrae wide and high. Spinous processes not pro-
nounced.
Sixth and seventh cervicals are built in the same
way as the previous three, but they have spinous pro-
cesses, largest on the seventh. Below the transverse
processes the seventh cervical has a well shaped fovea
costalis ! a concavity for the head of the first rib
which is between the seventh and the eighth vertebra
(measurements of all cervicals in Table 5).
Thoracic vertebrae (Pl. IX, Fig. 4 and 5, Pl. X, Fig. 1,
2, 3, 4, 5, and 6).
The first thoracic vertebra is very similar to the last
cervical, only its spinous process is larger. Thus it
resembles the second thoracic vertebra, where this
process is most developed. The second thoracic ver-
tebra (Pl. IX, Fig. 4) has a distinct structure. Its body
is small, cranially convex, caudally ! slightly round-
ed. Its cranial articular processes are less develped
than the caudal. Fovea costalis cranialis larger than
f. c. caudalis. Spinous process strong, widening at
the tip, tetragonal. Foramen vertebrae elongated to-
Table 5
Cervicals
Table 6
Thoracics
THORACICS
Dimensions
1 2 3 4 5 6 7 8 9 10 11 12 13 14
1. Diameter of corpus
vertebrae
250.0 240.0 230.0 220.0 210.0 200.0 196.0 192.0 188.0 186.0 182.0 180.5 180.0 179.0
2. Thickness of corpus
vertebrae
102.0 96.0 92.0 89.0 85.0 86.0 87.0 88.0 89.0 90.0 90.5 91.0 92.0
3. Length at proc. spinosus 400.0 396.0 390.0 384.0 380.0 362.0 340.0 320.0 310.0 295.0 290.0 287.0 280.0
4. Length of proc. spinosus 205.0 610.0 580.0 550.0 532.0 510.0 500.0 460.0 400.0 340.0 290.0 289.0 288.0 285.0
5. Length of proc.
transversus
93.0 102.0 112.0 117.0 120.0 116.0 112.0 110.0 106.0 100.0 96.0 94.0 87.0
6. Length of foramen
vertebrae
110.0 97.0 92.0 88.0 83.0 80.0 70.0 62.0 54.0 46.0 40.0 43.5 47.0 58.0
7. Width of foramen
vertebrae
78.0 70.0 62.0 50.0 41.0 30.0 38.0 45.0 52.0 57.0 60.0 60.0 60.5 59.0

CERVICSLES
Dimensions, mm
Atlas Axis 3 4 5 6 7
1. Height of the vertebra 300.0 340.0 330.0 340.0 343.0 350.0 352.0
2. Width of the vertebra 400.0 320.0 310.0 315.0 218.0 224.0 226.0
3. Length of corpus vertebrae 212.0 230.0 238.0 240.0 246.0 250.0
4. Width of corpus vertebrae 170.0 195.0 202.0 206.0 208.0 210.0
5. Length of articular surfaces 100.0 103.0 105.0 106.0 108.0
6. Height of processus spinosus 60.0 150.0
7. Length of foramen vertebrae 200.5 165.0 60.0 80.0
8. Width of foramen vertebrae 145.0 93.0

29
it probably has its functional meaning. It should be
related to the use of the hindlimbs. The accumula-
tion of nerve tissue close to the pelvis probably facil-
itated the movements of these huge #columns$ ! the
hindlimbs.
Sacral vertebrae (Pl. XI, figs. 1, 2 and 3). Three, fused,
building a single bone ! the sacrum. Body of the
sacrum large, with wide, slightly rounded wings. Two
foramina sacralia dorsalia on each of the two sides
of the wings. Spinous processes of the first two
vertebrae smaller and narrower, on third ! highest
and widening in its upper part. Two foramina on
each side ventrally, by the attachment of the
vertebrae. Of these four ventrally situated foramina,
the first two are wider, penetrating inside to the
sacral canal. Body of the sacrum oval, anteriorly
slightly concave and protruding in front of the
wings.
Caudal vertebrae. Preserved are only six original cau-
dals, but on the mounted skeleton there are 21. The
first, those after the sacrum, resemble the last lum-
bars, but their processes are gradually decreasing in
size. In the first, a chanal is still present, which grad-
ually disappears. The body of the caudals is elon-
gated, narrow and convex on both sides.
Comparison. Many of the vertebrae in the skeletons
we compare our materials with are reconstructed
which makes the comparison difficult. Nevertheless,
one can see that in our specimen the spinous pro-
cesses of the cervical vertebrae are weaker. These pro-
cesses are strong and high in all known skeletons. A
difference in D. thraceiensis sp. n. is the large tetrag-
onal widening of the tip of this process on the sec-
ond thoracic. There are no other visible differences.
The number of the vertebrae (mainly of the thoracics
and the caudals) varies. In fact their number is un-
known. As far as we know, there are no finds of a
whole tail.
Ribs (costae). Fragments of seven pairs of ribs were
found. As far as the fifteenth vertebra lacks fovea
costalis, the ribs must have been 14. First rib is wide
and thick. One half of caput costae articulating with
the last cervical is oval, the other half ! smaller and
spherical. Caput costae of the second vertebra has
the same structure but its neck is narrow and short.
Its body is wide and flat. The third rib is narrower,
Table 8
Sacrum
Table 9
Caudal vertebrae vert. coccigiae
Dimensions mm
1. Total length 315.0
2. Width at the wings 405.0
3. Anterior width of the corpus (at first fused vertebra) 180.0
4. Posterior width of the corpus (at third fused vertebra) 140.0
5. Anterior height of the corpus (at first fused vertebra) 118.0
6. Posterior height of the corpus (at third fused vertebra) 50.0
7. Height of processus spinosus 155.0
8. Anterior width of canalis sacralis (at first fused vertebra) 162.0
9. Posterior height of canalis sacralis (at third fused vertebra) 64.0

Dimensions, mm 1 2 3 5 7 8
1. Height of the vertebra 180 175 155 150 130 130
2. Width of the vertebra 290 240 180 190 190 190
3. Diameter of corpus vertebrae 110 109 109 100 100 100
4. Thickness of corpus vertebrae 106 104 94 85 90 90
5. Height of processus spinosus 100 90 - - - -
6. Vertical diameter of foram. vertebrae 60 60 60 40 - -
7. Horizontal diameter of foram. vertebrae 100 100 60 50 - -

Table 7
Lumbars
Dimensions 1 2 3 4 5 6 7 8 9 10
1. Diameter of corpus vertebrae 178.0 177.0 176.0 174.0 172.0 170.0 168.0 165.0 162.0 160.0
2. Thickness of corpus vertebrae 93.7 95.0 97.0 97.5 98.0 100.0 100.0 98.0 96.0 94.0
3. Length at proc. transversus 274.0 269.0 264.0 258.0 250.0 240.0 240.0 242.0 243.0 245.0
4. Length of proc. spinosus 282.7 280.5 277.8 275.5 273.0 270.0 268.0 266.5 262.0 258.0
5. Length of proc. transversus 82.0 77.0 73.0 69.8 66.5 62.0 63.0 63.0 64.5 66.0
6. Length of foramen vertebrae 64.0 75.0 81.0 90.0 102.0 110.0 120.0 130.0 140.0 150.0
7. Width of foramen vertebrae 58.5 60.0 61.0 59.5 60.0 60.0 61.0 61.5 62.0 63.0

30
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31
also with a short and narrow neck as the first two
ribs. The body of the fourth is long and almost round.
Caput costae spherical, divided by a well pronounced
furrow in the middle into two surfaces facies ar-
ticularis and facies articularis condilis, the first of
which is bigger. Near its distal end the rib gradually
becomes thinner. At the very end it becomes wider
ending with a rough surface for the attachment of
the cartilage. The other ribs are similar to the fourth.
Their length gradually decreases (measurements in
Table 10).
Sternum. Only a fragment preserved, with a rough
surface and places for the attachment of the rib car-
tilages.
Comparison. The most disputable problem concern-
ing the ribs is their number. There is still no explicit
opinion on that matter and we don!t know their pre-
cise number. D. gigantissimum Stefanescu has 32 pairs
of ribs restored, reaching up to the sacrum. What
was the reason for this number, we do not know. By
the restoration of D. bavaricum H. v. Meyer 11 pairs
of ribs were mounted. As we can see from Plate XXIV,
they had 5th, 9th, 17th and 18th thoracic vertebrae.
It seems they restricted themselves to the 12th.
Scapula (Pl. XII, Figs. 1 and 1a, 2 and 2a). Only the
right is entirely preserved. There are separate frag-
ments from the left, and it was restored after the right.
Dorsally strongly flattened, becoming thinner at the
anterior and posterior edge. Spina scapulae and the
edges surround two concavities of unequal size. The
crest takes almost the entire length of the scapula. It
is divided in two near the tubersitas spinae, forming
a wide long groove starting immediately behind the
acromion and ending in the dorsal thickening. The
acromion itself is high and curved towards the neck
of the scapula. Fossa infraspina shallower but wider
than fossa supraspina. Margo thoracales long and
pointing outwards. It has a regular triangular shape.
The articular surface for the attachment to the hu-
merus is shallow.
Humerus (Pl. XIII, Figs. 1. 1a and 2). Both humeri
entirely preserved. Caput humeri large and curved,
with no distinct neck. Tuberculum majus protrudes
upwards, above caput humeri. Tuberculum minus is
much smaller. Positioned cranio-medially. Between
the two tubercles there is a deep and wide furrow
sulcus intertubercularis. Shaft of the bone round
and smooth with weak sulcus spiralis. It starts on the
dorsal surface above the caput and ends in front on
the distal part over the joint. In the distal part of the
bone is trochlea humeri, with two large projections,
the lateral being the higher. In front, above them, is
the wide and shallow fossa radialis, and on the pos-
terior side fossa olecrani. It is deeper and wider.
Table 12
Humerus
D. thraceiensis
sp.n. Ezerovo,
Bulgaria
Dimensions, mm
sin dext
D. gigantissimum
St. Manzati,
Romania
D. giganteum
Kaup Saint
Yean Lartet
D. giganteum
Kaup M!nchen
Stromer
D. giganteum Kaup
M!nchen Stromer
495
1. Total length 1142.0 1146.0 1100.0 740.0 1090.0 900.0
2. Width of caput humeri 295.0 292.0 170.0 120.0 - -
3. Width in proximal part 325.0 320.0 370.0 280.0 - -
4. Width in diaphysis 190.0 195.0 - 124.0 115.0 90.0
5. Width in distal part 355.0 350.0 310.0 240.0 - -
6. Width at epicondyles 290.0 296.0 300.0 316.0 290.0 200.0
7. Diameter of caput
humeri
200.0 200.0 - - 260.0 190.0
8. Diameter of trochanter
major
202.0 207.0 260.0 108.0 260.0 210.0
9. Diameter of trochanter
minor
185.0 190.0 140.0

Table 13
Ulna
D. thraceiensic
sp.n. Ezerovo,
Bulgaria
Dimensions, mm
sin dext
D. gigantissimum
St. Manzati,
Romania
Stefanescu
D. giganteum
Kaup Saint Yean
Deperet, 1892
D. giganteum
Kaup M!nchen
495 Stromer,
1938
D. giganteum
Kaup M!nchen
Stromer, 1938
1. Total length 1150.0 1130.0 1050.0 - 850.0 1020.0
2. Width at proximal end proc. olecrani 320.0 300.0 300.0 320.0 - -
3. Height from proc. olecrani to proc.
styloideus
150.0 140.0 - - - -
4. Height of inc. semilunaris 127.0 120.0 - 150.0 - -
5. Width at inc. semilunaris prox. end 96.0 90.0 - 50.0 120.0 115.0
6. Width at inc. semilunaris dist. end 265.0 60.0 - - 260.0 225.0
7. Width in diaphysis 88.0 86.0 - 130.0 115.0 90.0
8. Width at distal end 255.0 50.0 -

32
The lateral concavity for the sinews is larger than
the medial.
Ulna (Pl. XIV, Fig. 1 and 1a). Both bones are entirely
preserved. They are wholly identical. Tuber olecrani
large and curved medially. Processus anconeus slight-
ly twisted and situated posteriorly, incisura semilu-
naris shallow and wide. Lateral coronoid processes
strongly protruding sideways. Their upper surfaces
form the whole semilunar incisure. Between them is
incisura radialis, with coarse surface.
According to V. M. Svistun, such a structure of
the elbow joint permitted different movements when
the forelimb was in flexion.
The diaphysis is flat, the bone ending distally with
well shaped processus styloideus ulnae and cicum-
ferentia articularis.
Radius (Pl. XIV, Fig. 2 and 2a). This bone is much
smaller than the ulna. Slightly widening in its prox-
imal part, with well pronounced articular surface.
Slightly rounded in the diaphysis. Ends with a clear-
ly outlined articular surface. With this part it articu-
lates with the ulna. Slightly convex laterally, in the
distal end.
Comparison. There are no significant differences
between the taxa of genus Deinotherium concerning
the structure of the scapula, humerus, ulna and ra-
dius. According to Svistun, the ulna of the skeleton
he describes has a triangular shape of the diaphysis
and is fused with the radius at ca. 1/3 of its length.
There is no such thing in D. thraceiensis sp. n.
Carpus (Pl. XV, Figs. 16; Fig. 10 A, B).
Made up of nine bones, arranged in two rows prox-
imal and distal. Five bones in the proximal: scaphoi-
deum, lunare, cuneiforme, pisiforme and trapezium;
four in the distal: trapezoideum, magnum, unciforme
and centrale. Trapezium and trapezoideum are miss-
ing and have been reconstructed. More bones are
missing from the right wrist, and have been restored
after the left. Here we!ll describe only the left bones
(measurements in Table 15).
Table 14
Radius
D.thraceiensis sp.n.
Ezerovo, Bulgaria Dimensions, mm
sin dext
D.gigantissimum
St. Manzati,
Romania Stefanescu
D.giganteum Kaup Pikermi
Gaudry, 1862
1. Total length 928.0 925.0 680.0 920.0
2. Width at proximal end 144.0 142.0 130.0 150.0
3. Width in diaphysis 95.0 95.0 40.0 -
4. Width at distal end 210.0 214.0 - 210.0
5.
Length of the articular
surfaces for inc. semilunaris
160.0 160.0 90.0 -
6. Length of same surface 130.0 132.0 - -

Fig. 10. Manus of Deinotherium thraciensis sp. n. A left manus; B right manus
33
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5 Geologica Balcanica, 3-4/2006
34
Scaphoideum (radiale) (Fig. 10 A, B No.1). Long,
narrow and almost flat with rough lateral surface.
Two large facets on the medial surface. Proximal
contacts lunare, distal centrale. A facet on the
very proximal end, articulating with the distal end of
the radius. The lower end of the scaphoid articu-
lates with the fifth metacarpal.
Lunare (Fig. 10 A, B No. 2). Triangular. Anterior
surace rounded. Two concave facets in the proximal
part, articulating with the distal parts of the ulna
and radius. Lateral larger. Two slightly concave fac-
ets in the distal part articulate with magnum and
centrale. The lateral facet of the scaphoid and the
medial of the cuneiform are divided in the middle
by a crest for the attachment of sinews.
Cuneiforme (ulnare) (Fig. 10 A, B No. 3). The bone
has a peculiar form. It is flat. Medially, a large pro-
cessus turns down to the proximal part of McI. On
the inner side of this processus is os unciforme, which
is part of the distal row. In the proximal posterior
part there is a small facet for the pisiform. The ante-
rior and the medial surfaces of the bone are rough.
Pisiforme (Fig. 10 A, B No. 4). Smallest bone of the
wrist. Its anterior part contacts the ulna and the cu-
neiform by two pronounced facets. Rounded in the
other parts. Slightly curving downward at the very end.
Magnum (Fig. 10 A, B No. 7). Almost square, elon-
gated anteriorly. Situated in the middle of the distal
row, between os unciforme and os centrale. Two fac-
ets on the distal part. Medial larger, touching the
proximal part of McIII, the other touches a small
part of McIV. Surface rough in the anterior part,
and the facets for articulation with os unciforme and
os centrale are concave in the middle this is the
place where sinews are attached.
Unciforme (Fig. 10 A, B No. 8). Largest of the distal
row. Tetragonal. Slightly rounded anteriorly, elon-
gated posteriorly. The proximal acet for the cunei-
form is large and medially inclined. Two facets in
distal end, medial larger and contacting McII, later-
al McIII.
Centrale (Fig. 10 A, B No. 9). Although slightly elon-
gated in the ends, it also has a square shape. Situat-
ed between the magnum and the scaphoid. Distal
surface slightly concave, entirely lying on the proxi-
mal articular surface of McIV. Facet for articulation
with the magnum concave in the middle with a place
for the attachment of sinews.
Metacarpus (Pl. XVII; Fig. 10 A, B; No. I, II, III, IV
and V). Here, as with the wrist, more bones are pre-
served from the left forelimb.
First metacarpal McI. Long, wide and flat. Facet for
articulation with the cuneiform and the unciform
narrow and long. Slightly convex in the proximal end,
slightly concave in the middle. Large rounded artic-
ular surface on distal end. Strong sinew processes in
posterior and anterior parts.
Second metacarpal McII. Also long, but narrow.
Rounded in the anteriror part, strongly elongated pos-
teriorly, gradually becoming narrower. Facet for unci-
form in the proximal end long and wide. Laterally to it
a facet for McIII, smaller than the first. Slightly con-
cave diaphysis, distal end widening with a large artic-
ular surface for the first phalanx of the second digit.
Sinew processes strong and on both sides of the bone.
McIII and Mc IV resemble McII. Facets in proximal
parts for magnum, unciform and central.
McV is a mirror image of McI. Facet for the scaphoid
in anterior part.
Phalanxes (Pl. XVII and XVIII; Fig. 10 A and B No.
1, 2 and 3).
First phalanx of the first digit PH-I,1 is medium-sized,
with arounded, elongated flat shaft. Deep articular
surface on proximal end for the distal surface of McI.
Phalanx rounded in posterior part. Narrowing dia-
physis, distal end thicker and wider. Articular facet
for the next phalanx rounded and convex. Above and
parallel to it the sinew concavity, weakly pronounced.
First phalanx of second digit PH-II,1 resembles McII.
PH-III,1 and PH-IV,1 do not differ from PH-II,1, but
PH-V,1 is a mirror image of PH-I,1.
Second phalanx of the first digit PH-I,2 is smaller
than the other phalanxes. In its posterior part, by the
sinew crests, it is more convex, and concave in the
middle. Proximal articular surface concave, distal
rounded and also concave in the middle.
PH-II,2 is also smaller than the other phalanxes.
It looks like a second phalanx of a horse, i.e. short
and wide. Very thick proximal end, thinner in the
middle, thick again distally at the articular surface.
PH-III,2 and PH-IV,2 look like PH-II,2. PH-V,2 is
a mirror image of PH-I,2.
PH-I,3, PH-II,3, PH-III,3, PH-IV,3 and PH-V,3,
third phalanxes of all digits have almost the same
structure. Only their size is different, so we!ll describe
just one phalanx.
D. thraceiensis sp. n. has a long and large hoof.
In its proximal part it is slightly convex, in the mid-
dle and in the distal part concave. Large articular
surface for the second phalanx on the posterior part.
Slightly rounded in front. Measurements of the dig-
its are given in Table 17.
Height of the entire manus (wrist and digits) is
460.0 mm; diameter with spread digits 902.0 mm.
Comparison
All carpal and metacarpal bones, as well as the pha-
lanxes are similar in D. giganteum Kaup, D. gigantis-
simum Stefanescu and D. bavaricum H. v. Meyer.
No taxonomically significant differences were found.
Maybe only the hoofs show some. It was already said
that in D. thraceiensis sp. nov. they are long. Their
walking surface is flat and concave in the middle
which means they were touching the ground with all
their surface. In the other species the hoofs seem to
touch the ground in most cases only with their ante-
rior part.
Pelvis (Pl. XIX, Fig. 1; Pl. XX, Fig. 1). Both hipbones
are entirely preserved.
The wing of the hipbone is a wide S-shaped curved
plate, anteriorly rounded, with a pronounced pubic
35
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36
D. thraceiensis sp.n.
Ezerovo, Bulgaria Dimensions, mm
sin dext
D. gigantissimum St.
Manzati, Romania
Stefanescu
D. giganteum Kaup
M!nchen
Stromer, 1938
D. giganteum Kaup
M!nchen
Stromer, 1938
1. Total length 1650.0 1650.0 1440.0 188.0 1060.0
2. Diameter of caput femuri 195.0 195.0 - 210.0 140.0
3. Height of caput femuri from [???] 160.0 160.0 -
4. Width at proximal end 375.0 375.0 350.0 400.0 295.0
5. Width in diaphysis 225.0 225.0 -
6. Width at distal end 330.0 340.0 440.0 315.0 175.0
7. Thickness at proximal end 150.0 150.0

Table 19
Femur
Table 21
Fibula
D. thraceiensis sp.n.
Ezerovo, Bulgaria Dimensions, mm
sin dext
D. gigantissimum St.
Manzati, Romania
Stefanescu
D. giganteum Kaup
Pikermi Gaudry, 1862
1. Total length 1125.0 565.0 830.0 870.0
2. Width at proximal end 90.0 72.0 70.0 70.0
3. Width in diaphysis 126.0 103.0 - 100.0
4. Width at distal end 275.0 250.0 170.0 140.0
5.
Length of the articular surface for os
calcaneus
80.0 72.0 - 70.0
6. Length of the small facet 76.0 60.0 - 55.0

Table 20
Tibia
D. thraceiensis sp.n. Ezerovo,
Bulgaria Dimensions, mm
sin dext
D. gigantissimum St.
Manzati, Romania
Stefanescu
D. giganteum Kaup
Pikermi Gaudry, 1862
1. Total length 1200.0 1250.0 920.0 950.0
2. Width at proximal end 370.0 365.0 330.0 310.0
3. Thickness at proximal end 320.0 320.0
4. Width in diaphysis 160.0 165.0 260.0 140.0
5. Width at distal end 300.0 330.0 260.0 340.0

crest. The wing significantly narrows posteriorly.
Dorso-lateral surface smooth and slightly concave,
medial almost flat. In the region of facies articularis
the surface is rough. Acetabulum well shaped. Fora-
men obturatorium oval.
Femur (Pl. XXI, Fig. 1 and 1a). We found only the
right thighbone. Left was restored after it. The bone
is long and solid. Caput femuri large, almost paral-
lel to the bones axis. A small neck separates it from
trochanter major, which is laterally situated. Fossa
trochanterica strongly pronounced. Shaft flat, slightly
curved medially and laterally. Two condyles in the
distal part of the bone. Condylus medialis larger and
some higher than condylus lateralis. Fossa inter-
condylaris which separates them is wide and deep.
Fossa muscularis cranialis and F. m. caudalis deep
and parallel to the condyles. Longitudinal crests sur-
rounding trochlea patellaris high and rounded in
their uppermost part.
Patella (Pl. XVIII, Fig. 4). Pear shaped. Concave
anteriorly. In the middle it is convex like a hemi-
sphere surrounded by a deep and wide ring. Flat-
tened posteriorly. The larger part of the bone was
covered by a cartilage with the shape of the thigh-
bones surface it articulated with.
From the lower hindlimbs, only distal part of the left
tibia and fragments of the proximal parts of the two
fibulae were found.
Tibia (Pl. XXII, Fig. 2 and 2a). Although the bone
was restored to its full size, well discuss only its dis-
tal part which is original. Down of the diaphysis the
bone becomes triangular. It gradually widens, form-
ing at the distal end the cochlea tibiae. Maleolus
medialis strongly developed, incisura fibularis deep
and wide.
Fibula. The original fragments used for the restora-
tion are from the proximal part. This is a long and
narrow bone. Almost flat in its proximal part. Thick
at the beginning, articular surfaces unclear.
Comparison. There are no differences of taxonomi-
cal value in the hipbones and the bones of the lower
leg.
37
Tarsus (Pl. XXIII, Fig. 1 and 1a; Fig. 11 A and B)
Consists of seven bones. Two in the proximal row:
calcaneus and astragalus; five in the distal: cuboid,
three cuneiforms and, between them, navicular. Only
the calcaneus was found. The rest were restored af-
ter D. gigantissimum Stefanescu. The toes were re-
stored, too.
Comparison. There are no differences in the struc-
ture of the tarsals in the various deinothere species.
There are such however in their digits. In D. thra-
ceiensis sp. n. the size of the first and second pha-
lanxes gradually decreases. In D. gigantissimum Ste-
fanescu all digits have large first phalanxes, the sec-
ond are much smaller and the third ! even more.
There is a big difference in their size. D. bavaricum
H. v. Meyer and D. giganteum demonstrate a much
more gradual transition from the first to the last pha-
Table 22
Tarsals tarsus sin et dext
Deinotherium traceiensis sp.n.
Astragalus Calcaneus Naviculare
Cuneiforme
mediale
Cuneiforme
intermedium
Cuneiforme
laterale
Dimensions, mm
sin dext sin dext sin dext sin dext sin dext sin dext
1. Diameter of the corpus 313.0 310.0
2. Proximal height 90.0 90.0
3.
Average thickness of the
corpus
130.0 130.0 190.0 200.0
4. Proximal width 150.0 150.0
5. Total width 245.0 250.0
6. Anterior length of the bone 170.0 170.0 222.0 220.0 114.0 113.0 118.0 117.0
7. Posterior length of the bone 180.0 180.0
8. Total length of the bone 350.0 350.0
9. Diameter of the bone 335.0 337.0
10. Thickness of the bone 78.0 80.0
11. Height of the bone 81.0 80.0 79.0 78.0 82.0 80.0

Fig. 11. Pes of Deinotherium thraciensis sp. n. A ! left pes; B ! right pes
lanx. In D. thraceiensis sp. n. the last phalanx - the
hoof ! in the hindlimbs is large, as with the fore-
limbs. In the compared species they are much small-
er. They are anteriorly situated like nails and touch
the ground only with the anterior surface, or with a
small part of the lower surface.
It can be seen from the description that D. thraceiensis
sp. n. has numerous characters distinguishing it from
all the other species. Here are some of them:
1. Skull ! short, high and, compared to the size of
the body ! very small.
2. External nares large and deep, with a peculiar
shape.
3. Nasal bones short, narrow and high, frontally fused.
4. Forehead high and wide.
5. Occipital bones high and wide, at an angle of 80
to the forehead.
38
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39
6. Eye socket differentiated and large.
7. Occipital condyles low.
8. P
3
and P
4
differently structured.
9. Ramus horizontalis strongly protruding backwards
at angulare.
10. Processus articularis and p. coronoideus high.
11. Incisura mandibulae deep.
12. Tusks differently structured.
13. P
3
differently structured.
14. Curve of the symphysis much more regular.
15. Spinous processes on the cervicals shorter and
weaker; first cervicals lack processes.
16. Spinal canal widening at the lumbar region.
17. Spinous process on second thoracic vertebra
strongly developed.
18. 14 ribs.
19. Manus and pes high and spread.
20. Hoofs larger and entirely lying! on the ground.
On the basis of these characters, we believe that
the specific status of D. thraceiensis n. sp. is entirely
proved. To this conclusion we shall add some short
notes on its anatomy.
We are acquainted with the reconstructions by
Prof. Abel of D. bavaricum H. v. Meyer, based on the
skeleton in Vienna, and another of D. giganteum
Kaup. We know also the paintings by Augusta and
Burian of D. giganteum Kaup, and the painting of
D. gigantissimum Stefanescu exhibited in the Bucha-
rest Museum in front of the skeleton itself. It is made
following the first two authors. Everywhere the skull
with the mandible takes about 1/3 of the whole skel-
eton. The neck is thick, merging with the vertebral
column because, according to them, the last cervi-
cals have high spinous processes. Frontally, the chest
is narrow, and the body " solid, with high legs. The
latter are narrower near the fingers because of the
small hoofs.
In D. thraceiensis sp. n. the skull and the mandi-
ble are much less than 1/3 of the whole skeleton. The
neck is short and only the last cervicals have spinous
processes. They are very high on the first thoracic
vertebrae, widening at the tip. This permits us to sup-
pose it had a high hump like Elephas.
On the other hand, as far as the first rib is wide
and solid, its chest was probably broad. The digits
were more spread.
For the time being, we cannot comment on the
problems of the phylogeny of genus Deinotherium.
They could become the subject of a future work.
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