Deinotherium thraceiensis sp. nov. from the Miocene near Ezerovo ...

GEOLOGICA BALCANICA, 35. 3—4. Sofia, Dec. 2006, p. 5—40. 

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) 

Introduction 

Ä. Êîâà÷åâ, È. Íèêîëîâ – Deinotherium thraceiensis sp. nov. èç ìèîöåíñêèõ îòëîæåíèé 

áëèçè äåðåâíè Åçåðîâî, Ïëîâäèâñêàÿ îáëàñòü. – Íàñòîÿùàÿ ñòàòüÿ ïðåäîñòàâëÿåò ìåæäóíàðîäíîìó 

ïàëåîíòîëîãè÷åñêîìó îáùåñòâó îïèñàíèå ïîëíîãî ñêåëåòà äåéíîòåðèÿ, îáíàðóæåííîãî 

è âûêîïàííîãî Ä. Êîâà÷åâûì â 1965 ãîäó èç íåîãåíà âáëèçè äåðåâíè Åçåðîâî, è 

îïèñàííîãî, ðåñòàâðèðîâàííîãî è ìîíòèðîâàííîãî ïîêîéíèì Èâàíîì Íèêîëîâûì. Ôîññèëüíûå 

îñòàíêè íàéäåííûå â ñåäèìåíòàõ ìýîòñêîãî âîçðàñòà. Äî ýòîãî, â òåõ-æå îòëîæåíèé 

íàéäåííûå òàêæå ôîññèëüíûå êîñòè Trilophodon angustidens Cuvier, Hipparion microtaton 

Nicolov è ÷àñòü ÷åëþñòè Mastodon sp. Ïðè âñêðûòèè ñêåëåòà òîëüêî íåìíîãèå êîñòè íå 

õâàòàëî, à èìåííî, çàäíàÿ ëåâàÿ áåäðåííàÿ êîñòü, çàäíàÿ ëåâàÿ ôèáóëà, áîëüøèíñòâî ðåáð è 

íåêîòîðûå èç êàóäàëüíûõ ïîçâîíêîâ. Îíè ðåñòàâðèðîâàííûå íà îñíîâå ñóùåñòâóþùèõ ñèììåòðè÷åñêèõ 

àíàëîãîâ. Ðàçìåðû êîñòåé è ñðàâíåíèå ñ äðóãèìè äåéíîòåðèÿìè îïèñàííûìè 

â ëèòåðàòóðå ïîêàçàëî íåêîòîðûå ñóùåñòâåííûå ðàçëè÷èÿ. Íåêîòîðûå ðàçëè÷èÿ ïî îòíîøåíèè 

÷åðåïà èìåþò áåç ñîìíåíèÿ òàêñîíîìè÷åñêîå çíà÷åíèå. Â ñâÿçè ñ ýòèì, îïèñàííûé 

äåéíîòåðèé îòíîñèòñÿ ê íîâîìó âèäó 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. Kovachev 

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 Trilophodon 

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. Therefore, 

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, 3—4; 5—40. 

Key words: Deinotherium thraceiensis, Miocene, Maeotian, Bulgaria. 

The fossil materials presented herewith to the scientific 

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 Department 

of Palaeontology at the Geological Insti- 

tute, Bulgarian Academy of Sciences, that the restoration 

process was finished. The assembled skeleton 

has been mounted and permanently exhibited in the 

Museum of Geology and Palaeontology, Sofia University 

“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 

5

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 species 

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 labour 

put into the description of the bones, the restoration 

and mounting of this unique skeleton, remained 

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 responsibility 

undertaken, and that the whole blame for eventual 

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 different 

time. Prof. Dr. R. Dehm (Director of Universität-Institut 

für Paläontologie und historische Geologie 

– München) and Prof. Dr. H. Tobien (Director 

of Paläontologisches Institut der Johanes-Gutenberg 

Universität – Meinz) helped I. Nikolov during the 

first studies and restoration. Academician T. Nikolov 

(Geological Institute, Bulgarian Academy of Sciences) 

gave valuable advice with the material. D. Kovachev 

wishes to express his gratitude to Dr. Marin 

Ivanov from the Sofia University for his responsiveness 

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 Töteboch. 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 Deinotherium 

remains found. 

One century later, in 1715, Réaumure admitted 

he could not refer the bones to any known animal. 

Kenedy supposed in 1775 that the materials were related 

to mammoths. G. Cuvier thought in 1779—1836 

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 mandible 

found near Eppelsheim, Germany. He thought 

this was an intermediary form between the sloth and 

the mastodons and referred them to the larger taxonomic 

unit Curtognati. In 1841 and 1857 he de- 

6 

scribed further remains of that genus found near 

Eppelsheim and Westhofen. According to him, the 

adult animals had five teeth in their upper and lower 

jaws. In 1833 he described the new species D. cuvieri, 

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 Mastodon 

should be referred to Proboscidea. He recognized 

the existence of three species: D. giganteum Kaup = 

Tapir gigantesque Cuvier; D. intermedium and D. cuvieri 

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 1858—1859 by M. Lartet, and namely, Deinotherium 

bavaricum H. v. Meyer, D. giganteum Kaup, D. sp. (an 

intermediary form between the first two), and D. cuvieri 

Kaup. The first species is known from the Miocene 

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 (Siwalik), 

Europe (France, Switzerland, Austria), etc. have 

been reported during the last quarter of the 19 th century 

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 Manzati, 

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 Hungary, 

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 species, 

and illustrating skulls/mandibles and teeth from 

D. giganteum Kaup, D. bavaricum H. v. Meyer, D. 

cuvieri Kaup, D. pentapotamiae Lydekker, D. indicum 

Falconer, etc. (Fig. 1). 

Numerous new finds of deinotheres have been reported 

during the whole 20 th century from Austria, 

Bohemia, Bulgaria, France, Germany, Greece, Hungary, 

Macedonia, Moldova, Pakistan, Romania, Russia, 

Serbia, Ukraina, etc. 

Studies on the genus Deinotherium 

in Bulgaria 

The first Deinotherium remains in Bulgaria have been 

found by G. Bonchev (Áîí÷åâ, 1897) who made ex-

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. 

Áàêàëîâ (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 determined 

two species – D. giganteum Kaup race mi- 

7

nor and D. giganteum Kaup race major. To the first 

he referred the smaller teeth, and to the second – 

the larger ones. Later on (Áàêàëîâ, 1949/1950) 

Bakalov described Deinotherium teeth found at the 

“Meander of the Konska river” near the village Noevtsi; 

the village (now – a town; for some time called 

Batanovtsi) Temelkovo (Pernik region) and Aksakovo, 

near Varna, as D. giganteum Kaup race major 

(from the first two localities, Pliocene) and D. giganteum 

Kaup race minor (from the Sarmatian limestones 

near Aksakovo). Íèêîëîâ (1960) reported a 

molar fragment of D. giganteum Kaup race major 

from the lignite coals near the village Hrabarsko (Sofia 

region), assuming a Pontian age. The whole fossil 

material belonging to genus Deinotherium then known 

from Bulgaria has been published by Áàêàëîâ, 

Íèêîëîâ (1962). They recognize two races – minor 

and major, and refer the finds as follows: two mandibles, 

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 Levantian. 

Íèêîëîâ (1962) referred the molar and fragment 

of a scapula found in the Levantian sediments 

near the village Popitsa, Vratsa region to D. giganteum 

Kaup. Íèêîëîâ, Êîâà÷åâ (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 because 

of the Anancus arvernensis Croizet et Jobert 

teeth found there, referring the find to D. giganteum 

Kaup. Êîâà÷åâ (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. Öàíêîâ, Íèêîëîâ 

(1966) published a brief review of the proboscideans 

in Bulgaria, and gave also some preliminary information 

about the finding of Kovachev reporting 

metrical data on some bones. 

PLATE I 

8 

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 remains, 

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 Viteževo and Porodin – D. giganteum 

Kaup. Near Veles and Ravanica, together 

with D. giganteum Kaup, D. bavaricum H. v. Meyer 

is also present. The sediments near the villages Meduhana, 

Ravanica and those between Viteževo and 

Porodin are of Lower Sarmatian, and the rest – of 

Pontian age. 

2. Drenovica, Chukovica, Bresnica, Prebrezka, Zdarski 

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 (Pontian), 

and Samos (Middle Pontian). From the Sarmatian 

of Chios D. bavaricum H. v. Meyer has been 

described. 

From Turkey (Asia Minor), D. giganteum Kaup 

has been described from Küçükçekmece (Maeotian). 

From the territory of Romania and part of Bessarabia, 

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 Solcustria; 

and D. gigantissimum Stefanescu, – from the Pliocene 

sediments near Manzati, Breaza and Cimislia. 

From the territory of Moldova and Bessarabia, two 

taxa are known too: 1. D. giganteum Kaup near Taraklia, 

Ciobruciu, Sofijevka, Tanovka, Chernevo, Ti- 

Deinotherium thracåiensis 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 


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 

→ 


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 VI 

PLATE V

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 XXI 

PLATE XX 

PLATE XIX 

PLATE XXII

PLATE XXIII 

PLATE XXIV

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) 

rasspol, Donau, Velikovo and Krivoy Rog. The sediments 

are of Maeotian, even Upper Sarmatian age. 

2. D. gigantissimum Stefanescu near Volchinec, Rahny-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), Noevtsi, 

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 (farmers, 

workers, etc.). Therefore, the exact location has 

been mentioned only approximately, and there is no 

stratigraphic profile. In such cases, a probable age 

← 

PLATE V 


1. Mandible. Scale ca. 8.3% coll. SU.M. No. SU Pl 312/2; Locality: Ezerovo, near Plovdiv; Level: Maeotian 

PLATE VI 


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 

2 Geologica Balcanica, 3-4/2006 

has been assumed after the age of the sediments in 

the noted areas. 

The vertical distribution of deinotheres in Bulgaria 

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 Sarmatian 

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- 

9

10 

Fig. 3. Position of the locality of Ezerovo in the Upper Thracian plain 

Fig. 4. Stratigraphic profile of the locality Ezerovo

hole (No 16; Áðúíêèí, Ñòàí÷åâà, 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 (Áðúíêèí, Ñòàí÷åâà, 1965). The 

maximum thickness of the Lower Miocene is about 

180 m. 

The Palaeogene and Lower Miocene marine sediments 

are unconformably covered by fluvial sediments 

referred to the Ahmatovo Formation (Êîþìäæèåâà, 

Äðàãîìàíîâ, 1979). They crop out at numerous 

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 conglomerate 

lenses. At places the sand layers are crossbedded. 

Petrified trees occur in the sands. The thickness 

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 Formation 

is referred to the Maeotian, Pontian and Dacian 

regional Paratethys stages. Three macrocycles 

have been distinguished (Êîþìäæèåâà, Äðàãîìàíîâ, 

1979). The first macrocycle is referred to the 

Maeotian (possibly also upper parts of the Sarmatian) 

on the basis of Lower to Middle Turolian (“Pikermian”) 

mammal fauna. The Pontian age of the 

lower parts of the second cycle is determined by Upper 

Pikermian (= Upper Turolian) fauna, and the upper 

parts of the second cycle are referred to the Dacian 

Regional Stage by Ruscinian fossil remains. The 

fossil locality of Deinotherium thraceiensis sp. n. belongs 

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 4—5 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 mediterraneum 

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 resembles 

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 longirostris 

Kaup. 

6. M 3 sin – Anancus sp. 

7. M 2 and M 3 dext and M 3 sin – Zygolophodon borsoni 

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 Pleistocene. 

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 


(Pl. I – XXII) 

Derivatio nominis: thraceiensis from Greek “Thraceia” 

– Thrace, the region of the find. 

Holotype. As such we design all the bones of the skeleton 

– 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. Nasal 

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. Forehead 

high and, compared to the external nares – 

short. The occipital bone is high and wide. Symphysis 

of the mandible curved downwards and backwards. 

Incisors of moderate length, slightly curving 

outwards. Their tips end exactly beneath processus 

angularis. Neck short, with seven thin cervical vertebrae. 

First ones lack processus spinosus, in the last 

ones it is weakly developed. Skull and mandible, relatively 

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. 

11

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 rugosity. 

Nasal bones (os nasale) are short, high in their anterior 

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 suture. 

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 wider 

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 forehead. 

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 (frontoparietal 

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 rounded, 

under the condyles and by the parietal bone almost 

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 relation 

to the forehead this bone is almost perpendicularly 

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 occipital 

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 lacrimale, 

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).

Table 1 

Skulls – Crania 

D.bavaricum 

H. v. Meyer 

Bavaria 

D.gigantissimum 

Stefanescu Manzati 

D.giganteum Kaup 

Munich 

D.giganteum Kaup 

Eppelsheim 

№ Dimensions, mm D.thraceiensis 

Ezerovo 

1. Total length 1320.0 121.0 1035.0 1260.0 860.0 

2. Length from os nasale to cond. occipitalis 1290.0 900.0 950.0 1118.0 780.0 

3. Length from os nasale to os occipitale 1012.0 840.0 - - - 

4. Width at the basis of os occipitale 165.0 240.0 252.0 270.0 - 

5. Width at the basis of the external nares (nasale) 570.0 - 108.0 - - 

6. Width in the middle of the external nares (nasale) 680.0 - 170.0 - 

7. Width in eye sockets 730.0 - 500.0 - - 

8. Width in ear area 540.0 570.0 - - - 

9. Height of the skull at M 1 715.0 - 360.0 - - 

10. Height of the skull at M 3 805.0 550.0 380.0 630.0 385.0 

11. Length of ear area 380.0 530.0 - 540.0 260.0 

12. Height of eye socket 185.0 

13. Width of eye socket 120.0 

14. Depth of eye socket 160.0 - 

15. Length of os nasale to the anterior end of P 3 350.0 324.0 330.0 430.0 300.0 

16. Width of os nasale in premaxillaria 260.0 

17. Width of os nasale to the anterior end of P 3 130.0 

18. Width of os nasale in the middle between these two points 185.0 

19. Minimum height of os nasale 250.0 160.0 108.0 210.0 - 

20. Beginning of external nares from the beginning of os nasale 150.0 

21. Width of external nares at the first curve 160.0 

22. Width of external nares at the second curve 300.0 

23. Width of external nares at the eye socket 360.0 

24. Depth of the external nares at the first curve 154.0 

25. Depth of the external nares at the second curve 165.0 

26. Depth of the external nares in the middle 500.0 

27. Length of os frontale 308.0 - - - - 

28. Minimum width of os frontale 500.0 - - - - 

29. Maximum width of os frontale 540.0 - - - - 

30. Length of os occipitale 850.0 - - - - 

31. Height of os occipitale 700.0 450.0 - 420.0 450.0 

32. Height of condylus occipitalis 196-sin 

176-dext 

33. Maximum width of condylus occipitalis 126-sin 

134-dext 

34. Maximum height of foramen magnum 155.0 

35. Maximum width of foramen magnum 75.0 

36. Length of arcus zygomaticus 440.0 

37. Width of arcus zygomaticus 95-100 

38. Maximum depth of palatum durum 176.0 

39. Width of palatum durum at P 3 550.0 

40. Length of tooth row P 3 -M 3 510.0 470.0 390.0 540.0 252.0 

13

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 (measurements 

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 dentin 

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 differences, 

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 correspondingly. 

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 cingulum 

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 developed 

on the whole anterior and the posterior outer 

part of the crown. 

The fourth premolar (P 4 dext) has an almost tetragonal 

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 divided 

in two parts, and V-shaped. Thus the ridges surround 

a deep triangular valley with steep walls. On 

the inner and outer posterior side and on the anterior 

side of the crown there is a slightly developed cingulum. 

The left P 4 has a weaker cingulum on its anterior 

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 middles. 

The second and third ridges are slightly concave 

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 anterior 

wall of the first ridge, surrounds a long and 

shallow valley. 

14 

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 posterior 

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 posterior 

talon. 

Comparison 

According to the literature, there are only two completely 

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 referred 

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 skeleton 

recently mounted in Kishinev only part of the 

maxilla with the upper teeth is preserved. 

In conclusion, it seems that the only possible comparisons 

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 preorbital 

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 rugged 

with a tuberosity protruding forward. On it, according 

to Svistun 1974, there could be horn-like structures 

substituting for the upper tusks. There is no such 

thing in our material, neither in the Eppelsheim skull.

Table 2 

Upper cheek-teeth 

D. giganteum Kaup 

D. thraceensis 

D. gigantissimum 

Measurements in mm 

sp.n. Ezerovo, Bulgaria Stefanescu, 1909, Romania racemjor Bakalov, Nikolov, 

raceminor, Bakalov, 

1962, Bulgaria 

Nikolov, 1962, Bulgaria 

sin dext sin dext sin dext sin dext sin dext 

Length of the tooth 100.0 100.0 - - 102.0 - - - - 36.0 

Width of the tooth 105.0 106.5 - - 100.0 - - - - 30.0 

P 3 Width of first ridge 90.5 91.8 - - 97.0 - - - - 17.0 

Width of second ridge 106.5 106.0 - - 106.0 - - - - 36.0 

Enamel thickness 4.0 4.2 - - - - - - - - 

Length of the tooth 75.0 81.5 87.0 - 80.0 - - - - 51.0 

Width of the tooth 100.8 98.5 - - - - - - - 50.0 

P 4 Width of first ridge 96.3 87.0 96.0 - 68.0 - - - - 42.5 

Width of second ridge 100.5 102.5 - - 62.0 - - - - 50.0 


Length of the tooth 110.0 110.0 119.0 115.0 99.0 103.0 92.6 - - 74.0 

Width of the first ridge at the base 94.6 100.0 100.0 93.0 82.0 73.0 75.0 - - 52.0 

Width of the first ridge at the top 70.0 73.0 - - - - - - - 29.5 

Width of the second ridge at the base 94.0 97.0 98.0 92.0 65.0 70.0 61.0 - - 57.0 

M 1 Width of the second ridge at the top 69.0 71.0 - - - - - - - 33.0 

Width of the third ridge at the base 83.5 84.5 86.0 83.0 - - 53.0 - - 48.5 

Width of the third ridge at the top 63.5 62.0 - - - - - - - 31.0 

Width of posterior talon - - - - - - - - - - 


Length of the tooth 107.7 106.5 116.0 114.0 103.0 92.0 86.0 91.0 70.0 71.0 

Width of the first ridge at the base 113.6 115.0 129.0 111.0 103.0 98.0 96.0 101.0 62.0 70.5 

Width of the first ridge at the top 88.8 87.5 - - - - - - 44.0 52.0 

M 2 Width of the second ridge at the base 110.0 110.0 117.0 114.0 102.0 90.0 84.5 83.0 59.0 73.0 

Width of the second ridge at the top 85.3 84.5 - - - - - - 38.0 51.5 



Length of the tooth 110.0 108.0 115.0 107.0 96.6 105.0 89.0 96.0 - - 

Width of the first ridge at the base 118.0 115.0 114.0 114.0 103.0 107.0 95.0 105.6 - - 

Width of the first ridge at the top 100.0 98.0 - - - - - - - - 

M 3 Width of the second ridge at the base 107.2 107.0 107.0 104.0 98.0 101.0 89.0 89.0 - - 

Width of the second ridge at the top 87.0 89.0 - - - - - - - - 

Width of posterior talon 78.0 78.0 - - - - - - - - 


15

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. 

18 

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 situation 

is closer to our case. 

7. The zygomatic arch (Figs. 5, 6, and 7 – zyg) of D. 

thraceiensis sp. n. with its developed proc. postorbitalis 

resembles the one of D. levius Jourdan, but in its 

posterior part, behind its connection with the temporal 

bone, it lacks the processus that Svistun notes. 

He regards this processus as related to the development 

of the lower tusks. No comparison with the zygomatic 

arch of the Eppelsheim skull is possible, 

because it is not preserved.

8. There are significant differences between the three 

skulls in the occipital region. Os occipitale in D. thraceiensis 

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 lacking 

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 lower 

part of the occipital bone, almost the same is their 

position in D. giganteum Kaup, and in D. levius Jourdan 

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 includes 

D. giganteum Kaup, D. gigantissimum Stefanescu, 

D. thraceiensis sp. n., D. levius Jourdan and 

D. indicus Falconer, which have larger teeth, and the 

second – D. bavaricum H. v. Meyer, D. pentapotamiae 

Lydekker and others, all with smaller teeth. 

As a whole, the teeth are rather similar and differences 

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 perpendicular 

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. gigantissimum 

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 diagonal 

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 valley 

surrounded by the posterior inner wall of the outer 

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 anterior 

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 ridges 

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 posterior 

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. thraceiensis 

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 tubercles, 

closely arranged along the end of the tooth. 

There are no significant differences in the structure 

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 strongly 

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 symphysis. 

Together with the tusks, it is curved down and 

back. In D. thraceiensis sp. n. the symphysis is solid, 

19

16 

Table 2 A 


D. giganteum 

Deperet, 1887 


D. bavaricum H. v. Meyer, 

Germany 

D. thraceiensis 

sp.n. Ezerovo, 

after 

after 

after 

from from 

Bulgaria 

Петронjевиh, 

А. Белокрые, Stromer, 

Athanasia, 1907, 

Seint Eppel 

1954, 

1960, Krivoi Münhen 

Romania 

Yean sheim 

Yugoslavia 

Rog, Moldova 

sin dext sin dext sin dext sin dext sin dext sin dext 


Length of the tooth 100.0 100.0 - 73.5 87.0 - 98.0 96.0 72.0 71.0 85.0 83.0 66.0 66.0 52.0 53.0 

Width of the tooth 105.0 106.5 - 69.0 100.0 - - - - - - - 61.0 61.0 33.0 - 

Width of second ridge 90.5 91.8 - 68.2 98.0 - 97.0 95.0 69.0 71.0 - - - - 47.5 - 

Width of second ridge 106.5 106.0 - 70.4 70.0 - - - - - - - - - - - 

Enamel thickness 4.0 4.2 - - 5.0 - - - - - - - - - - - 

Length of the tooth 75.0 81.5 - 67.5 62.0 - 88.0 86.0 68.0 63.0 77.0 78.0 61.0 61.0 50.0 - 

Width of the tooth 100.8 98.5 - 79.0 50.0 - - - - - - - 66.0 66.0 51.0 - 

Width of first ridge 96.3 87.0 - - 55.0 - 99.0 98.0 74.0 73.0 - - - - - - 

Width of second ridge 100.5 102.5 - 75.0 37.0 - - - - - - - - - - - 

Enamel thickness 4.2 4.3 - - - - - - - - - - - - - - 

Length of the tooth 110.0 110.0 61.2 - 119.0 - 104.0 - 81.0 82.0 - 95.0 75.0 75.0 72.0 - 

Width of the first ridge at the base 94.6 100.0 50.5 - 100.0 - 93.0 - 68.0 68.0 - - 62.0 62.0 54.0 - 

Width of the first ridge at the top 70.0 73.0 - - 48.0 - - - - - - - - - - - 

Width of the second ridge at the base 94.0 97.0 70.8 - 98.0 - - - - - - - - - 54.0 - 

Width of the second ridge at the top 69.0 71.0 - - 45.0 - - - - - - - - - - - 

Width of the third ridge at the base 83.5 84.5 - - 86.0 - - - - - - - - - 44.0 - 

Width of the third ridge at the top 63.5 62.0 - - - - - - - - - - - - - - 

Width of posterior talon - - - - - - - - - - - - - - - - 


Length of the tooth 107.7 106.5 79.5 - - 115.0 99.0 - 80.0 78.0 - 91.0 68.0 68.0 60.0 64.0 

Width of the first ridge at the base 113.6 115.0 80.5 - - 88.0 - - 82.0 81.0 - - 66.0 66.0 - 61.5 

Width of the first ridge at the top 88.8 87.5 - - - - - - - - - - - - - - 

Width of the second ridge at the base 110.0 110.0 79.0 - - 89.0 108.0 - - - - - - - - 61.0 

Width of the second ridge at the top 85.3 84.5 - - - - - - - - - - - - - - 

Width of posterior talon - - - - - 75.0 - - - - - - - - - - 


Length of the tooth 110.0 108.0 - 70.0 - - - - 81.0 82.0 - 91.0 67.0 63.0 62.0 63.0 

Width of the first ridge at the base 118.0 115.0 - 73.0 - - - - 89.0 89.0 - - 70.0 70.0 - 63.0 

Width of the first ridge at the top 100.0 98.0 - - - - - - - - - - - - - - 

Width of the second ridge at the base 107.2 107.0 - 69.0 - - - - - - - - - - - 56.0 

Width of the second ridge at the top 87.0 89.0 - - - - - - - - - - - - - - 

Width of posterior talon 78.0 78.0 - - - - - - - - - - - - - - 


P 3 

P 4 

M 1 

M 2 

M 3

Table 2 B 


Rhone – Deperet, 1887 after Lydekker, 1880 

D. pentapotamiae 

D. thraceiensis sp.n. 

after Dehm 1963, 


Ezerovo, Bulgaria D. levius D. giganteum D. cuvieri D. 

D. indicum Fal. 

Pakistan 

Jordan Kaup Kaup pentapotamiae Narbada, India 

sin dext 

sin dext sin 

Length of the tooth 100.0 100.0 66.0 70.0 53.0 - - 50.0 51.0 58.4 

Width of the tooth 105.0 106.5 - - - - - 45.0 51.0 58.2 

P 3 Width of second ridge 90.5 91.8 - - - - - 51.0 52.0 60.0 

Width of second ridge 106.5 106.0 - - - - - - - - 


Length of the tooth 75.0 81.5 78.0 - - - - - 52.4 

Width of the tooth 100.8 98.5 - - - - - - 64.4 

P 4 Width of first ridge 96.3 87.0 - - - - - - 62.2 

Width of second ridge 100.5 102.5 - - - - - - - 

Enamel thickness 4.2 4.3 - - - - - - - 

Length of the tooth 110.0 110.0 95.0 72.0 2.3 3.9 72.0 70.0 68.4 

Width of the first ridge at the base 94.6 100.0 - - 2.1 2.5 62.0 52.2 62.2 

Width of the first ridge at the top 70.0 73.0 - - - - - - - 

Width of the second ridge at the base 94.0 97.0 - - - - 60.3 - 62.2 

M 1 Width of the second ridge at the top 69.0 71.0 - - - - - - - 

Width of the third ridge at the base 83.5 84.5 - - - - - - - 

Width of the third ridge at the top 63.5 62.0 - - - - - - - 

Width of posterior talon - - - - - - - - - 


Length of the tooth 107.7 106.5 91.0 60.0 2.1 2.5 - - 71.3 

Width of the first ridge at the base 113.6 115.0 - - 2.5 2.4 - - 73.2 


M 2 Width of the second ridge at the base 110.0 110.0 - - - - - - 71.8 

Width of the second ridge at the top 85.3 84.5 - - - - - - - 

Width of posterior talon - - - - - - - - - 


Length of the tooth 110.0 108.0 91.0 62.0 2.75 - 67.0 66.0 68.4 

Width of the first ridge at the base 118.0 115.0 - - 2.4 - - 70.0 74.0 


M 3 Width of the second ridge at the base 107.2 107.0 - - - - 69.0 59.6 64.3 

Width of the second ridge at the top 87.0 89.0 - - - - - - - 

Width of posterior talon 78.0 78.0 - - - - - - - 



Note: In the columns after Lydekker, 1880, the size is in inches. 

17

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. 

18 

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 situation 

is closer to our case. 

7. The zygomatic arch (Figs. 5, 6, and 7 – zyg) of D. 

thraceiensis sp. n. with its developed proc. postorbitalis 

resembles the one of D. levius Jourdan, but in its 

posterior part, behind its connection with the temporal 

bone, it lacks the processus that Svistun notes. 

He regards this processus as related to the development 

of the lower tusks. No comparison with the zygomatic 

arch of the Eppelsheim skull is possible, 

because it is not preserved.

8. There are significant differences between the three 

skulls in the occipital region. Os occipitale in D. thraceiensis 

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 lacking 

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 lower 

part of the occipital bone, almost the same is their 

position in D. giganteum Kaup, and in D. levius Jourdan 

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 includes 

D. giganteum Kaup, D. gigantissimum Stefanescu, 

D. thraceiensis sp. n., D. levius Jourdan and 

D. indicus Falconer, which have larger teeth, and the 

second – D. bavaricum H. v. Meyer, D. pentapotamiae 

Lydekker and others, all with smaller teeth. 

As a whole, the teeth are rather similar and differences 

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 perpendicular 

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. gigantissimum 

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 diagonal 

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 valley 

surrounded by the posterior inner wall of the outer 

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 anterior 

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 ridges 

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 posterior 

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. thraceiensis 

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 tubercles, 

closely arranged along the end of the tooth. 

There are no significant differences in the structure 

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 strongly 

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 symphysis. 

Together with the tusks, it is curved down and 

back. In D. thraceiensis sp. n. the symphysis is solid, 

19

20 

Table 3 

Mandibles – mandibulae 

D. thraceiensis sp.n. D. gigantissimum D. giganteum Kaup D. bavaricum H. v. Mayer 

№ Dimensions, mm 

Ezerovo 

Romania 

Moldova München H. v. Meyer München 

1. Total length – from the beginning to the end of ramus ascendens 1120.0 1200.0 1000.0 1120.0 820.0 840.0 

2. Length of symphysis – horizontal 360.0 420.0 - 320.0 230.0 235.0 

3. Length of symphysis – vertical 435.0 - - - - - 

4. Diameter of the symphysis at the angle with the mandible 310.0 - - 

5. Diameter of the symphysis at the beginning of I 280.0 - - 

6. Width of the through of the symphysis 145.0 - - 

7. Length of symphysis and ramus ascendens 1100.0 1150.0 - 

8. Height of the mandible at P3 235.0 - - 

9. Height of the mandible at M1 230.0 190.0 230.0 210.0 160.0 154.0 

10. Height of the mandible at M3 185.0 - 192.0 

11. Height of ramus ascendens at proc. articularis 685.0 680.0 455.0 

12. Height of ramus ascendens at proc. coronoideus 505.0 - - 

13. Width – middle of ramus ascendens 470.0 482.0 - 

14. Distance between the two semimandibles at P3 105.0 - - 

15. Distance between the two semimandibles at M1 150.0 - - 

16. Distance between the two semimandibles at M3 120.0 - - 

17. Length of the incisors – I 855.0 608.0 900.0 795.0 530.0 525.0 

18. Diameter of I – at the basis 132.0 172.0 142.0 

19. Diameter of I – in the middle 125.0 120.0 125.0 

20. Distance between the points of I 470.0 - - 

21. Distance between the point of I and ramus ascendens 895.0 - - 

22. Length of the entire tooth row P3-M1 510.0 540.0 470.0 485.0 370.0 365.0

Table 4 

Lower cheek teeth 




sp.n. Ezerovo, Bulgaria Stefanescu, 1909, Romania 

Bakalov, Nikolov, 1962, 

raceminor, Bakalov, 


Bulgaria 

Nikolov, 1962, Bulgaria 


Length of the tooth 80.5 75.5 - - - 80.0 - - - - 

Width of the tooth 68.0 67.0 - - - - - - - - 

P 3 Width of first ridge 58.0 55.0 - - - 68.0 - - - - 

Width of second ridge 70.0 73.0 - - - 62.0 - - - - 


Length of the tooth 89.0 89.0 90.0 - - 92.0 - - - - 

Width of the tooth 80.0 81.0 - - - - - - - - 

P 4 Width of first ridge 76.0 78.0 72.0 - - 88.0 - - - - 

Width of second ridge 81.1 83.0 73.0 - - 86.0 - - - - 


Length of the tooth 107.5 107.0 111.0 109.0 - 111.0 - - 109.0 73.0 

Width of the first ridge at the base 82.0 81.5 72.0 77.0 - 85.0 - - 81.0 73.0 

Width of the first ridge at the top - - - - - - - - - - 

Width of the second ridge at the base 81.0 81.5 72.0 - - 82.0 - - 76.0 37.0 

M 1 Width of the second ridge at the top - - - - - - - - - - 

Width of the third ridge at the base 75.0 75.5 71.0 70.0 - 75.0 - - - - 

Width of the third ridge at the top - - - - - - - - - - 



Length of the tooth 115.5 112.5 111.0 - 106.0 104.0 - - 71.0 72.0 

Width of the first ridge at the base 103.0 110.5 99.0 - 84.0 103.0 - - 70.5 67.0 

Width of the first ridge at the top 80.0 81.0 - - - - - - - - 

M 2 Width of the second ridge at the base 97.5 107.0 97.0 - 82.0 99.5 - - 73.0 76.0 

Width of the second ridge at the top 83.0 75.5 - - - - - - - - 



Length of the tooth 120.0 118.0 115.0 - 100.5 110.0 - 115.5 - - 

Width of the first ridge at the base 106.0 105.0 114.0 - 84.0 110.0 - 92.0 - - 

Width of the first ridge at the top 85.0 82.0 - - 68.0 - - - - - 

M 3 Width of the second ridge at the base 95.0 100.0 107.0 - 83.0 93.0 - 96.0 - - 

Width of the second ridge at the top 78.5 80.0 - - 66.5 - - - - - 

Width of posterior talon 64.0 67.0 - - 65.0 - - 56.0 - - 


21

22 

Table 4 A 




Simionescu. 

Петронjевиh, 1954, А. Белокрые, 1960, Stromer, 1938, R. Deim, 1949, South Bavaria, 



1939, 

Yugoslavia Krivoi Rog, Moldova München 

Museum of München 

Bulgaria 

Romania 

sin dext sin dext sin dext sin dext sin dext sin dext sin 

Length of the tooth 80.5 75.5 58.2 40.0 81.0 - 60.0 - 62.0 - 58.0 - - 

Width of the tooth 68.0 67.0 - - 64.0 - - - 55.0 - 47.0 - - 

P 3 Width of first ridge 58.0 55.0 38.5 28.0 - - - - - - - - - 

Width of second ridge 70.0 73.0 49.1 - - - 55.0 - - - - - - 

Enamel thickness 4.0 3.8 - - - - - - - - - - - 

Length of the tooth 89.0 89.0 64.4 50.5 - 88.0 68.0 - 68.0 57.0 69.0 67.0 - 

Width of the tooth 80.0 81.0 - - - 99.0 - - 60.0 47.0 62.0 61.0 - 

P 4 Width of first ridge 76.0 78.0 57.5 39.0 - - 55.0 - - - - - 21.5 

Width of second ridge 81.1 83.0 - 40.0 - - 56.0 - - - - - 27.5 


Length of the tooth 107.5 107.0 - - - - 75.0 - 80.0 - 83.0 82.0 - 

Width of the first ridge at the base 82.0 81.5 - - - - 55.0 - - - - - 22.0 

Width of the first ridge at the top - - - - - - - - - - - - - 

Width of the second ridge at the base 81.0 81.5 59.7 - - - - - - - 62.0 - 28.0 

M 1 Width of the second ridge at the top - - - - - - - - - - - - - 

Width of the third ridge at the base 75.0 75.5 - - - 52.0 - 60.0 - - - - 

Width of the third ridge at the top - - - - - - - - - - - - 

Width of posterior talon - - - - - - - - - - - - 

Enamel thickness 4.3 4.2 - - - - - - - - - - 

Length of the tooth 115.5 112.5 76.7 102.0 - 79.0 - 82.0 76.0 80.0 78.0 - 

Width of the first ridge at the base 103.0 110.5 68.7 - 86.0 71.0 - 76.0 68.0 76.0 74.0 - 

Width of the first ridge at the top 80.0 81.0 - - - - - - - - - - 

M 2 Width of the second ridge at the base 97.5 107.0 66.0 - - 70.0 - - - 76.0 76.0 - 

Width of the second ridge at the top 83.0 75.5 - - - - - - - - - - 

Width of posterior talon - - - - - - - - - - - - 

Enamel thickness 4.6 4.5 - - - - - - - - - - 

Length of the tooth 120.0 118.0 - 105.0 107.0 100.0 - 93.0 81.0 86.0 83.0 - 

Width of the first ridge at the base 106.0 105.0 - 93.0 100.0 ? - 78.0 68.0 76.0 76.0 - 

Width of the first ridge at the top 85.0 82.0 - - - - - - - - - - 

M 3 Width of the second ridge at the base 95.0 100.0 62.0 - - - - - - 76.0 76.0 - 

Width of the second ridge at the top 78.5 80.0 - - - - - - - - - - 

Width of posterior talon 64.0 67.0 - - - - - - - - - - 

Enamel thickness 5.2 5.3 - - - - - - - - - -

Table 4 B 



Rhone 

from Siwaliku Narbada, 

D. bavaricum H.v. 

D. pentapotamiae 


Deperet, 1887 

after Lydekker, 1889, India 

Meyer, R.Dehm, 1949, 

Lydekker, after 


Bulgaria 

D. D. 

D. 

South Bavaria, Museum of 

D. cuvieri D. 

R. Dehm, 1963, 

levius giganteum 

Lydekker indicum 

sin dext 

Münhen 

Kaup pentapotamiae 

Pakistan 

Jordan Kaup 

Falconer 

Length of the tooth 80.5 75.5 44.0 43.0 - 55.0 63.0 43.0 1.8 1.47 - - 54.0 

Width of the tooth 68.0 67.0 - 36.0 - - - - 2.1 2.05 - - 33.0 

P 3 Width of first ridge 58.0 55.0 - - - - - - - - - - 42.5 

Width of second ridge 70.0 73.0 - - - - - - - - - - - 


Length of the tooth 89.0 89.0 43.0 50.0 57.0 70.0 70.0 48.0 1.8 - 2.9 56.0 - 

Width of the tooth 80.0 81.0 - 42.0 - - - - 1.7 - 2.6 42.0 - 

P 4 Width of first ridge 76.0 78.0 - - 47.0 - - - - - - - - 

Width of second ridge 81.1 83.0 - - - - - - - - - 45.8 - 


Length of the tooth 107.5 107.0 72.0 67.0 68.0 60.0 2.34 1.8 4.0 67.0 - 

Width of the first ridge at the bas 82.0 81.5 47.0 - - - 2.4 2.1 2.8 47.0 - 

Width of the first ridge at the top - - - - - - - - - - - 

Width of the second ridge at the base 81.0 81.5 50.0 41.0 45.0 - - - - - - 

M 1 Width of the second ridge at the top - - - - - - - - - - - 

Width of the third ridge at the base 75.0 75.5 - - - - - - - - - 

Width of the third ridge at the top - - - - - - - - - - - 

Width of posterior talon - - - - - - - - - - - 

Enamel thickness 4.3 4.2 - - - - - - - - - 

Length of the tooth 115.5 112.5 67.0 64.0 71.0 59.0 2.4 2.90 3.9 73.0 - 

Width of the first ridge at the base 103.0 110.5 57.0 54.0 - - 2.15 2.15 3.5 65.0 - 

Width of the first ridge at the top 80.0 81.0 - - 56.0 - - - - - - 

M 2 Width of the second ridge at the base 97.5 107.0 58.5 53.0 - - - - - 58.0 - 

Width of the second ridge at the top 83.0 75.5 - - - - - - - - - 

Width of posterior talon - - - - - - - - - - - 


Length of the tooth 120.0 118.0 68.0 70.0 77.0 72.0 2.9 3.18 - 65.0 73.0 

Width of the first ridge at the base 106.0 105.0 59.5 55.0 - - 2.25 2.45 - - - 

Width of the first ridge at the top 85.0 82.0 - - - - - - - - - 

M 3 Width of the second ridge at the base 95.0 100.0 - 51.0 - - - - - - - 

Width of the second ridge at the top 78.5 80.0 - - - - - - - - - 

Width of posterior talon 64.0 67.0 - - - - - - - - - 


Note: In the columns after Lydekker, 1880, the size is in inches. 

23

PLATE VII 


1. Part of vertebral column (vertebrae, ribs). Scale ca. 5% 

Locality: Ezerovo, near Plovdiv; Level: Maeotian 

PLATE VIII 


1—3. 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 


PLATE IX 


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 


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 


1—3. 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 


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 


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 


PLATE XIV 


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 


PLATE XV 


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 


24

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 XXI 

PLATE XX 

PLATE XIX 

PLATE XXII

PLATE XXIII 

PLATE XXIV

PLATE XVI 


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 


PLATE XVII 


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 


PLATE XVIII 


1. Left hand. Scale ca. 7.7%. Coll. SU.M. No. SU Pl 312/13 


PLATE XIX 


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 


PLATE XX 


1. Pelvis in lateral view. Scale ca. 4.0%. Coll. SU.M. No. SU Pl 312/16-1 


PLATE XXI 


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 


PLATE XXII 


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 


PLATE XXIII 


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 


Restored skeleton 

Coll. SU.M. No SU Pl 312 



25

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 protruding 

backwards. This thickness reaches as far as 

proc. articularis. The anterior part of ramus ascendens 

is significantly thinner, however, incl. proc. coronoideus 

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 backwards; 

at the posterior ends there is a moderate concavity 

(incisure). Processus articularis thick and high. 

Perpendicularly positioned, with well shaped articular 

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 symphysis, 

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 inside. 

At the end of the symphysis the tusks are already 

wholly solid. They gradually become thinner 

and, curving in two directions – outwards and backwards, 

go apart from each other. Their tips are pointed, 

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 situated 

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 between 

them in two not quite equal parts. The anterior 

wall of the first ridge is strongly concave. The posterior 

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 

26 

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 tetragonal 

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 mandible 

is close to the other species (Fig. 9). 

The symphysis is moderately large. There are differences 

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. thraceiensis 

sp. n. the curving is even stronger and more 

gradual.

Fig. 9. Comparison of deinothere mandibles: a – Deinotherium 

bavaricum H. v. Meyer; b – Deinotherium levius Jourdan; 

c – Deinotherium giganteum Kaup; d – Deinotherium thraciensis 

sp. n. 

2. In D. thraceiensis sp. n. ramus horizontalis (in the 

region of proc. angualris) strongly protrudes backwards 

with a pronounced curve. It almost reaches 

the level of incisura mandibulae. 

3. Ramus ascendens is almost perpendicular to ramus 

horizontalis in its anterior part. 

4. Processus articularis is high and abruptly separated 

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. articularis. 

6. Incisura mandibulae is deep and almost horizontal 

in relation to the mandible, perpendicular to ramus 

ascendens. 

In D. gigantissimum Stefanescu the curve at proc. 

angularis is weaker. Poc. articularis is not so abruptly 

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 longitudinal 

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 ridges 

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 posterior 

wall of the anterior ridge. The latter is “Cshaped”. 

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 displacement 

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, probably 

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 partially 

broken, without losing the shape of the vertebra. It 

consists of two arches – dorsal and ventral, limiting 

a large foramen vertebrae. The foramen widens downwards 

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 process. 

The atlas articulates with the skull by wide oval 

concavities, cranially positioned on the arches. Caudally, 

on the ventral arch, there is an articular surface 

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 

27

Table 5 

Cervicals 

Table 6 

Thoracics 

№ Dimensions 

1 2 3 4 5 6 

THORACICS 

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 

crests on tip with a deep furrow between them. Dens 

epistrophei connects the axis with the atlas. Foramen 

vertebrae much smaller than in atlas, tetragonal 


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 surfaces 

on the cranial and caudal processes. Foramen 

vertebrae wide and high. Spinous processes not pronounced. 

Sixth and seventh cervicals are built in the same 

way as the previous three, but they have spinous processes, 

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 vertebra 

(Pl. IX, Fig. 4) has a distinct structure. Its body 

is small, cranially convex, caudally – slightly rounded. 

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- 

28 


Atlas Axis 3 

CERVICSLES 

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 

wards the spinous process. All other thoracic vertebrae 

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 foramina 

decreases, then increases to the last. In the fifteenth 

and all the following there is no fovea costalis. 

So, there were no ribs attached to these vertebrae 


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 lumbars 

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 difficult 

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 diameters 

of foramen vertebrae increase as in the last thoracic 

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

it probably has its functional meaning. It should be 

related to the use of the hindlimbs. The accumulation 

of nerve tissue close to the pelvis probably facilitated 

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 caudals, 

but on the mounted skeleton there are 21. The 

first, those after the sacrum, resemble the last lumbars, 

but their processes are gradually decreasing in 

size. In the first, a chanal is still present, which grad- 

Table 7 

Lumbars 

Table 8 

Sacrum 

№ 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 

Table 9 

Caudal vertebrae – vert. coccigiae 

ually disappears. The body of the caudals is elongated, 

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 processes 

of the cervical vertebrae are weaker. These processes 

are strong and high in all known skeletons. A 

difference in D. thraceiensis sp. n. is the large tetragonal 

widening of the tip of this process on the second 

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 unknown. 

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, 

№ 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 

№ 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 - - 

29

30 

Table 10 

Ribs – costae 

№ Dimensions, mm 1 2 3 4 5 6 7 8 9 10 11 12 13 14 

1. Total length 1245.0 1390.0 1450.0 472.0 148.0 1430.0 1376.0 1322.0 1274.0 1230.0 1195.0 1102.0 950.0 840.0 

2. Diameter of caput costae 80.0 82.0 96.0 68.0 69.9 70.5 68.5 67.0 64.0 62.3 60.0 57.2 54.0 50.0 

3. Diameter at the 

98.0 105.0 81.0 85.0 67.0 54.5 52.0 56.3 48.1 46.5 45.0 46.0 45.5 45.0 

thickening 

4. Maximum width of the 80.0 153.0 114.0 91.0 75.0 60.0 60.5 60.9 61.3 61.7 62.0 64.5 67.3 70.0 

rib 

5. Distance between the 50.0 140.0 120.0 136.0 162.0 194.0 198.0 203.5 206.0 213.0 220.0 195.0 180.0 160.0 

beginning of caput 

costae to the thickening 

Note: The measurments in the table are taken for the left ribs 

Table 11 

Scapula 


Munchen - Stromer 


Munchen - Stromer 

D. giganteum 

Kaup 


Pikermi 


St. Manzati, Romania 

Stefanescu 


Ezerovo, Bulgaria 

№ Dimensions 

sin dext 

1. Total length 1135.0 1135.0 970.0 730.0 - 756.0 - 

2. Width c-d 500.0 500.0 300.0 525.0 - 570.0 - 

3. Maximum width m-l 895.0 895.0 650.0 - - 

4. Maximum width of l 650.0 650.0 - - 

5. Maximum width of m 245.0 245.0 205.0 225.0 

6. Diameter at the neck 254.0 245.0 260.0 - - 

7. Diameter at the articular surface 260.0 260.0 280.0 170.0 185.0 170.0 250.0 

8. Height at the articular surface 96.0 110.0 110.0 160.0 

9. Length of tuberositas spinae 635.0 635.0 150.0 - - 

10. Height of acromion 215.0 215.0 90.0 - - 

11. Angle at l 85 o

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 articularis 

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 cartilages. 

Comparison. The most disputable problem concerning 

the ribs is their number. There is still no explicit 

opinion on that matter and we don’t know their precise 

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- 

Table 12 

Humerus 




Bulgaria 

sin dext 


St. Manzati, 

Romania 

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 humerus 

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 posterior 

side – fossa olecrani. It is deeper and wider. 

D. giganteum 

Kaup Saint 

Yean – Lartet 

D. giganteum 

Kaup München – 

Stromer 


München – 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 202.0 207.0 260.0 108.0 260.0 210.0 

major 

9. Diameter of trochanter 

minor 

Table 13 

Ulna 

185.0 190.0 140.0 


D. thraceiensic 


Bulgaria 

sin dext 


St. Manzati, 

D. giganteum 

Kaup Saint Yean 

Romania – 

– Deperet, 1892 

Stefanescu 

D. giganteum 

Kaup München 

№ 495 – Stromer, 

1938 

D. giganteum 

Kaup München – 

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 - 

31

Table 14 

Radius 

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 slightly 

twisted and situated posteriorly, incisura semilunaris 

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 cicumferentia 

articularis. 

Radius (Pl. XIV, Fig. 2 and 2a). This bone is much 

smaller than the ulna. Slightly widening in its proximal 

part, with well pronounced articular surface. 

Slightly rounded in the diaphysis. Ends with a clear- 

32 


D.thraceiensis sp.n. 


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 

ly outlined articular surface. With this part it articulates 

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 radius. 

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. 1–6; Fig. 10 A, B). 

Made up of nine bones, arranged in two rows – proximal 

and distal. Five bones in the proximal: scaphoideum, 

lunare, cuneiforme, pisiforme and trapezium; 

four in the distal: trapezoideum, magnum, unciforme 

and centrale. Trapezium and trapezoideum are missing 

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 15 

Carpals – carpus sin et dext 


Deinotherium thraceiensis sp.n. 


scaphoid lunare pisiforme cuneiforme magnum centrrale unciforme 

sin dext sin dext sin dext sin dext sin dext sin dext sin dext 

1. Length 187.0 188.0 166.0 160.0 206.0 206.0 162.0 160.0 110.0 110.0 115.0 115.0 162.0 160.0 

2. Width at proximal end 104.0 102.0 

3. Width of the facet for the radius 110.0 113.0 115.0 140.0 

4. Width of the facet for McI 106.0 104.0 

5. Height of the facet for the radius 120.0 140.0 

6. Height of the tubercle 114.0 150.0 

7. Lateral height 77.0 78.0 77.0 78.0 

8. Anterior width 86.0 86.0 

9. Length of the facet for the ulna 45.0 43.0 

10. Length of the facet for the cuneiform 66.0 65.0 

11. Width of the facet for the cuneiform 150.0 150.0 

12. Length of the processus 150.0 150.0 

13. Length of proximal articular surface 140.0 140.0 

14. Width of proximal articular surface 142.0 140.0 

15. Width of the bone 100.0 102.0 98.0 97.0 100.0 100.0 

16. Length of the facet for os lunare 160.0 160.0 

17. Width of the facet for os lunare 98.0 98.0 

Table 16 

Metacarpals 



Mc-I Mc-II Mc-III Mc-IV Mc-V 


1. Length 226.0 225.0 230.0 230.0 248.0 250.0 232.0 230.0 205.0 205.0 

2. Width at proximal end 140.0 140.0 125.0 125.0 112.0 112.0 98.0 108.0 146.0 145.0 

3. Width at distal end 115.0 115.0 108.0 125.0 114.0 113.0 109.0 98.0 97.0 97.0 

4. Width at distal sinew processes - - 104.0 108.0 105.0 106.0 103.5 103.5 114.0 113.0 

5. Length of small facet - - 126.0 108.0 135.0 135.0 115.0 115.0 140.0 140.0 

6. Length of large facet - - 110.0 126.0 136.0 138.0 - - 112.0 140.0 

33

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 articulates 

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 facets 

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 processus 

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 anterior 

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 cuneiform 

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, elongated 

anteriorly. Situated in the middle of the distal 

row, between os unciforme and os centrale. Two facets 

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, elongated 

posteriorly. The proximal acet for the cuneiform 

is large and medially inclined. Two facets in 

distal end, medial larger and contacting McII, lateral 

– McIII. 

Centrale (Fig. 10 A, B No. 9). Although slightly elongated 

in the ends, it also has a square shape. Situated 

between the magnum and the scaphoid. Distal 

surface slightly concave, entirely lying on the proximal 

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 preserved 

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 articular 

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 posteriorly, 

gradually becoming narrower. Facet for unciform 

in the proximal end long and wide. Laterally to it 

34 

a facet for McIII, smaller than the first. Slightly concave 

diaphysis, distal end widening with a large articular 

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 diaphysis, 

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 middle 

and in the distal part – concave. Large articular 

surface for the second phalanx on the posterior part. 

Slightly rounded in front. Measurements of the digits 

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 phalanxes 

are similar in D. giganteum Kaup, D. gigantissimum 

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 anterior 

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

Table 17 

Bones of the digits – forelimb 


1-st digit 2-nd digit 3-rd digit 


1 2 3 1 2 3 1 2 3 

sin dext sin dext sin dext sin dext sin dext sin dext sin dext sin dext sin dext 

1. Length 110 110 100 100 98 96 150 150 80 80 130 130 178 180 105 102 167 167 

2. Width at proximal end 125 125 91 91 66 68 104 105 106 107 95 95.5 115 116 108 108 111 112 

3. Width at distal end 88 88 82.6 83 - - 94 95 97 97 - - 107 107 102 103 - - 

Table 17 

Bones of the digits – forelimb (continued) 


4-th digit 5-th digit 


1 2 3 1 2 3 


1. Length 178 180 101 102 120.5 123 140 140 104 102 116 117.5 

2. Width at proximal end 110 111 96 98 87 88 95 96 96 96 96 97.5 

3. Width at distal end 96 96 90 89 - - 90 90 80 80 - - 

Table 18 

Pelvis 

D. thraceiensis sp.n. D. gigantissimum D. giganteum Kaup D. giganteum Kaup D. giganteum, Krivoi 


Ezerovo, Bulgaria St. Manzati, Romania – München – München – Stromer, Rog, Moldova – 

Stefanescu 

Stromer, 1938 

1938 

Белокрые, 1960 

sin dext 

1. Length of wing d-c 1110.0 1165.0 1130.0 1000.0 780.0 

2. Total length of the wing h-d 1160.0 1175.0 1180.0 - - 

3. Length of wing to for. obturatum 820.0 740.0 900.0 830.0 560.0 

4. Length of for. obturatum 266.0 280.0 - - - 245.0 

5. Width of for. obturatum 126.0 150.0 - - - 

6. Diameter at the acetabulum 190.0 188.0 - - - 230.0 

7. Height of sacral process 320.0 320.0 - - - 250.0 

8. Length of pelvic symphysis 462.0 460.0 - 470.0 340.0 

9. Height of same 385.0 275.0 - - - - 

35

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. Foramen 

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 parallel 

to the bone’s 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 intercondylaris 

which separates them is wide and deep. 

Fossa muscularis cranialis and F. m. caudalis deep 

and parallel to the condyles. Longitudinal crests surrounding 

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 hemisphere 

surrounded by a deep and wide ring. Flat- 

Table 19 

Femur 

36 




sin dext 

D. gigantissimum St. 

Manzati, Romania – 

Stefanescu 


München – 

Stromer, 1938 


München – 

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 20 

Tibia 


D. thraceiensis sp.n. Ezerovo, 

Bulgaria 

sin dext 



Stefanescu 


Pikermi – Gaudry, 1862 

1. Total length 1200.0 1250.0 920.0 950.0 


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 

Table 21 

Fibula 




sin dext 

tened posteriorly. The larger part of the bone was 

covered by a cartilage with the shape of the thighbone’s 

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, we’ll discuss only its distal 

part which is original. Down of the diaphysis the 

bone becomes triangular. It gradually widens, forming 

at the distal end the cochlea tibiae. Maleolus 

medialis strongly developed, incisura fibularis deep 

and wide. 

Fibula. The original fragments used for the restoration 

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 taxonomical 

value in the hipbones and the bones of the lower 

leg. 



Stefanescu 


Pikermi – Gaudry, 1862 

1. Total length 1125.0 565.0 830.0 870.0 


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 22 

Tarsals – tarsus sin et dext 

Deinotherium traceiensis sp.n. 

№ Dimensions, mm Astragalus Calcaneus Naviculare 

Cuneiforme 

mediale 

Cuneiforme 

intermedium 

Cuneiforme 

laterale 


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 

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 after 

D. gigantissimum Stefanescu. The toes were restored, 

too. 

Comparison. There are no differences in the structure 

of the tarsals in the various deinothere species. 

There are such however in their digits. In D. thraceiensis 

sp. n. the size of the first and second phalanxes 

gradually decreases. In D. gigantissimum Stefanescu 

all digits have large first phalanxes, the second 

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- 

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 forelimbs. 

In the compared species they are much smaller. 

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. 

37

38 

Table 23 

Metatarsals – metatarsus sin et dext 



Mt-I Mt-II Mt-III Mt-IV Mt-V 


1. Length 212.0 212.0 220.0 220.0 225.0 225.0 238.0 240.0 222.0 222.0 

2. Width at proximal end 137.0 138.0 114.0 114.0 120.0 125.0 120.0 120.0 146.0 146.0 

3. Width at distal end 120.0 120.0 115.0 115.0 115.0 115.0 114.0 115.0 116.0 116.0 

4. Width at distal sinew processes 135.0 136.0 112.0 112.0 102.0 103.0 112.0 112.0 117.0 117.0 

5. Length of small facet 131.0 130.0 - - - - - - - - 

6. Length of large facet 126.0 126.0 - - - - - - - - 

Table 24 

Bones of the digits – hindlimb – Phalangi sin et dext 


1-st digit 2-nd digit 3-rd digit 


1 2 3 1 2 3 1 2 3 

sin dext sin dext sin dext sin dext sin dext sin dext sin dext sin dext sin dext 

1. Length 136 136 107 107 130 130 160 160 100 100 142 142 180 180 125 125 165 167 

2. Width at proximal end 12 122 105 105 94.5 94 112 112 104 105 98 98 117 116 110 111 107 107 

3. Width at distal end 100 100 92 92 - - 101 101.5 87 87.5 - - 104 104 95 94 - - 

Table 24 A 

Bones of the digits – hindlimb – Phalangi sin et dext 


4-th digit 5-th digit 


1 2 3 1 2 3 


1. Length 169 170 105 105 125 125 146 147 104 104 103 103 

2. Width at proximal end 117 117 110 110 87.5 89 110 111.5 103 104 85 86 

3. Width at distal end 106 108 93 94 - - 105 103 92 92 - -

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 

References 

Atanasiu, S. 1907. Contributiuni la studiul faunei tertiare de 

mamifere din Romania. – Anuar. Inst. Geol. a Rom. Mol. 

I. Fasc. 1; 129—214, Taf. I—XII. 

Augusta, J. 1942. Swieta praebjeta. Praga; 43—49. 

Biber, V. 1884. Ein Dinotherium-Skelet aus dem Eger-Franzesbader 

Tertiärbecken. – Verh. d. k.k. Geol. Reichs.; 

299-305. 

Bleinville, H. 1837—1854. Osteographie ou description econographique 

comparee du squelette et du systeme dentaire des 

Mammiferes. 

Cuvier, G. 1836. Recherches sur les ossements fossiles. 4-eme 

Ed. Text et Atlas. Paris. 

Dehm, R. 1936. Paläontologische und Geologische Untersuchungen 

im Tertiär von Pakistan. Dinotherium in der Chinji 

Stufe der Uneren Siwalik Schichten. – Bayer. Akad. d. Wissen. 

Math. Naturw. Kl. Abh. N. F., 4; 1—34, Taf. I—II. 

Dehm, R. 1949. Das jüngere Tertiär in Südbayern als Laderstätte 

von Sängetieren, besonders Dinotherien. – N. Jahr. f. 

Min. ets. Abh. Bd. B; 1—3. Taf. I—II. 

Deperet, Ch. 1890. La faune de Mammiferes miocenes de la 

Grivesaint-Alban (Isere) et de quelques autres localites du 

Bassin du Rhone. – N. Jahr. f. Min. Abh., Bd 5; 1—93, Taf. 

I—IV. 

Deperet, Ch. 1887. Recherches sur la succession des Faunes de 

Vertebres miocenes de la vallee du Rhone. – Arch. d. Mus. 

d’Hist. Nat. Lyon, 4; 45—313. pls. XIV—XVIII. 

Dietrich, W. O. 1916. Über die Hand und den Fuss von Deinotherium. 

– Zeitschr. d. Deutsch. Geol. Ges., 68; 44—53. 

Ehik, J. 1930. Prodinotherium hungaricum n. g. n. sp., with an 

appendix by Szalai T. Geologica Hungar. Ser. Facs. 6; 1—24. 

Gaudry, A. 1862. Animaux fossiles de l’Attique. 

Gaudry, A. 1873. Animaux fossiles du Mont Leberon. 

Gervais, P. 1848. Zoologie et Paleontologie Francaises. 

Gervais, P. 1867/69. Zoologie et Paleontologie generales. 

Jaquemim 1837. Memoire sur les Pachydermes fossiles. – 

Magazin de Zoologie; 18—20. 

Lartet, E. 1858—59. Sur la dentition des Proboscidiens fossiles 

(Dinotherium, Mastodonta, et Elephas) et sur la discribbution 

geographique et stratigraphique de leur debris en Europe. 

– Soc. Geol. d. Fr. ser. z. t 16; 477—482. 

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 Bucharest 

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 skeleton. 

The neck is thick, merging with the vertebral 

column because, according to them, the last cervicals 

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 mandible 

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 suppose 

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. 

Lattet, E., K. Chantre. 1876. Recherches sur les Mastodontes 

et les Faunes mammalogiques qui les accompagnent. Arch. 

d. mus. d’Hist. Nat.1 Lyon, 2; 285—311, Pls. I—X. 

Lydekker, R. 1876/80/84. Indian tertiary and post-tertiary Vertebrata. 

– Mem. Geol. Surv. India, Pal. Ind. ser. 10, 1—3. 

Nikolov, I. 1985. Catalogue of the localities of Tertiary Mammals 

in Bulgaria. – Paleontol., strat. and lithol., 21; 43—62. 

Osborn, H. F. 1936. Proboscidea – a monograph of the discovery 

evolution migration and exinction of the Mastodonts, 

Elephants of the world. I – Moeritheriodea, Deinotherioidea, 

Mastodontoidea. 

Owen, R. 1840-45. Odontography or a treatise of the comparative 

anatomy of teeth. 

Schlosser, M. 1921. Die Hipparionenfaunen von Weles in Mazedonien. 

– Abh. Bayer. Ak. Wiss. Math-phys. Kl. Bd. 29, 4; 1—50. 

Stefanescu, Gr. 1891. On the existence of the Deinotherium in 

Romania. – Bull. Geol. Soc., 2; 1—10. 

Stefanescu, Gr. 1910. Dinotherium gigantissimum. – An. Mus 

d. Geol. Und Paleont., 4; 1—46, Tabl. I—X. 

Strömer, Er. 1938. Huftier Reste aus dem unterstpliocänen 

Flintdsande München. – Abh. d. Bayer. Akad. d. Wiss. 

Mothnat. wiss Kl. N F, 44; 1—40, Taf. I—II. 

Tobien, H. 1962. Über Carpus und Tarsus von Deinotherium 

giganteum Kaup (Mamm. Proboscides). – Pal. Zeitschr. 

H. Schmidt-Festband; 231—238. 

Vacek, M. 1882. Uber neue de von Dinotherium in Wiener Becken. 

– Verh. k. k. geol. Reichs.; 1—17 . 

Wagner, A. 1850 (1848). Urweltliche Säugetierüberreste aus 

Griechenland. – Abh. Nat.-phis. Kl. Bayr. Ak. Wiss., Bd. 5. 

Áàêàëîâ, Ï. 1911—1913. Ïðèíîñè êúì ïàëåîíòîëîãèÿòà íà 

Áúëãàðèÿ. II. Dinotherium’îâè îñòàíêè â Áúëãàðèÿ. – 

Ãîä. ÑÓ, 8—9; 1—29, òàáë. I—VIII. 

Áàêàëîâ, Ï. 1949—1950. Íÿêîëêî íîâè íàõîäêè îò Dinotherium 

â Áúëãàðèÿ. – Ãîä. ÑÓ, Ïðèðîä.-ìàòåì. ôàê., 46, 3. 

Áàêàëîâ, Ï., Èâ. Íèêîëîâ. 1962. Ôîñèëèòå íà Áúëãàðèÿ. Õ 

Òåðöèåðíè áîçàéíèöè. Ñ., Èçä. ÁÀÍ; 162 ñ. 

Áåëîêðûñ, Ë. Ñ. 1960. Ê ñèñòåìàòèêå è ôèëîãåíèè äèíîòåðèåâ 

(â ñâÿçè ñ íîâîé íàõîäêîé äèíîòåðèÿ â âåðõíåñàðìàòñêèõ 

îòëîæåíèÿõ Êðèâîãî ðîãà). – Ïàëåîíò. Æóðíàëü, 

4; 95—103. 

39

Áîí÷åâ, Ã. 1897. Ðàçêîïêè îêîëî Ìåñåìâðèÿ. – Ãîä. Áúëã. 

ïðèð. ä-âî, 2; 49—50. 

Áîí÷åâ, Ã. 1900. Âúðõó òåðöèåðíàòà ôàóíà îò Ìåñåìâðèÿ. 

– Ãîä. ïðèðîäîèçï. ä-âî, 3; 113—115. 

Áðúíêèí, K., Ñòàí÷åâà, Ì. 1965. Çà ïðèñúñòâèåòî íà ìèîöåíñêè 

îòëîæåíèÿ â Þæíà Áúëãàðèÿ. – Ñïèñ. Áúëã. 

ãåîë. ä-âî, 26, 1; 106—108. 

Äàâèä, À. È. 1987. Âûìåðøèå õîáîòíûå Ìîëäîâèè. Àêàä. 

Íàóê Ìîëäîâñêîé ÑÑÐ, È-ò çîîëîãèè è ôèçèîëîãèè 

„Øòèíöà“, Êèøèíåâ. 

Èâàíîâ, Ñò. 1960. Àíàòîìèÿ íà äîìàøíèòå æèâîòíè. ×àñò 

I. Ñîôèÿ, 1—368. 

Êîâà÷åâ, Ä. 1966. Åçåðîâñêèÿò äåéíîòåðèóì. – Ïðèðîäà è 

çíàíèå, 19, 3; 18—19. 

Ëåòðîíèjåâèh, Æ. Ï. 1954. Íîâè íàëàñöè Dinotherium giganteum 

Kaup ó Ñðáèjè. – Ãåîë. Àí. Áàëê. ï-â, 22; 99—113, 

òàáë. I—IV. 

Ìîðîçàê, Í. 1936. Äèíîòåðèóì â Áåñàðàáèÿ. – Ãîä. Óíèâ., 

22; 256—279. 

Íèêîëîâ, È. 1960. Ïëèîöåíñêà áîçàéíà ôàóíà îò Õðàáúðñêèÿ 

ëèãíèòåí áàñåéí. – Òðóä. ãåîë. Áúëã., ñåð. 

ïàëåîíò., 2; 295—314, òàáë. I—IÕ. 

Íèêîëîâ, È. 1962. Íÿêîëêî íîâè íàõîäêè íà ïëèîöåíñêà 

áîçàéíà ôàóíà îò Áúëãàðèÿ. – Òðóä. ãåîë. Á-ÿ, ñåð. 

ïàëåîíò., 4; 182—202, òàáë. I—IÕ. 

Íèêîëîâ, È. 1969. Äîêëàä íà òåìà „Ðåñòàâðèðàíå ñêåëåòà 

íà äåéíîòåðèóìà îò ñ. Åçåðîâî“. – ÃÈ ïðè ÁÀÍ, ÌÕÌ 

¹ 181; 1—12, IÕ òàáë. 

Íèêîëîâ, È., Êîâà÷åâ, Ä. 1966. Ïëèîöåíñêà áîçàéíà ôàóíà 

40 

îò Àñåíîâãðàä. – Òðóä. ãåîë. Á-ÿ, ñåð. ïàëåîíò., 8; 

131—142, òàáë. 1. 

Ïàâëîâà, Ì. 1907. Dinotherium giganteum Kaup èç îêðåñíîñòåì 

ã. Òèðàñïîëÿ. – Åæåã. ïî Ãåîë. è Ìèíåð. Ðîññèè, 

9; 21—22. 

Ïàâëîâèh, Ì. 1969. Ìèîöåíñêèõ ñèñàðè Òîïëè÷êîå êîòëèíå 

– ïàëåîíòîëîøêî-ñòðàòèãðàôñêè ñòóäjà. – Ãåîë. 

Àí. Áàëê. ï-â, 34; 269—394. 

Ïàâëîâèh, Ì., Â. ×èëîâèh. 1964. Î ôîñèëíèõ ñèñàðèêà íà 

òåðöèåíîã áàñåéíà Êðèâà ðåêà (Ñðájà). – Ãåîë. Àí. 

Áàëê. ï-â, 31; 145—160. 

Ïàâëîâèh, Í. 1967. Çíà÷àj ôîñèëíèõ ñèñàðà çà ñòðàòèãðàôñêî 

ðàç÷ëàíàâàíå íåîãåíèõ òâîðåâèíà òîïëè÷êîã 

áàñåéíà. – Ãåîë. Àí. Áàëê. ï-â, 33; 127—138. 

Ïåòðîíèåâèh, Æ. Ì. 1956. Ôîñèëíè îñòàíöè ñóðåàëêà èç 

îêîëèíå Êðàëüåâà è íèõîâ ñòðàòèãðàôñêè çíà÷àj. – 

Ãåîë. Àí. Áàëê. ï-â, 24; 185—198, òàáë. I—VI. 

Ðîäåâ, Ö. 1966. Äåéíîòåðèóìà ïðè ñ. Åçåðîâî. – Ñï. Êîñìîñ. 

Ñâèñòóí, Â. È. 1974. Äèíîòåðèè Óêðàèíû. Àêàä. Íàóê 

Óêðàèíñêîé ÑÑÐ, È-ò çîîëîãèè, Êèåâ, „Íàóêîâà äóìêà“. 

Ñòåâàíîâèh, Ï. Ì. 1951. Äîíè ïëèîöåí Ñðáèjå è ñóñåäíèõ 

îáëàñòè. – Ñðïñêè Àêàä. íàóê, 2; 1—361, òàáë. ÕVIII. 

Òàðàáóêèí, Á. À. 1968. Î ðàñêîïêàõ ñêåëåòà äåéíîòåðèÿ â 

Ðåçèíñêîé ðàéîíà Ìîëäàâñêîé ÑÑÐ. – Èçâ. Àêàä. Ìîëä. 

ÑÑÐ, ñåð. áèîë. õèì., 3; 37—42. 

Öàíêîâ, Â., Íèêîëîâ, È. 1966. Äåéíîòåðèóìúò ïðè ñ. Åçåðîâî 

Àñåíîâãðàäñêî. – Ïðèðîäà è çíàíèå, 4; 3—6.

Deinotherium thraceiensis sp. nov. from the Miocene near Ezerovo ...

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?