Deinotherium Thraceiensis Sp

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

Deinotherium thraceiensis sp. nov. from the Miocene near Ezerovo, Plovdiv District

Dimitar Kovachev1, Ivan Nikolov2

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)

Ä. Êîâà÷åâ, È. Íèêîëîâ – Deinotherium thraceiensis sp. nov. èç ìèîöåíñêèõ îòëîæåíèé áëèçè äåðåâíè Åçåðîâî, Ïëîâäèâñêàÿ îáëàñòü. – Íàñòîÿùàÿ ñòàòüÿ ïðåäîñòàâëÿåò ìåæäó- íàðîäíîìó ïàëåîíòîëîãè÷åñêîìó îáùåñòâó îïèñàíèå ïîëíîãî ñêåëåòà äåéíîòåðèÿ, îáíà- ðóæåííîãî è âûêîïàííîãî Ä. Êîâà÷åâûì â 1965 ãîäó èç íåîãåíà âáëèçè äåðåâíè Åçåðîâî, è îïèñàííîãî, ðåñòàâðèðîâàííîãî è ìîíòèðîâàííîãî ïîêîéíèì Èâàíîì Íèêîëîâûì. Ôîñ- ñèëüíûå îñòàíêè íàéäåííûå â ñåäèìåíòàõ ìýîòñêîãî âîçðàñòà. Äî ýòîãî, â òåõ-æå îòëîæå- íèé 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 ñèì- ìåòðè÷åñêèõ àíàëîãîâ. Ðàçìåðû êîñòåé è ñðàâíåíèå ñ äðóãèìè äåéíîòåðèÿìè îïèñàííûìè â ëèòåðàòóðå ïîêàçàëî íåêîòîðûå ñóùåñòâåííûå ðàçëè÷èÿ. Íåêîòîðûå ðàçëè÷èÿ ïî îòíî- øåíèè ÷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. Ko- vachev in 1965, and described, restored and mounted by the late Ivan Nikolov. The fossil remains are found in sediments of Maeotian age. Former excavations have yielded fossil bones of Trilo- phodon angustidens Cuvier, Hipparion microtaton Nicolov and parts of mandible of Mastodon sp. When the whole skeleton was excavated only a few bones were missing, and namely, the whole rear left thighbone, the right rear fibula, most of the ribs as well as some caudal vertebrae. They are restored on the basis of symmetrical analogues existing. The measurements on the bones found and the comparisons with other deinotheres described in the literature showed certain differences. Some of the differences relative to the skull have certainly a taxonomic value. There- fore, the deinothere described is referred to a new species, Deinotherium thraceiensis sp. n.

Kovachev, D., Nikolov, I. 2006. Deinotherium thraceiensis sp. nov. from the Miocene near Ezerovo, Plovdiv District. – Geologica Balc., 35, 3—4; 5—40. Key words: Deinotherium thraceiensis, Miocene, Maeotian, Bulgaria.

Introduction tute, Bulgarian Academy of Sciences, that the restoration process was finished. The assembled skeleton The fossil materials presented herewith to the scien- has been mounted and permanently exhibited in the tific community were found by Dimitar Kovachev in Museum of Geology and Palaeontology, Sofia Uni- 1965. The late Ivan Nikolov has worked for a long versity “St. Kliment Ohridski”. A diminished (3/4 of time on their restoration, with minor participation of normal size) copy has been prepared and is now on D. Kovachev. In 1972 Nikolov reported to the De- exhibit at the Asenovgrad Palaeontological Branch partment of Palaeontology at the Geological Insti- of the National Natural History Museum. Both Ivan

5 Nikolov and Dimitar Kovachev were fully aware of scribed further remains of that genus found near the importance of this very rare case of finding a Eppelsheim and Westhofen. According to him, the whole skeleton of such a huge extinct animal, and adult animals had five teeth in their upper and low- they estimated it to be a representative of a new spe- er jaws. In 1833 he described the new species D. cu- cies of genus Deinotherium. Ivan Nikolov, well-known vieri, but because it was often found in Bavaria, H. v. for his thoroughness, wanted to make an exemplary Meyer called it D. bavaricum. This name is still used publication but his early death stopped for a long today. However, arguments about the exact place and time the work on it. The results of his enormous la- the life environment of deinotheres still continued. bour put into the description of the bones, the resto- P. Gervais accepted in 1848 the opinion of Kenedy ration and mounting of this unique skeleton, re- and Koch that the genera Deinotherium and Masto- mained in the archives of the Geological Institute. don should be referred to Proboscidea. He recognized After the sudden untimely death of Nikolov in 1984, the existence of three species: D. giganteum Kaup = the work on the publication remained unfinished. Tapir gigantesque Cuvier; D. intermedium and D. cu- Now, D. Kovachev decided to prepare the materials vieri Kaup. S. Pictet reported in 1853 about the finding for publication with the clear idea about the respon- near the village of Absdorf of a whole skeleton of D. sibility undertaken, and that the whole blame for even- giganteum Kaup. Only the head, the first two vertebrae, tual flaws and errors should be addressed to himself. the corpus of a thoracic and a caudal vertebra and Such publication would be of importance for the fragments of the limbs were preserved. These remains future studies on genus Deinotherium. showed that it is close to the mastodon and belongs to Many scientists have helped the authors in dif- Pachydermae. Four deinothere species were recognized ferent time. Prof. Dr. R. Dehm (Director of Univer- in 1858—1859 by M. Lartet, and namely, Deinotherium sität-Institut für Paläontologie und historische Geolo- bavaricum H. v. Meyer, D. giganteum Kaup, D. sp. (an gie – München) and Prof. Dr. H. Tobien (Director intermediary form between the first two), and D. cu- of Paläontologisches Institut der Johanes-Gutenberg vieri Kaup. The first species is known from the Mi- Universität – Meinz) helped I. Nikolov during the ocene of France and Bavaria, the second from the Late first studies and restoration. Academician T. Nikolov Miocene of France, Germany and Greece, the third (Geological Institute, Bulgarian Academy of Scienc- and fourth from the Miocene of France. After Lartet, es) gave valuable advice with the material. D. Ko- the dental formula of genus Deinotherium is: I 0-0/1-1; vachev wishes to express his gratitude to Dr. Marin Pm 2-2/2-2 and M 3-3/3-3 for the permanent teeth, Ivanov from the Sofia University for his responsive- and for the deciduous – I 0-0/1-1; deciduous molars ness to the problems of this research. Dr. N. Spassov 3-3/3-3. New fossil finds in Greece (Pikermi), India (Si- (National Natural History Museum, Sofia) kindly walik), Europe (France, Switzerland, Austria), etc. have helped with the literature I. Nikolov had used at the been reported during the last quarter of the 19th centu- library of the Museum. ry by A. Gaudry, H. Hensel, B. A. Lydekker, E. Chantre, M. Vacek, V. Biber, C. Deperet, and other authors. Gr. Stefanescu first reported in 1891, and then described Studies on the genus Deinotherium – (1895–1910) a deinothere skeleton found near Man- a historical review zati, Romania. Because of the large dimensions of all its bones and teeth, he named it D. gigantissimum. Ehik As early as the 17 century a place near Lyon, France, (1930) published upper and lower molars and some was known as “the field of the giants” because of the finger bones, found in the Pliocene sediments of Hun- large animal bones often found there. Some of those gary, as a new species – Prodinotherium hungaricum. bones came to Matsorier – a surgeon, who used to Osborn (1936) changed the name of the genus (the show them for years in France and Germany as the Latinized form Dinotherium) to the ancient Greek bones of king Töteboch. Much later, the real tomb of form Deinotherium. He revised all then known the king was found, and the deceit was exposed. The deinothere materials, discussing in details each spe- huge bones were transferred to the Natural History cies, and illustrating skulls/mandibles and teeth from Museum in Paris. Probably these are the first Deinothe- D. giganteum Kaup, D. bavaricum H. v. Meyer, D. rium remains found. cuvieri Kaup, D. pentapotamiae Lydekker, D. indi- One century later, in 1715, Réaumure admitted cum Falconer, etc. (Fig. 1). he could not refer the bones to any known animal. Numerous new finds of deinotheres have been re- Kenedy supposed in 1775 that the materials were reported during the whole 20th century from Austria, lated to mammoths. G. Cuvier thought in 1779—1836 Bohemia, Bulgaria, France, Germany, Greece, Hun- that the animal had been a large tapir coexisting with gary, Macedonia, Moldova, Pakistan, Romania, Rus- mastodons. He called it Tapir gigantesque and sia, Serbia, Ukraina, etc. thought that its tusks were curved upwards. Kaup created in 1829 the genus Deinotherium with the species D. giganteum upon the skull and mandi- Studies on the genus Deinotherium ble found near Eppelsheim, Germany. He thought in Bulgaria this was an intermediary form between the sloth and the mastodons and referred them to the larger taxo- The first Deinotherium remains in Bulgaria have been nomic unit Curtognati. In 1841 and 1857 he de- found by G. Bonchev (Áîí÷åâ, 1897) who made ex-

6 Fig. 1. Distribution of genus Deinotherium in Southeastern Europe and Asia Minor

cavations in the Middle Sarmatian limestones near other single teeth from the village Sovolyano, near Nessebar, studying the hipparion fauna. Three years Pleven, the village Archar and the village Mazgosh. later he reported new finds from the same locality. They were all brought to the Institute of Geology by Áàêàëîâ (1911/1913) published the two deinothere different people. From these materials Bakalov de- mandibles excavated by G. Bonchev, together with termined two species – D. giganteum Kaup race mi-

7 nor and D. giganteum Kaup race major. To the first Distribution of genus Deinotherium he referred the smaller teeth, and to the second – in Southeast Europe and parts of Asia the larger ones. Later on (Áàêàëîâ, 1949/1950) Bakalov described Deinotherium teeth found at the Minor “Meander of the Konska river” near the village No- evtsi; the village (now – a town; for some time called Until now, more than 80 localities (partly shown on Batanovtsi) Temelkovo (Pernik region) and Aksako- Fig. 1) within the region have yielded deinothere re- vo, near Varna, as D. giganteum Kaup race major mains, and their number is constantly growing. (from the first two localities, Pliocene) and D. gigan- There are 16 localities from former Yugoslavia: teum Kaup race minor (from the Sarmatian lime- 1. Veles, Mokranje, Smederevska palanka, Resnik, stones near Aksakovo). Íèêîëîâ (1960) reported a Meduhana, Vracevic, Duboko, Tsiganski potok, and molar fragment of D. giganteum Kaup race major between the villages Viteževo and Porodin – D. gi- from the lignite coals near the village Hrabarsko (So- ganteum Kaup. Near Veles and Ravanica, together fia region), assuming a Pontian age. The whole fossil with D. giganteum Kaup, D. bavaricum H. v. Meyer material belonging to genus Deinotherium then known is also present. The sediments near the villages Medu- from Bulgaria has been published by Áàêàëîâ, hana, Ravanica and those between Viteževo and Íèêîëîâ (1962). They recognize two races – minor Porodin are of Lower Sarmatian, and the rest – of and major, and refer the finds as follows: two man- Pontian age. dibles, an upper jaw and three separate teeth to race 2. Drenovica, Chukovica, Bresnica, Prebrezka, Zdar- minor, and a mandible, seven complete upper and ski and Kriva reka – D. bavaricum H. v. Meyer. The lower teeth and three tooth fragments to race major. age cited is Helvetian-Tortonian, Lower Sarmatian, The age of both races varies from Sarmatian to Le- Sarmatian and Middle Miocene. vantian. Íèêîëîâ (1962) referred the molar and frag- In Greece D. giganteum Kaup is known from four ment of a scapula found in the Levantian sediments localities: Pikermi, Thessaloniki and Tanagra (Pon- near the village Popitsa, Vratsa region to D. gigan- tian), and Samos (Middle Pontian). From the Sar- teum Kaup. Íèêîëîâ, Êîâà÷åâ (1966) described a matian of Chios D. bavaricum H. v. Meyer has been lower molar found by the latter author in the sands described. in the valley of Cherkezitsa river, near the village From Turkey (Asia Minor), D. giganteum Kaup Ahmatovo. They assume the age to be Levantian be- has been described from Küçükçekmece (Maeotian). cause of the Anancus arvernensis Croizet et Jobert From the territory of Romania and part of Bessa- teeth found there, referring the find to D. giganteum rabia, two taxa are known – D. giganteum Kaup from Kaup. Êîâà÷åâ (1966) reported the finding of an the Pontian and Dacian sediments near Vernesti, almost complete skeleton of Deinotherium near the Culm, Lichtental, Pelineu, Telenesti, Visterniceni, village Ezerovo, Plovdiv region. This same skeleton is Baimaclia, Cimislia, Comanesti, Curtea and Solcus- the subject of the present paper. Öàíêîâ, Íèêîëîâ tria; and D. gigantissimum Stefanescu, – from the Plio- (1966) published a brief review of the proboscideans cene sediments near Manzati, Breaza and Cimislia. in Bulgaria, and gave also some preliminary infor- From the territory of Moldova and Bessarabia, two mation about the finding of Kovachev reporting taxa are known too: 1. D. giganteum Kaup near Tara- metrical data on some bones. klia, Ciobruciu, Sofijevka, Tanovka, Chernevo, Ti-

PLATE I → 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 Deinotherium thraceiensis sp. n. 1. Skull and mandible. Scale ca. 2.9% coll. SU.M. No. SU Pl 312/1, SU Pl 312/2 2. Skull from behind – os occipitale. Scale ca 5.6% coll. SU.M. No. SU Pl 312/1; Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE IV Deinotherium thraceiensis sp. n. 1. Upper cheek teeth – P3 to M3 sin et dext. Scale ca. 25% coll. SU.M. No. SU Pl 312/1; Locality: Ezerovo, near Plovdiv; Level: Maeotian

8 PLATE I PLATE II

PLATE III PLATE IV PLATE V

PLATE VI PLATE VII

PLATE VIII PLATE IX PLATE X PLATE XI

PLATE XII PLATE XIII PLATE XIV PLATE XV

PLATE XVI PLATE XVII PLATE XVIII PLATE XIX

PLATE XX

PLATE XXI 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 sedi- has been assumed after the age of the sediments in ments are of Maeotian, even Upper Sarmatian age. the noted areas. 2. D. gigantissimum Stefanescu near Volchinec, Rah- The vertical distribution of deinotheres in Bul- ny-Lessovye and Pripecheni. The host sediments are garia starts in the Sarmatian, includes the Maeotian also of Maeotian and Upper Sarmatian age. and Pontian and ends in the Pliocene. The remains A considerable number of deinothere localities of D. bavaricum H. v. Meyer are found only in Sar- are known from the territory of Bulgaria (Fig. 2). matian sediments, while D. giganteum Kaup is present Some of the first Deinotherium finds in Bulgaria from the Middle Sarmatian up to the Romanian. 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 Stratigraphic notes on the sediments part is better studied. Some of the more important near Ezerovo, Plovdiv region finds come from Nessebar (by G. Bonchev), No- evtsi, locality “Meander of the Konska river” (by P. The village of Ezerovo is situated ca. 30 km east of Bakalov), and the village Ezerovo, “Kolnu dere” Plovdiv (Fig. 3). Sediments of Eocene, Miocene, (by D. Kovachev). Pliocene and Pleistocene Age crop out in the area Some of the materials have been found and (Fig. 4). brought to the specialists by common people (farm- The Upper Eocene (Priabonian) is represented by ers, workers, etc.). Therefore, the exact location has light organogenic limestones, rich of nummulites and been mentioned only approximately, and there is no corals. South of the village the dam wall is built onto stratigraphic profile. In such cases, a probable age them. The thickness exposed is about 50 m. In a bore-

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

PLATE VI Deinotherium thraceiensis sp. n.

1. Lower cheek teeth – P3 to M3. Scale ca. 25% coll. SU.M. No. SU Pl 312/2; Locality: Ezerovo, near Plovdiv; Level: Maeotian

2 Geologica Balcanica, 3-4/2006 9 Fig. 3. Position of the locality of Ezerovo in the Upper Thracian plain

Fig. 4. Stratigraphic profile of the locality Ezerovo

10 hole (No 16; Áðúíêèí, Ñòàí÷åâà, 1965) situated 2. A whole mandible with a preserved symphysis and near the village of Ezerovo, these rocks are covered M3 dext et sin of Choerolophodon sp. by massive limestones, marls, and well-cemented 3. Two maxillary fragments with M3 also belonging quartz sandstones. They are referred to the Oligocene to Choerolophodon sp. on the basis of rich foraminifer and ostracod fauna. 4. M3 sin et M2 dext – Choerolophodon sp. These rocks pass into the Lower Miocene dark gray 5. Right semimandible with M3 – Tetralophodon lon- aleurolites and marls with foraminifer and ostracod girostris Kaup. fauna similar to the Aquitanian-Burdigalian faunas 6. M3 sin – Anancus sp. reported in France (Áðúíêèí, Ñòàí÷åâà, 1965). The 7. M2 and M3 dext and M3 sin – Zygolophodon bor- maximum thickness of the Lower Miocene is about soni Hays. 180 m. 8. M3 dext – Zygolophodon borsoni Hays. The Palaeogene and Lower Miocene marine sed- 9. M3 sin Deinotherium giganteum Kaup. iments are unconformably covered by fluvial sedi- 10. Right semimandible – Microstonyx major. ments referred to the Ahmatovo Formation (Êîþì- They will be subject of other publications. äæèåâà, Äðàãîìàíîâ, 1979). They crop out at nu- The Neogene sands are covered by a layer of soil merous places in the valley of the river Cherkezitsa. 1.0 to 1.5 m thick including isolated rock fragments At the basis there are aleurolites and shales covered of different size. These soils are referred to the Pleis- by sands of light rusty colour, alternating with con- tocene. glomerate lenses. At places the sand layers are cross- bedded. Petrified trees occur in the sands. The thickness of the sand layers varies from 1.5 m to 7.5 m; Palaeontological part and that of the conglomerates is up to 1.5 m. The whole thickness of the sands has been revealed only Order Proboscidea Illiger, 1821 by drillings and is up to 515 m. Ahmatovo Forma- Suborder Deinotherioidea Osborn, 1921 tion is referred to the Maeotian, Pontian and Da- Family Deinotheriidae Bonaparte, 1845 cian regional Paratethys stages. Three macrocycles Genus Deinotherium Kaup, 1829 have been distinguished (Êîþìäæèåâà, Äðàãîìà- Type species of the genus Deinotherium giganteum íîâ, 1979). The first macrocycle is referred to the Kaup, 1829 Maeotian (possibly also upper parts of the Sarma- Deinotherium thraceiensis sp. n. tian) on the basis of Lower to Middle Turolian (“Pik- ermian”) mammal fauna. The Pontian age of the (Pl. I – XXII) lower parts of the second cycle is determined by Upper Derivatio nominis: thraceiensis from Greek “Thra- Pikermian (= Upper Turolian) fauna, and the up- ceia” – Thrace, the region of the find. per parts of the second cycle are referred to the Da- cian Regional Stage by Ruscinian fossil remains. The Holotype. As such we design all the bones of the skel- fossil locality of Deinotherium thraceiensis sp. n. be- eton – No. 312/1 to No. 312/23-30, collection of the longs to the first cycle of the Ahmatovo Formation Museum of Geology and Palaeontology of the Sofia (Nikolov, 1985, p. 58). University “St. Kliment Ohridski”. About 800 m west of the village of Ezerovo, right Locality. Light, medium-grained sands (Ahmatovo from the road to the dam, at the fork with the road Formation, first cycle) near Ezerovo, Plovdiv region, for the pumping station, in light medium-grained just at the turn for the pumping station from the sands, D. Kovachev found in 1965 the skeleton of road to the dam. Deinotherium thraceiensis sp. n. described in the present paper. It was lying at a depth of ca. 40 m Age. Maeotian. relatively to the level of borehole No 16. At the same Diagnosis. Large animals, skull short and high. Na- level, but 50 m to the east, V. Tsankov and I. Vaptzarov sal bones short, fusing at the anterior ends, curved have found in 1956 a part of a skull, determined by and slightly flattened laterally. Large external nares. P. Bakalov as Trilophodon angustidens Cuvier. In 1962 Ear area also large and with trapezoid shape. Eye St. Stoykov and I. Nikolov found 4—5 m below the sockets almost separated from the ear areas. Fore- find described here, teeth of Hipparion microtaton head high and, compared to the external nares – Nikolov and a part of a mandible of Mastodon sp. short. The occipital bone is high and wide. Symphy- They all indicate a Maeotian age. Above the sis of the mandible curved downwards and back- deinothere have been found teeth of Hipparion med- wards. Incisors of moderate length, slightly curving iterraneum Gervais and Tetralophodon longirostris outwards. Their tips end exactly beneath processus Kaup. They unequivocally prove a Pontian age. angularis. Neck short, with seven thin cervical verte- After 1965, D. Kovachev collected the following brae. First ones lack processus spinosus, in the last fossils during numerous field trips: ones it is weakly developed. Skull and mandible, rel- 1. A skull, a mandible and bones from the limbs of atively to the size of the whole skeleton, are much Mastodon sp. In the structure of its molars it resem- smaller than in the other proboscideans. Fingers high, bles T. angustidens Cuvier, but the tusks and the skull with large phalanxes – hoofs, strongly developed on are different from this species. the third finger.

11 Fig. 5. Deinotherium thraceiensis sp. n.

Description and comparison ta conchalis dorsalis is outlined. (Measurements of the external nares in Table 1). Skull. (Pl. I, fig. 1 and 2; Pl. II, fig. 1 and 2; pl. III, fig. The calvaria – os frontale and os parietale (fronto- 2; Fig. 5 – A, B, C and D). parietal part) starts against the beginning of the ear The skull described here is entirely preserved. It is area ends with os occipitale. In the middle it is slightly very short and its posterior part – high (Fig. 5). concave, and in the posterior part – strongly convex The premaxillary bones (os premaxilarae), which are (measurements in Table 1). strongly developed in Proboscidea, do not merge at Occipital bone (os occipitale). (Fig. 5 A, C, D ocs; their anterior ends and are slightly curved downwards. Pl. III, Fig. 2). High and wide, laterally slightly round- There are no upper tusks and alveoli for them in ed, under the condyles and by the parietal bone al- these bones. They are widening in front of the eye most straight and parallel. On both sides its wings sockets. All their surface is smooth, without any rug- are curved backwards, and in the region of the osity. condyles and above them it is slightly convex. In re- Nasal bones (os nasale) are short, high in their ante- lation to the forehead this bone is almost perpendic- rior part, rounded and laterally flattened. On the ularly situated, at an angle of 80° (measurements in inner side, in the region of the maxillary bones, the Table 1). nasals are slightly concave, so that together with the Occipital condyles. (Fig. 5; Pl. I, Fig. 1 and 2). premaxillary bones they have a rounded, wedge-like They are situated in the middle and in the lower part shape (Fig. 5, A). They are connected by a strong su- of the occipital bone. Their shape is irregular and ture. The nasal canal has an inner width of 70.0 mm oval. Wider and convex in their upper part. They and length to the end of the external nares 710.0 mm. surround foramen magnum – fm, which has an oval External nares (Fig. 5, ex. n., and Pl. II, figs 1 and 2) shape and is situated perpendicularly to the occipi- pear-shaped and rather large. They start almost at tal bone (measurements in Table 1). the beginning of os nasale, gradually becoming wid- Eye socket (orbita) (Fig. 5 A orb.) is preserved only er and deeper. By the beginning of the eye socket on the right side of the skull. In front it is limited by they narrow, then becoming straight and ending with the massive and laterally strongly protruding os lac- a regular, arch-shaped curve just at the beginning rimale, and in the back – by the well pronounced of the ear area. They penetrate for 70–80 mm more processus postorbitale, which is long and thin. From and end in the oscular area with the dorsal nasal below it is limited by os zygomaticum. Thus the eye opening, which is small and oval, parallel to the fore- socket is almost entirely separated from the ear area head. On the bottom, parallel to the nasal cavity, cris- (measurements in Table 1).

12 Bavaria Bavaria H.Meyer v. D.bavaricum D.gigantissimum Stefanescu Manzati Kaup Munich D.giganteum Kaup

Eppelsheim Eppelsheim D.giganteum 176-dext 176-dext 134-dext Ezerovo D.thraceiensis D.thraceiensis 350.0 324.0 330.0 430.0 300.0 300.0 430.0 330.0 324.0 350.0 3 130.0 130.0 3 550.0 3 510.0 470.0 390.0 540.0 252.0 252.0 540.0 390.0 470.0 510.0 3 805.0 550.0 380.0 630.0 385.0 385.0 - 630.0 - 380.0 360.0 - 550.0 715.0 805.0 1 3 -M 3 Dimensions, mm mm Dimensions, 1. Total length length 1. Total 2. occipitalis cond. to nasale os from Length 3. occipitale os to nasale os from Length 4. occipitale os of basis the at Width 5. (nasale) nares external the of basis the at Width 6. (nasale) nares external the of middle the in Width 7. sockets eye in Width 8. area ear in Width 9. M at skull the of Height 1290.0 570.0 680.0 1012.0 165.0 900.0 1320.0 - 840.0 - 240.0 121.0 950.0 730.0 540.0 108.0 - 170.0 1035.0 252.0 1118.0 - 570.0 1260.0 - 780.0 - 270.0 860.0 500.0 ------10. 10. M at skull the of Height 11. area ear of Length 12. socket eye of Height 13. socket eye of Width 14. socket eye of Depth 15. Length of os nasale16. to the anterior end of P premaxillaria in nasale os of Width 17. Width of os nasale18. to the anteriorof P end points two these between middle the in nasale os of Width 19. nasale os of height Minimum 20. Beginning 21. of external nares from the beginning of os nasale curve first the at nares external of Width 22. 185.0 curve second the at nares external of Width 23. Width of24. external nares at the eye socket 150.0 curve first the at nares external the of Depth 25. curve second the at nares external the of Depth 26. middle the in nares external the of Depth 27. frontale os of Length 28. 260.0 frontale os of width Minimum 380.0 29. 185.0 frontale os of width Maximum 30. 120.0 occipitale os of Length 31. 160.0 300.0 160.0 occipitale os of Height 32. occipitalis condylus of Height 154.0 165.0 250.0 360.0 33. occipitalis 530.0 condylus of width Maximum 34. 500.0 magnum foramen of height Maximum 35. magnum foramen of width Maximum 36. zygomaticus arcus of Length 37. 160.0 zygomaticus arcus of Width 38. durum palatum of depth Maximum 39. P at durum palatum of Width 500.0 40. - 540.0 Length of tooth P row 308.0 126-sin 108.0 196-sin - 850.0 155.0 700.0 - 540.0 75.0 - - 210.0 176.0 440.0 - 450.0 260.0 95-100 ------420.0 - - - 450.0 - - №

Table 1 Skulls – Crania

13 Ear area. (Fig. 5 A, C or. r and Pl. I, Fig. 2). It is high, The second molar (M2 dext) is large and almost wide and deep. Begins immediately behind the eye square. It is built by two thick and high ridges which socket. In the posteriror part it borders os occipitale. are slightly curved backwards on the outer side. Thus Maxillary bone (Maxilla) (Fig. 5 B m and Pl. I, Fig. 2). on the posterior wall of the ridge there is a small Wide, slightly concave at the palatum. Length concavity, while in front they are straight. A deep 400.0 mm, height at M1 – 250 mm. Facial crest and valley with steep walls separates them. Both teeth (left infraorbital canal not pronounced. and right) have a clear cingulum on the anterior Palatum durum. (Fig. 5 B; Pl. I, Fig. 2). Long and and the posterior sides. In the anterior part of the very narrow. Strongly concave in the middle (mea- crown the cingulum is something like an anterior surements in Table 1). talon and has a worn-out tip. Upper cheek teeth (Pl. I, Fig. 2 and Pl. IV, Fig. 1). The last molar (M3 dext) is the largest. It has two Both toothrows are entirely preserved. Only the first thick ridges, slightly convex forwardly. A deep and molars are slightly worn out, so that part of the den- wide valley separates them. The ridges are wider on tin is seen. There is no difference in the structure of the inner side and slightly curved backwards on the the left and the right ones. Their sizes differ with 1 to outer side. They form a small concavity on the poste- 3 mm (see Table 2) but this is assumed to be normal. rior side. The valley between the two ridges is free. We shall discuss only the morphological differenc- The cingulum is well developed both at the anterior es, where there are such. and the posterior part, shaping an anterior and pos- The third premolar (P3 dext) is large, of almost terior talon. 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 Comparison anterior and the posterior part of the outer ridge correspondingly. Thus the three ridges surround a deep According to the literature, there are only two com- triangular valley. The anterior inner ridge is wider, pletely preserved skulls found up till now. The first with a triangular shape, slightly worn out at the tip. has been found near Eppelsheim in 1835, described Its inner side is concave and, together with the cin- by Andrews (1921) as D. giganteum Kaup and is gulum on this side of the crown, surrounds a small stored at the British Museum. The second has been and shallow triangular valley. On the inner side it is found near the village Gussiatin, Tirasspol region, slightly more worn out. The cingulum is well devel- by a palaeontological expedition from the Institute oped on the whole anterior and the posterior outer of Zoology, Academy of Sciences of the Ukrainian part of the crown. SSR in 1963, described by Svistun in 1974 and re- The fourth premolar (P4 dext) has an almost tet- ferred by him to D. levius Jourdan. In Vienna there ragonal shape. It consists of one outer and two inner is a mounted skeleton of D. bavaricum H. v. Meyer ridges, situated perpendicularly to the outer. The but its skull is entirely reconstructed (see Pl. XXII). anterior inner ridge is wider than the posterior and Similar is the case with D. gigantissimum Stefanescu less worn out. It has the shape of a isosceles triangle, in Bucharest. From the skull of the deinothere skele- the base of which touches the inner anterior part of ton recently mounted in Kishinev only part of the the outer ridge. At this place the two ridges surround maxilla with the upper teeth is preserved. a shallow valley. The posterior ridge is narrower and In conclusion, it seems that the only possible com- less worn out. By touching the outer ridge it is divid- parisons are with the skulls from Eppelsheim and ed in two parts, and V-shaped. Thus the ridges sur- Gussiatin. The main differences between them and round a deep triangular valley with steep walls. On the skull described by us (cf. Figs. 5, 6 and 7) are as the inner and outer posterior side and on the anteri- follows: or side of the crown there is a slightly developed cin- 1. Relatively to the body size, the skull described here gulum. The left P4 has a weaker cingulum on its an- is short, but high. Compared to the other two, the terior side. Ezerovo skull is highest, and the one from Gussiatin The first molar (M1 dext) is long and narrow. It – the lowest. consists of three ridges. First two are more worn out. 2. The premaxillary bones have interesting differences The three ridges on the inner side are wider. They (Figs. 5, 6 and 7 pmx). In the skull described by us gradually become thinner toward the outer wall of they are curved downwards as in the Eppelsheim skull, the crown. At the very end they are slightly curved but are not so strongly flattened, the part not knit downwards but they don’t touch each other. They together is much smaller and the incisure between are separated by small valleys narrowing in their mid- them is narrower and shallower. In the Gussetin skull dles. The second and third ridges are slightly con- the curving is weaker. In this species, in their pre- cave on the posterior side, while the anterior side of orbital part the lateral processes are much stronger than the third ridge is more convex. A tubercle at the base in both our and the Eppelsheim skull. Besides, in the of each valley, but only from the inner side, closes Gussiatin skull the premaxillary bones are coarsely rug- them. The valleys are open outwards. In front, the ged with a tuberosity protruding forward. On it, ac- crown has a cingulum which, together with the an- cording to Svistun 1974, there could be horn-like struc- terior wall of the first ridge, surrounds a long and tures substituting for the upper tusks. There is no such shallow valley. thing in our material, neither in the Eppelsheim skull.

14 , Bakalov, raceminor Nikolov, 1962,Bulgaria Kaup D. giganteum Bakalov, Nikolov, 1962, Bulgaria racemjor

D. gigantissimum Stefanescu, 1909, Romania Romania 1909, Stefanescu,

D. thraceensis D. sin dext sin dext sin dext sin dext sin sin dext dext sin dext sin dext sin dext sin sp.n. Ezerovo, Bulgaria Width of the second ridgethe at top 69.0 71.0 base the at ridge second the of Width 110.0 - 110.0 base the at ridge second the of Width 117.0 107.2 - 107.0 114.0 - 107.0 102.0 - 90.0 104.0 84.5 - 98.0 83.0 101.0 - 59.0 89.0 - 73.0 89.0 - 33.0 - Measurements in mm 1 2 3 Width of first ridge Width of first ridge 90.5 91.8 96.3 - 87.0 96.0 - - 97.0 - 68.0 ------17.0 42.5 3 4 thickness P tooth the of Length tooth the of Width Enamel ridge second of Width thickness P tooth the of Length Width of the tooth 4.0 Enamel 4.2 ridge second of Width 100.0 - 105.0 tooth the of Length 106.5 base the at ridge first the of Width 4.2 M 100.0 Width of the first ridge at the top 4.3thickness base the - at ridge second the of Width 106.5 106.0 75.0 - 94.6 base the at ridge third the of Width - 100.8 Enamel 100.5 - 94.0 Width of the third ridge atthetop - 70.0 - Width of posterior talon - 81.5 - 100.0 - 83.5 - 110.0 102.5 98.5 tooth the of Length M 97.0 thickness base the at ridge first the of Width 4.3 - 73.0 63.5 - - Width of the first ridge at the top 4.3 100.0 - 87.0 110.0 84.5 Enamel - top the at ridge second the - of Width 113.6 - 98.0 - - - - 62.0 Width of posterior talon - - - - 119.0 88.8 tooth the of Length 93.0 85.3 86.0 M 115.0 102.0 thickness base the - at ridge first the of Width 4.7 - 107.7 100.0 92.0 - Width of the first ridge at the top 106.0 - 4.6 - - 87.5 Enamel - - 115.0 - 84.5 - Width of the second ridgethe 129.0 at top 82.0 - 118.0 - - 83.0 - 106.5 - Width of posterior talon 80.0 65.0 73.0 100.0 - - - 87.0 115.0 99.0 - 70.0 - 62.0 - - 5.1 111.0 75.0 - - 116.0 - - - 110.0 103.0 - - 5.2 98.0 - - 61.0 - - - - 114.0 89.0 92.6 - - - 103.0 - 108.0 - - 114.0 - 78.0 - - - - 98.0 ------114.0 - 115.0 53.0 - - - - - 96.0 103.0 78.0 ------92.0 - 103.0 - - - 101.0 - 107.0 - - 107.0 36.0 - 86.0 ------52.0 30.0 - - 36.0 - 62.0 - 95.0 - - 57.0 - 96.6 - 91.0 - 74.0 - - 105.0 51.0 - - 105.6 70.5 - - - - 70.0 - 89.0 - - 50.0 48.5 - 50.0 - - - 29.5 - - 71.0 96.0 - - - - - 31.0 44.0 - - - 38.0 - - - 52.0 51.5 ------Table 2 Upper cheek-teeth

15 Fig. 6. Deinotherium levius Jourdan

Fig. 7. Deinotherium giganteum Kaup

3. The external nares (Figs. 5, 6, and 7 – exn) in D. In the Gussiatin species it is even strongly concave thraceiensis sp. n. is large, deep and pear-shaped. It in its anterior part. starts from the very beginning of the nasal bones, 6. The eye socket (Figs. 5, 6, and 7 – orb) in the gradually widening and taking two thirds of the Ezerovo skull is large and separated from the ear length of the whole skull. In the other species it is area. In D. giganteum Kaup they are almost fused, shorter and almost of the same width in its anterior and in D. levius Jourdan from Gussiatin the situa- and posterior parts. tion is closer to our case. 4. The nasal bones in D. thraceiensis sp. n. are short, 7. The zygomatic arch (Figs. 5, 6, and 7 – zyg) of D. narrow, frontally rounded and fused. In the other thraceiensis sp. n. with its developed proc. postorbit- species they are longer, wider, almost flat for most of alis resembles the one of D. levius Jourdan, but in its their length, and separated. In the Gussiatin species posterior part, behind its connection with the tem- they protrude above the external nares. This is not poral bone, it lacks the processus that Svistun notes. the case with the two other skulls (Figs. 5, 6, and 7). He regards this processus as related to the develop- 5. The calvaria of the skull (fronto-parietal part, Figs. ment of the lower tusks. No comparison with the zy- 5, 6, and 7 – Ic in D. thraceiensis sp. n.) is high and gomatic arch of the Eppelsheim skull is possible, wide). In the other two species it is lower and shorter. because it is not preserved.

18 8. There are significant differences between the three D. bavaricum H. v. Meyer has small P3s. Their skulls in the occipital region. Os occipitale in D. thra- inner ridges are some more perpendicular to the outer ceiensis sp. n. is high, wide and at an angle of 80° one. This is rather clear with the anterior inner ridge. toward the forehead, and of ca. 70° to the plane of The valley between them is wider and deeper. the teeth. In D. giganteum Kaup these angles are 70° In D. pentapotamiae Lydekker, D. levius Jourdan and 50° correspondingly; in D. levius Jourdan the and D. indicum Falconer the differences are almost declination of the occipital bone is even larger, so the same, concerning the position of the two inner the angle is only 60°. As a whole, the skull of D. levius ridges and the size of the tooth and its cingulum. Jourdan is very flat and low. Os occipitale is visibly The fourth premolar of D. thraceiensis sp. n. has concave and very wide, forming two lateral wings inner ridges perpendicular to the outer one. The (Fig. 6 C and D – OS). Those wings are almost lack- posterior is narrower and longer. The triangular val- ing in D. thraceiensis sp. n. and in D. giganteum ley surrounded by the posterior inner wall of the out- Kaup they are much less developed. er ridge and the inner wall of the posterior ridge is 9. The position of the occipital condyles is rather clearer than in all the other species. different (Figs. 5, 6, and 7 – oc). In D. thraceiensis D. gigantissimum Stefanescu has a longer P4. Its sp. n. they are situated in the middle and in the low- transverse ridges are at a larger distance from each er part of the occipital bone, almost the same is their other. Besides, it has a strong cingulum on the ante- position in D. giganteum Kaup, and in D. levius Jour- rior and the posterior side, lacking in our species. dan they are on the upper half of the noted bone. In D. bavaricum H. v. Meyer P4s are smaller and According to Svistun, this permitted the animal to of a more regular square shape. The transverse ridg- raise its head almost at right angle to its neck, thus es are more distant at the inner side of the crown, fully using its back curved lower tusks. and the valley surrounded by them and the outer 10. Worth mentioning is a characteristic peculiarity ridge is wider and larger. of the Gussiatin skull lacking in all the others, In D. levius Jourdan the posterior inner ridge is including the skull from Ezerovo. This skull has a also attached to the outer one with its longer posteri- double articulation with the mandible. Once by an or branch. articular surface on the zygomatic process of the In D. indicum Falconer and D. pentapotamiae temporal bone and then by a second surface on the Lydekker the last premolars have a square shape and petromastoideum. Between those two surfaces is the are smaller. Their transverse ridges are parallel and external meatus of the auditory canal. How and of equal size. The valley between them is larger and when this second articular surface was used, is everywhere equally wide. unclear. In the M1s of D. giganteum Kaup, unlike D. thra- After comparing their anatomy we can see that ceiensis sp. n., the three ridges are almost of equal deinotheres could be divided in two groups. One in- size, with an equally developed cingulum on both cludes D. giganteum Kaup, D. gigantissimum Stefa- sides of the valleys, blocking them entirely on the nescu, D. thraceiensis sp. n., D. levius Jourdan and outer side. The latter is of significant taxonomical D. indicus Falconer, which have larger teeth, and the value. second – D. bavaricum H. v. Meyer, D. pentapota- In D. gigantissimum Stefanescu has a developed miae Lydekker and others, all with smaller teeth. cingulum on the outer anterior and on the posterior As a whole, the teeth are rather similar and differ- side of the crown, resembling a chain of large tuber- ences are seen mainly in the two premolars P3 and P4. cles, closely arranged along the end of the tooth. In D. giganteum Kaup P3 is more rounded on the There are no significant differences in the struc- inner side. The two inner ridges are more perpendic- ture of the first molars of D. bavaricum H. v. Meyer, ular to the outer one. The anterior inner ridge is not except in their size. connected with the outer and is at an acuter angle to it The other molars (M2 and M3) of D. thraceiensis than in other taxa. They almost don’t touch each other are not very different from those of the other species. on the inner side. If anything like that is observed in these species, than it is down at the very basis. The Mandible. (Pl. III, Fig. 1 and Pl. V). ridges merge when they are worn. The cingulum is Like the skull, it is entirely preserved. Compared to more clearly seen here. It consists of numerous large the size of the body, it is not large (measurements in and small tubercles on all sides of the crown. Table 3). Ramus horizontalis long, high, wide and No comparison with the third premolars of D. gi- laterally slightly flattened. Below P3 there is a well gantissimum Stefanescu is possible, because they have shaped double opening – foramen mentale. The been restored after D. giganteum Kaup. highest point of the mandible is at the symphysis, it In D. levius Jourdan the anterior inner ridge of P3 becomes lower towards the posterior end (see Plate is situated diagonally to the outer one and touches V). Because of this, the last molar is not horizontal, it. The posterior one is wider and V-shaped in its but inclined backwards. Processus angularis is strong- upper part, with a shorter anterior part. It is also di- ly developed, and fossa masseterica is wide but not agonal to the outer ridge, but does not touch it. very deep. The most typical feature of the animals In D. indicum Falconer the inner ridges are some belonging to this group is the shape of the symphys- more diagonally situated, compared to D. giganteum is. Together with the tusks, it is curved down and Kaup, but their tips do not contact. back. In D. thraceiensis sp. n. the symphysis is solid,

19 H. v. Meyer, Meyer, v. H. Germany Germany D. bavaricum

from Eppel Eppel sheim D. giganteum Deperet, 1887 1887 Deperet, from Seint Seint Yean Münhen Stromer, , е ы after after Белокр . 1960, Krivoi Kaup А Rog, Moldova D. giganteum after after Romania Romania Athanasia, 1907, 1907, Athanasia, h, h, еви j after after 1954, Yugoslavia Петрон

Bulgaria Bulgaria sin dext sin dext sin dext sin dext sin dext dext sin sin dext sin dext sin sin dext dext dext sin sp.n. Ezerovo, D. thraceiensis D. Enamel thickness Enamel thickness 4.2 - 4.0 - 5.0 - Enamel ------4.3 - - 4.2 - - Enamel thickness Enamel ------4.3 4.3 thickness Enamel ------4.6 4.7 thickness Enamel ------5.2 5.1 Length of the tooth tooth the of Length tooth the of Width ridge second of Width Width of second ridge tooth the of Length tooth the of Width Width of first ridge 100.0 Width of second ridge 105.0 90.5 100.0 106.5 tooth the of Length 106.5 base the at ridge 91.8 first the of Width 106.0 - Width of the first ridge at the top - Width of the second ridgethe base at 75.0 - - 94.6 73.5 Width of the second ridge 100.8 the at top 100.5 Width of the third ridge atthebase 94.0 81.5 69.0 100.0 96.3 70.0 68.2 87.0 98.5 Width of the third ridge atthetop 70.4 102.5 69.0 100.0 Width 97.0 of posterior talon 50.5 87.0 73.0 98.0 - 110.0 83.5 70.0 - 71.0 - - 70.8 tooth the of Length - 110.0 63.5 - - 84.5 - base 67.5 the at ridge first the of Width - - 79.0 98.0 - Width of the first 61.2 ridge at the top 75.0 - 62.0 100.0 62.0 base the at ridge second the of Width - - 113.6 96.0 50.0 97.0 - - Width of the second 37.0 ridgethe at top 110.0 - - 98.0 - 115.0 - - 72.0 Width of posterior talon 95.0 - - 88.8 110.0 - - 55.0 - 48.0 119.0 85.3 71.0 - 80.5 69.0 tooth the of Length - 45.0 93.0 87.5 - 79.0 - 88.0 107.7 base the at ridge first the of Width 85.0 71.0 84.5 - - 86.0 - - - - Width of the first ridge at the top - 106.5 86.0 - - - - - Width of the second ridgethe base at 83.0 - 118.0 - - - 68.0 - 104.0 Width of the second ridgethe 79.5 at top 99.0 - - 107.2 115.0 - - 68.0 66.0 Width of posterior talon - - 100.0 - 63.0 - 98.0 - - - 107.0 - - - - 66.0 68.0 87.0 - - - 98.0 77.0 - - - 74.0 88.0 52.0 110.0 - - - 81.0 89.0 - 89.0 - - - - - 73.0 - 53.0 78.0 108.0 - - 73.0 - - 108.0 - 82.0 ------61.0 69.0 - - - 115.0 - - 78.0 ------61.0 ------99.0 61.0 - - 78.0 50.0 - - - 70.0 - 62.0 - 47.5 - - 61.0 95.0 - - 82.0 ------62.0 - - - 33.0 - - - - - 75.0 81.0 - - - 54.0 - - - - 80.0 - 75.0 ------66.0 72.0 78.0 - - - - - 75.0 - - 66.0 ------51.0 ------89.0 ------91.0 66.0 - - - 89.0 ------54.0 66.0 68.0 - - - - - 68.0 - 81.0 ------60.0 - - 82.0 - 44.0 61.5 - 64.0 ------70.0 - - - - - 70.0 - 91.0 61.0 - - - - - 67.0 - - - - - 63.0 - - - 63.0 - 62.0 - - - - - 63.0 ------56.0 ------Table 2 A Upper cheek-teeth

1 2 3 Measurements in mm 3 4 P P M M M

16 Pakistan after Dehm 1963, D. pentapotamiae sin dext dext sin sin Fal. Narbada, India D. indicum D. pentapotamiae pentapotamiae

Kaup Kaup D. cuvieri D.

Kaup Kaup D. giganteum

Rhone – Deperet, 1887 1887 – Deperet, Rhone 1880 Lydekker, after Jordan D. levius sp.n. sp.n. sin dext dext sin Ezerovo, Bulgaria D. thraceiensis D. : In the columns after Lydekker, 1880, size is in inches. Width of the second ridgethe at top 69.0 71.0 base the at ridge second the of Width 110.0 110.0 base the at ridge second the of Width - 107.2 107.0 ------71.8 69.0 59.6 64.3 Measurements in mm 1 2 3 ridge second of Width Width of first ridge 90.5 91.8 96.3 - 87.0 ------51.0 52.0 - 60.0 - - 62.2 3 4 thickness P tooth the of Length tooth the of Width Enamel ridge second of Width thickness P tooth the of Length tooth the of Width Enamel 4.0 4.2 ridge second of Width 100.0 - 105.0 tooth the of Length 106.5 base the at ridge first the of Width - 100.0 M 4.2 Width of the first ridge at the top 106.5 4.3 base the at ridge thickness second the of Width 106.0 - 75.0 66.0 94.6 100.8 Width of the third ridge atthebase 100.5 Enamel 94.0 Width of the third ridge atthetop - 70.0 - - - Width of posterior talon 81.5 100.0 70.0 98.5 102.5 110.0 83.5 - 97.0 tooth the of Length 73.0 M base the at ridge thickness first the of Width 63.5 4.3 - Width of the first ridge at the top 110.0 - 53.0 4.3 - 84.5 - Enamel Width of the second ridgethe at top 113.6 62.0 Width of posterior talon - - - 78.0 - 88.8 - 115.0 - tooth the of Length - 85.3 M - - - - base the at ridge thickness first - the of Width 4.7 107.7 - - Width of the first ridge at the top - 4.6 87.5 - Enamel 95.0 84.5 - - - Width of the second ridgethe at top 106.5 118.0 - - - Width of posterior talon - - - - 100.0 - - - - - 115.0 72.0 - 87.0 - 5.1 110.0 - - 98.0 - 5.2 - - - 2.1 89.0 - 108.0 - - - 2.3 - - - 91.0 78.0 - 50.0 ------51.0 - 78.0 - - 60.0 - 58.4 - - - 2.5 - 45.0 - - 3.9 - - - 2.5 - 51.0 - 91.0 ------2.1 - 62.0 58.2 - - - 72.0 - 52.2 - - 62.0 - - - - 70.0 - - 60.3 - - - 62.2 - 2.4 - 2.4 68.4 - - - - - 2.5 2.75 52.4 ------62.2 - 64.4 ------73.2 - - - - 71.3 - - 67.0 - 70.0 66.0 - - - - - 74.0 - 68.4 ------Note Table 2 B Upper cheek-teeth

3 Geologica Balcanica, 3-4/2006 17 Fig. 6. Deinotherium levius Jourdan

Fig. 7. Deinotherium giganteum Kaup

19 H. v. Mayer Mayer v. H. D. bavaricum Kaup D. giganteum Moldova München H. v. Meyer Meyer München v. München Moldova H.

Romania Romania D. gigantissimum sp.n. sp.n. Ezerovo D. thraceiensis D. 150.0 ------150.0 120.0 1 3 105.0 - - - - 105.0 3 510.0 540.0 470.0 485.0 370.0 365.0 365.0 370.0 485.0 470.0 540.0 510.0 1 -M 3 230.0 190.0 230.0 210.0 160.0 154.0 154.0 160.0 210.0 230.0 192.0 190.0 - 230.0 185.0 1 3 235.0 - - - - 235.0 3 Dimensions, mm Dimensions, 1. ascendens ramus of end the to beginning the – from length Total 2. horizontal – symphysis of Length 3. – vertical symphysis of Length 4. Diameter of the symphysis5. at the angle with the mandible I of beginning the at symphysis the of Diameter 6. symphysis the of through the of Width 7. 1120.0 ascendens ramus and symphysis of Length 8. at P mandible the of Height 9. M at mandible the of Height 1200.0 310.0 280.0 1000.0 360.0 1100.0 435.0 1120.0 145.0 - - 420.0 820.0 1150.0 - - - 840.0 - - - - 320.0 - 230.0 - 235.0 - - 10. 10. M at mandible the of Height 11. articularis proc. at ascendens ramus of Height 12. coronoideus proc. at ascendens ramus of Height 13. ascendens ramus of middle – Width 14. at P semimandibles two the between Distance 15. M at semimandibles two the between Distance 16. M at semimandibles two the between Distance 17. – I incisors the of Length 18. basis the – at I of Diameter 19. middle the – in I of Diameter 20. of I points the between Distance 21. ascendens and ramus I of point the between Distance 22. P row tooth entire the of Length 685.0 505.0 470.0 680.0 895.0 - 455.0 482.0 855.0 132.0 125.0 470.0 - - - 608.0 172.0 120.0 - - 900.0 142.0 125.0 - 795.0 530.0 525.0 №

Table 3 Mandibles – mandibulae

20 , Bakalov, raceminor Nikolov, 1962,Bulgaria Kaup D. giganteum D. Bulgaria Bulgaria Bakalov, Nikolov, 1962, D. gigantissimum Stefanescu, 1909, Romania D. thraceiensis thraceiensis D. sin sin dext sin dext dext sin dext sin sin dext sp.n. Ezerovo, Bulgaria Width of the second ridgethe at top - base the at ridge second the of Width - 97.5 base the at ridge second the of Width 107.0 - 95.0 97.0 100.0 - - 107.0 - 82.0 - - 99.5 - - 83.0 - - 93.0 - 73.0 - 96.0 76.0 - - - Measurements in mm 1 2 3 Width of first ridge Width of first ridge 58.0 55.0 76.0 78.0 - 72.0 - - - 68.0 - - 88.0 ------3 4 Enamel thickness P Length of thetooth Enamel Width of the tooth thickness Width of second ridge P 4.0 tooth the of Length Enamel 3.8 Width of the tooth - ridge second of Width 4.0 tooth the of Length 80.5 4.0 base the at - thickness ridge first the of Width M 68.0 Width of the first ridge at the top 70.0 - - - base the at ridge second the of Width - Enamel 75.5 base the at ridge third the of Width 89.0 67.0 - - 82.0 Width of the third ridge atthetop 73.0 80.0 81.0 Width of posterior talon 81.1 - - thickness - - 4.3 tooth the of Length M 89.0 75.0 4.2 107.5 base the at ridge first the of - Width 81.5 81.0 - Width 81.5 of the first ridge at the top 83.0 - Enamel - - - Width of the second ridgethe at top 107.0 90.0 75.5 - Width of posterior talon 72.0 103.0 - thickness 72.0 73.0 4.6 tooth the of Length M - - - 80.0 - - 4.5 base the at ridge first the of Width 111.0 83.0 - - - Width of the first ridge at the top 71.0 110.5 Enamel - - 115.5 77.0 - - Width of the second ridgethe at top - 81.0 talon posterior of Width 106.0 75.5 - - - 109.0 - - 112.5 99.0 5.2 - 70.0 - - - - 85.0 - 5.3 - 78.5 - 80.0 - 105.0 - - 120.0 - 111.0 - 62.0 ------85.0 82.0 92.0 - - 80.0 - - - 114.0 118.0 64.0 - 111.0 82.0 - 86.0 ------75.0 ------115.0 84.0 67.0 ------103.0 ------106.0 - - 81.0 - - - - - 104.0 - - - - 109.0 - - - 84.0 76.0 ------110.0 - 73.0 ------100.5 73.0 - 68.0 37.0 - - - - 66.5 70.5 - 110.0 ------92.0 - - - - - 71.0 - - 67.0 65.0 - - - 115.5 - - - - - 72.0 ------56.0 ------Table 4 Lower cheek teeth

21 1939, Romania Romania Simionescu. Museum of München München of Museum R. Deim, 1949, South Bavaria, München München Kaup Stromer, 1938, D. giganteum , 1960, Белокрые . А Krivoi Rog,Moldova h, 1954, еви j Yugoslavia Петрон Bulgaria Bulgaria sp.n. Ezerovo, D. thraceiensis thraceiensis D. sin sin dext sin dext sin dext sin dext dext sin sin dext sin Width of the second ridgethe at top - base the at - ridge second the of Width 97.5 - 107.0 base the at ridge second the of Width 66.0 - 95.0 - 100.0 62.0 - - - - - 70.0 ------76.0 - 76.0 - - - - 76.0 76.0 - Measurements in mm 1 2 3 Width of first ridge Width of first ridge 58.0 55.0 76.0 38.5 78.0 28.0 57.5 - 39.0 - - - - - 55.0 ------21.5 3 4 thickness P tooth the of Length tooth the of Width Enamel Width of second ridge thickness P tooth the of Length 4.0 tooth the of Width Enamel 3.8 - Width of second ridge 80.5 - - tooth the of Length 68.0 70.0 4.0 Width of the first ridge at the base - M 75.5 4.0 Width of the first ridge at the top - thicknessbase 67.0 the at ridge second the of Width - 73.0 89.0 - - 58.2 - 82.0 - Width of the third - ridge atthebase 80.0 - Enamel 81.0 - 81.1 Width of the third ridge atthetop - - 89.0 49.1 - 40.0 Width of posterior talon - 81.5 81.0 - 81.5 75.0 107.5 83.0 tooth the of Length - 81.0 M 64.4 - - thickness base the at 4.3 ridge first the of Width - - - 4.2 - - - 107.0 Width of the first ridge at the top 59.7 - - 75.5 - 50.5 - Enamel - - Width of the second ridgethe at top 64.0 103.0 - Width of posterior talon ------80.0 40.0 - 110.5 tooth the of Length 83.0 M 60.0 - thickness base the at ridge 4.6 first the of Width - - 4.5 115.5 - - Width 81.0 of the first ridge at the top 68.7 88.0 ------75.5 - - Enamel - - 112.5 - Width of the second ridgethe at top 106.0 - Width of posterior talon - 55.0 68.0 - - - - - 62.0 99.0 85.0 - - - - 76.7 - - 105.0 - 78.5 5.2 - - - - - 5.3 120.0 82.0 ------80.0 - 56.0 - 55.0 - - - 58.0 - - 118.0 - 68.0 55.0 - - 64.0 - 86.0 57.0 102.0 - - - - 75.0 ------69.0 52.0 - - - - 67.0 - 47.0 71.0 - 67.0 60.0 - - - - 93.0 ------47.0 ------80.0 62.0 - 100.0 - 79.0 - - 60.0 - - - 105.0 61.0 - - 76.0 - - - - 62.0 ------? 68.0 - 107.0 83.0 - - - 76.0 - - 82.0 - - - 82.0 - - - - 100.0 - - 74.0 - - 76.0 - - 28.0 - 80.0 - - - - 27.5 78.0 22.0 - - - - 78.0 - - - 68.0 93.0 - - - - 76.0 - - 81.0 - - 76.0 - - - 86.0 - - - - - 83.0 ------Table 4 A Lower cheek teeth

22 Pakistan Lydekker, after R. Dehm, 1963, 1963, R. Dehm, D. pentapotamiae D. indicum indicum Falconer Lydekker Lydekker

from Siwaliku Narbada, after Lydekker, 1889, India India 1889, Lydekker, after D. pentapotamiae Kaup Kaup D. cuvieri cuvieri D. D. Rhone Rhone Kaup Kaup giganteum giganteum Deperet, 1887 1887 Deperet, D. levius levius Jordan H.v. Münhen D. bavaricum Meyer, R.Dehm,1949, South Bavaria,Museum of Bulgaria Bulgaria sin dext dext sin sp.n. Ezerovo, D. thraceiensis thraceiensis D. Width of the second ridgethe at top - - base the at ridge second the of Width - 97.5 107.0 - 58.5 53.0 - base the at ridge second the of Width - 95.0 100.0 - 51.0 ------58.0 ------Width of first ridge Width of first ridge 58.0 55.0 - 76.0 - 78.0 - - - - 47.0 ------42.5 - - : In the columns after Lydekker, 1880, size is in inches. 1 2 3 3 4 Measurements tooth in the mm of Length thickness tooth the of Width Enamel Width of second ridge tooth the of Length thickness tooth the of Width Enamel 80.5 4.0 Width of second ridge 3.8 - 68.0 75.5 tooth the of Length - 70.0 bas the at ridge first the of Width 67.0 44.0 - 73.0 Width of the first ridge at the top 43.0 base the at ridge second the of 89.0 4.0 Width - - 4.0 82.0 - - 80.0 thickness 81.0 89.0 Width of the third ridge atthebase - 81.1 36.0 - - 81.5 Width of the third ridge atthetop 81.0 Enamel - 43.0 81.5 - - 83.0 Width of posterior talon 47.0 107.5 75.0 50.0 - 50.0 - 55.0 - - tooth 107.0 the of Length - - 41.0 - 75.5 57.0 - base the at ridge first the of - Width 42.0 72.0 Width of the first ridge at the top - 45.0 - thickness 63.0 - - - 67.0 4.3 70.0 103.0 4.2 - - - - Width of the second ridgethe at top - Enamel 110.5 - 68.0 Width of posterior talon - - - - 80.0 - - 43.0 57.0 70.0 83.0 - - tooth the of - Length - 81.0 115.5 - 54.0 base the at ridge first the of Width - 75.5 - - - 112.5 Width of the first ridge at the top - - - thickness - - 48.0 4.6 - 106.0 1.8 - 67.0 4.5 Width - of the second ridgethe at top - Enamel 105.0 - - 64.0 Width of posterior talon - - - - 85.0 - - - 59.5 78.5 - - - 71.0 - 82.0 - 1.8 120.0 55.0 - 56.0 1.47 - - 80.0 - 118.0 - - - 2.1 - - 5.2 - 60.0 - - 68.0 5.3 - - - - - 64.0 - 70.0 - - - - - 67.0 - - - 77.0 2.05 2.34 - 1.7 - - 2.4 - - - 2.9 ------54.0 56.0 - 1.8 - - - 59.0 - - 2.1 ------4.0 2.15 - - - 2.8 - - 2.6 2.4 - 33.0 - 67.0 - - - 47.0 - - - 2.15 42.0 - 72.0 - - - - 2.90 - - - 3.5 - - 2.25 - - - - 45.8 - - - - 2.9 - 3.9 65.0 - - - 2.45 73.0 ------3.18 ------65.0 - - - - - 73.0 ------P P M M M

Note Table 4 B Lower cheek teeth

23 PLATE VII Deinotherium thraceiensis sp. n. 1. Part of vertebral column (vertebrae, ribs). Scale ca. 5% Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE VIII Deinotherium thraceiensis sp. n. 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 Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE IX Deinotherium thraceiensis sp. n. 1. Third cervical vertebra. Coll. SU.M. No. SU Pl 312/3-3 2. Fourth cervical vertebra. Coll. SU.M. No. SU Pl 312/ 3-4 3. Fifth cervical vertebra. Coll. SU.M. No. SU Pl 312/3-5 4. Second thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-9 5. Sixth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-13 Scale of all figures ca. 11.9%; Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE X Deinotherium thraceiensis sp. n. 1. Ninth thoracic vertebra. Coll. SU.M. No. SU Pl 312/ 3-16 2. Tenth thoracic vertebra. Coll. SU.M. No. SU Pl 312/ 3-17 3. Eleventh thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-18 4. Twelfth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-19 5. Thirteenth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-20 6. Fourteenth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-21 Scale of all figures ca. 11.8%; Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XI Deinotherium thraceiensis sp. n. 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 Deinotherium thraceiensis sp. n. 1, 1a. Right scapula in medial and lateral view. Coll. SU. M. No. SU Pl 312/6 2, 2a. Left scapula in medial and lateral view. Coll. SU.M. No. SU Pl 312/5 Scale ca. 4.6%; Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XIII Deinotherium thraceiensis sp. n. 1, 1a. Right humerus in lateral and cranial view. Coll. SU.M. No. SU Pl 312/8 2. Left humerus in cranial view. Coll. SU.M. No. SU Pl 312/7 Scale ca. 5.4%; Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XIV Deinotherium thraceiensis sp. n. 1, 1a. Right ulna in lateral and medial view. Coll. SU.M. No. SU Pl 312/10 2, 2a. Right radius in lateral and medial view. Coll. SU.M. No. SU Pl 312/12 Scale ca. 6.2%; Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XV Deinotherium thraceiensis sp. n. Carpals 1. Cuneiforme (ulnare) sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-1 2. Lunare sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-2 3. Magnum sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-7 4. Unciforme sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-8 5. Pisiforme sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-4 Locality: Ezerovo, near Plovdiv; Level: Maeotian

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 XIX

PLATE XX

PLATE XXI PLATE XXII PLATE XXIII

PLATE XXIV PLATE XVI Deinotherium thraceiensis sp. n. 1, 1a. Metacarpus I of the first digit in frontal and lateral view. Scale ca. 18.6%. Coll. SU.M. No. SU Pl 312/14-1 2, 2a. Metacarpus II of the second digit in frontal and lateral view. Scale ca. 18.6%. Coll. SU.M. No. SU Pl 312/14-2 3, 3a. Second phalanx of the second digit in frontal and lateral view. Scale ca. 14.1%. Coll. SU.M. No. SU Pl 312/15-2 Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XVII Deinotherium thraceiensis sp. n. 1, 1a. Metacarpus V sin in frontal and lateral view. Scale ca. 22.2%. Coll. SU.M. No. SU Pl 312/14-5 2, 2a. Hoof of the third digit from above and from below. Scale ca. 65%. Coll. SU.M. No. SU Pl 312/15-10. 3, 3a. Second phalanx of the fourth digit in frontal and lateral view. Scale ca. 22.2%. Coll. SU.M. No. SU Pl 312/15-8 4. Patella. Scale ca. 22.2%. Coll. SU.M. No. SU Pl 312/18-2 Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XVIII Deinotherium thraceiensis sp. n. 1. Left hand. Scale ca. 7.7%. Coll. SU.M. No. SU Pl 312/13 Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XIX Deinotherium thraceiensis sp. n. 1. Pelvic bones with the sacrum. Scale ca. 4.0%. Coll. SU.M. No. SU Pl 312/16-1 and 2; SU Pl 312/3-32 Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XX Deinotherium thraceiensis sp. n. 1. Pelvis in lateral view. Scale ca. 4.0%. Coll. SU.M. No. SU Pl 312/16-1 Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XXI Deinotherium thraceiensis sp. n. 1, 1a. Right femur in dorsal and caudal view. Coll. SU.M. No. SU Pl 312/17-2 Figs. 2 and 2a. Left femur (restored) in caudal view. coll. SU.M. No. SU Pl 312/17-1 Scale ca. 6.8%; Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XXII Deinotherium thraceiensis sp. n. 2, 2a. Left tibia (restored) in dorsal and caudal view. Coll. SU.M. No. SU Pl 312/19-1 3. Right tibia (restored) in dorsal view. Coll. SU.M. No SU Pl 312/19-2 Scale ca. 8.0%; Locality: Ezerovo, near Plovdiv; Level: Maeotian

PLATE XXIII Deinotherium thraceiensis sp. n. 1, 1a. Pes (restored). Scale ca. 9.0%. Coll. SU.M. No. SU Pl 312/14 2. Calcaneus sin. Scale ca 20.3%. Coll. SU.M. No SU Pl 312/21-1 Locality: Ezerovo, near Plovdiv; Level: Maeotian.

PLATE XXIV Deinotherium thraceiensis sp. n. Restored skeleton Coll. SU.M. No SU Pl 312 Locality: Ezerovo, near Plovdiv; Level: Maeotian

4 Geologica Balcanica, 3-4/2006 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 M1 a widening begins, rami coming closer again at the level of M3, 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 al- Fig. 8. Shape of the tusks of Deinotherium thraceisensis sp. n. ready wholly solid. They gradually become thinner A – shape of the tusks at the basis of the alveolus; B – shape of and, curving in two directions – outwards and back- the tusks at 115.0 mm from their basis in the symphysis; C – wards, go apart from each other. Their tips are point- cross-section of I dext at 600.0 mm from the symphysis – ed, slightly smoothed only at the foremost part of the natural size 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 they don’t touch the ridge in front of them. On the five teeth each – two premolars and three molars. inner side, at the bottom of each valley there is a weak Unlike the upper premolars, the lower are much tubercle, and on the anterior side – a small cingulum. narrower, and their structure is very different, Left M1 resembles the right but is more worn. especially P3. M2 dext is large and tetragonal. It has two thick P3 has a long and narrow crown, pointed at the transversal ridges, wider on the outer side. There both anterior end. One large longitudinal ridge is situat- ridges form a small concavity. The valley between ed along its axis, and another, transversal – in the them is deep and free but on the inner side there is a posterior part. They form the occlusial surface of small tubercle. On its anterior and posterior sides the the tooth. The longitudinal ridge is wider at its basis, crown has a cingulum. gradually narrowing towards the apex of the crown. M3 dext is large too but with an irregular tetrago- Its widest part is in the middle of the crown. There it nal shape. Built by two transversal ridges. The first is is more worn out and part of the dentine is seen. On wider. Both are widening outwards. At the ends – the inner part of the tooth, at the place where the inner and outer – they are slightly curved to the front, longitudinal ridge contacts the inner one, the first forming with their anterior walls shallow valleys. The narrows abruptly thus shaping a large triangular valley between them is deep and unblocked. The valley. Outside the crown is smooth. posterior talon on the inner side of the crown is formed P4 dext has a larger crown, elongated and wide. by numerous tubercles of different size. The occlusal surface is moderately worn. It is formed by two transversal ridges, contacting at the outer part Comparison. The structure of the described mandi- of the crown. At this place they divide the valley be- ble is close to the other species (Fig. 9). tween them in two not quite equal parts. The anterior The symphysis is moderately large. There are dif- wall of the first ridge is strongly concave. The poste- ferences in the shape of ramus horizontalis, proc. rior part of the crown has a weak cingulum. articularis, proc. coronoideus and the tusks. M1 dext has three transversal ridges. First two are 1. The symphysis of D. bavaricum H. v. Meyer is more more worn. Their structure is generally the same as protruding. In D. levius Jourdan it is more rounded in M1. All three ridges are wider on the outer side. and curved inwards and in D. giganteum Kaup, as The valleys dividing them become gradually narrower well as in D. gigantissimum Stefanescu and D. thra- towards the outer part. In this part the anterior walls ceiensis sp. n. the curving is even stronger and more of the second and third ridge are slightly convex but gradual.

26 shallow. The symphysis is some more elongated, but it has been restored. No P3 is known for this species. 7. The third premolar – P3 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 P3 which surrounds a deep valley together with the longitudinal ridge. A high cingulum encircles the crown. In D. bavaricum H. v. Meyer P3 is smaller and built by two smaller transversal ridges and a large longitudinal one, positioned on the outer side of the crown. 8. P4 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 P4 has the same structure as D. giganteum Kaup but without the large cingulum. P4 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 “C- shaped”. It surrounds a tetragonal valley anteriorly. Strong cingulum. As a whole, the fourth premolars in the studied taxa are rather close in their structure. 9. M1, M2 and M3 in the species described show no significant differences when compared to the other studied species. A small exception is the fact that our species lacks a developed cingulum. The dis-

placement of the third ridge in M3 is not regarded as Fig. 9. Comparison of deinothere mandibles: a – Deinotheri- a distinct taxonomic peculiarity by us. um bavaricum H. v. Meyer; b – Deinotherium levius Jourdan; We had no opportunity to compare the material c – Deinotherium giganteum Kaup; d – Deinotherium thra- to the species Kaup, Jourdan, ciensis sp. n. D. cuvieri D. levius D. indicum Falconer, D. pentapotamiae Lydekker.

Vertebral column (Pl. VII). 2. In D. thraceiensis sp. n. ramus horizontalis (in the In D. thraceiensis sp. n. it consists of 7 cervical, prob- region of proc. angualris) strongly protrudes back- ably 14 thoracic, probably 14 lumbar, 3 sacral and wards with a pronounced curve. It almost reaches probably 6 caudal vertebrae. the level of incisura mandibulae. Atlas (Pl. VIII, Figs. 1, 2 and 3). 3. Ramus ascendens is almost perpendicular to ra- Almost entirely preserved. Processes are only partial- mus horizontalis in its anterior part. ly broken, without losing the shape of the vertebra. It 4. Processus articularis is high and abruptly separat- consists of two arches – dorsal and ventral, limiting ed from ramus ascendens, forming with it an obtuse a large foramen vertebrae. The foramen widens down- angle in its posterior part. Anteriorly it descends wards and is slightly constricted in the middle. There’s steeply toward incisura mandibulae. no corpus vertebrae. In fact the atlas is a wide bone 5. Processus coronoideus is high too and anteriorly ring. The transverse processes have become large bone almost vertical, nearly at the same level as proc. ar- plates – wings of the atlas. There is no spinous pro- ticularis. cess. The atlas articulates with the skull by wide oval 6. Incisura mandibulae is deep and almost horizon- concavities, cranially positioned on the arches. Cau- tal in relation to the mandible, perpendicular to ra- dally, on the ventral arch, there is an articular sur- mus ascendens. face for the dens of the axis. Foramen ovale and for. In D. gigantissimum Stefanescu the curve at proc. ventrale lateralis on both sides of the vertebra and angularis is weaker. Poc. articularis is not so abrupt- large (measurements in Table 5). ly separated from it. Proc. coronoideus posterioly Axis (Pl. VIII, Fig. 4 and 5). Body narrow and oval, turns into incisura mandibulae. The latter is very spinous process high, wide and long. Two strong

27 Table 5 Cervicals

CERVICSLES № Dimensions, mm Atlas Axis 3 4 5 6 7 1. Height of the vertebra 300.0 340.0 330.0 340.0 343.0 350.0 352.0 2. Width of the vertebra 400.0 320.0 310.0 315.0 218.0 224.0 226.0 3. Length of corpus vertebrae 212.0 230.0 238.0 240.0 246.0 250.0 4. Width of corpus vertebrae 170.0 195.0 202.0 206.0 208.0 210.0 5. Length of articular surfaces 100.0 103.0 105.0 106.0 108.0 6. Height of processus spinosus 60.0 150.0 7. Length of foramen vertebrae 200.5 165.0 60.0 80.0 8. Width of foramen vertebrae 145.0 93.0

Table 6 Thoracics

THORACICS № Dimensions 1 2 3 4 5 6 7 8 9 10 11 12 13 14 1. Diameter of corpus 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 vertebrae 2. Thickness of corpus 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 vertebrae 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. 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 transversus 6. Length of foramen 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 vertebrae 7. Width of foramen 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 vertebrae

crests on tip with a deep furrow between them. Dens wards the spinous process. All other thoracic verte- epistrophei connects the axis with the atlas. Fora- brae resemble the second. There are differences only men vertebrae much smaller than in atlas, tetragonal in the size of proc. transversus and proc. spinosus, (measurements in Table 5). which are smaller in each next vertebra. From the Third, fourth and fifth cervicals (Pl. IX, Figs. 1, 2 first to the twelfth the size of the corpus and foram- and 3). All have similar structure. Narrow bodies. ina decreases, then increases to the last. In the fif- Cranially slightly convex, caudally slightly concave. teenth and all the following there is no fovea costal- Transverse processes small and rounded in the ends. is. So, there were no ribs attached to these vertebrae Transverse foramina limited by the bodies and the (measurements in Table 6). transverse processes. Well pronounced articular surfaces on the cranial and caudal processes. Foramen Lumbar vertebrae (Pl. XI, Fig. 4) vertebrae wide and high. Spinous processes not pro- Resemble the last thoracic vertebrae. Here, too, fovea nounced. costalis is missing. The diameter and thickness of Sixth and seventh cervicals are built in the same corpus vertebrae gradually decreases from the first way as the previous three, but they have spinous pro- to the last. Spinous processes become shorter. As in cesses, largest on the seventh. Below the transverse the thoracic vertebrae, they are inclined backwards. processes the seventh cervical has a well shaped fovea In the other vertebrates these processes on the lum- costalis – a concavity for the head of the first rib bars are pointing anteriorly. This peculiarity shouldn’t which is between the seventh and the eighth vertebra be of taxonomic importance for genus Deinotherium (measurements of all cervicals in Table 5). because it is present in the other proboscideans as well. The transverse processes are not long as should Thoracic vertebrae (Pl. IX, Fig. 4 and 5, Pl. X, Fig. 1, be expected for lumbars so we cannot speak of proc. 2, 3, 4, 5, and 6). costarius. These peculiarities in the development of The first thoracic vertebra is very similar to the last the processes of the lumbars and the lack of fovea cervical, only its spinous process is larger. Thus it costalis in the last thoracic vertebrae make it diffi- resembles the second thoracic vertebra, where this cult for us to decide unequivocally where one group process is most developed. The second thoracic ver- of vertebrae ends and the other begins. This is why tebra (Pl. IX, Fig. 4) has a distinct structure. Its body we cannot say for sure how many the thoracic and is small, cranially convex, caudally – slightly round- correspondingly the limbar vertebrae are. The diam- ed. Its cranial articular processes are less develped eters of foramen vertebrae increase as in the last tho- than the caudal. Fovea costalis cranialis larger than racic vertebrae. They are largest in the last vertebra f. c. caudalis. Spinous process strong, widening at (Table 7). This character too creates difficulties in the tip, tetragonal. Foramen vertebrae elongated to- the determination of the two groups of vertebrae but

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

Table 7 Lumbars

№ Dimensions 1 2 3 4 5 6 7 8 9 10 1. Diameter of corpus vertebrae 178.0 177.0 176.0 174.0 172.0 170.0 168.0 165.0 162.0 160.0 2. Thickness of corpus vertebrae 93.7 95.0 97.0 97.5 98.0 100.0 100.0 98.0 96.0 94.0 3. Length at proc. transversus 274.0 269.0 264.0 258.0 250.0 240.0 240.0 242.0 243.0 245.0 4. Length of proc. spinosus 282.7 280.5 277.8 275.5 273.0 270.0 268.0 266.5 262.0 258.0 5. Length of proc. transversus 82.0 77.0 73.0 69.8 66.5 62.0 63.0 63.0 64.5 66.0 6. Length of foramen vertebrae 64.0 75.0 81.0 90.0 102.0 110.0 120.0 130.0 140.0 150.0 7. Width of foramen vertebrae 58.5 60.0 61.0 59.5 60.0 60.0 61.0 61.5 62.0 63.0

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

№ 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 Kaup Munchen - Stromer Stromer - Munchen D. giganteum Kaup Munchen - Stromer D. giganteum

Kaup Kaup D. giganteum Kaup

o Pikermi D. giganteum

Stefanescu D. gigantissimum St. Manzati, Romania Romania Manzati, St. sp.n. sp.n. sin dext Ezerovo, Bulgaria D. thraceiensis D. 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 70.0 67.3 64.5 62.0 61.7 45.0 52.0 54.5 46.5 46.0 48.1 45.5 61.3 67.0 60.5 45.0 56.3 60.0 60.9 85.0 81.0 105.0 75.0 91.0 98.0 114.0 160.0 180.0 195.0 220.0 213.0 153.0 206.0 198.0 194.0 203.5 162.0 80.0 136.0 120.0 140.0 50.0 thickening thickening rib beginning of caput thickening the to costae Dimensions Dimensions : The measurments in the table are taken for left ribs Dimensions, mm mm 1 2 5 6 3 4 7 12 14 13 11 10 8 9 Dimensions, 1. Total length length c-d 1. Total 2. Width 3. 4. m-l width Maximum 5. l of width Maximum 6. m of width Maximum 1135.0 7. 1135.0 neck the at Diameter 8. surface articular the at Diameter 9. 500.0 surface articular the at Height 500.0 spinae tuberositas of Length 260.0 895.0 970.0 650.0 245.0 260.0 895.0 254.0 300.0 635.0 650.0 245.0 245.0 635.0 730.0 280.0 650.0 525.0 260.0 150.0 - - 170.0 756.0 570.0 185.0 - 96.0 - 170.0 110.0 - 250.0 205.0 - - - 110.0 - 225.0 160.0 - - - № 10. 10. 11. acromion of Height at l Angle 215.0 215.0 90.0 85 - - 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 1102.0 950.0 840.0 840.0 950.0 1102.0 1195.0 1230.0 1274.0 1322.0 1376.0 1430.0 length 148.0 472.0 1450.0 1390.0 Total 1245.0 1. 2. 3. costae caput of Diameter Diameter at the the 4. 80.0 between the of width Maximum 5. Distance 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 №

Table 10 Ribs – costae Table 11 Scapula Note

30 also with a short and narrow neck as the first two ments from the left, and it was restored after the right. ribs. The body of the fourth is long and almost round. Dorsally strongly flattened, becoming thinner at the Caput costae spherical, divided by a well pronounced anterior and posterior edge. Spina scapulae and the furrow in the middle into two surfaces – facies ar- edges surround two concavities of unequal size. The ticularis and facies articularis condilis, the first of crest takes almost the entire length of the scapula. It which is bigger. Near its distal end the rib gradually is divided in two near the tubersitas spinae, forming becomes thinner. At the very end it becomes wider a wide long groove starting immediately behind the ending with a rough surface for the attachment of acromion and ending in the dorsal thickening. The the cartilage. The other ribs are similar to the fourth. acromion itself is high and curved towards the neck Their length gradually decreases (measurements in of the scapula. Fossa infraspina shallower but wider Table 10). than fossa supraspina. Margo thoracales long and pointing outwards. It has a regular triangular shape. Sternum. Only a fragment preserved, with a rough The articular surface for the attachment to the hu- surface and places for the attachment of the rib car- merus is shallow. tilages. Humerus (Pl. XIII, Figs. 1. 1a and 2). Both humeri Comparison. The most disputable problem concern- entirely preserved. Caput humeri large and curved, ing the ribs is their number. There is still no explicit with no distinct neck. Tuberculum majus protrudes opinion on that matter and we don’t know their pre- upwards, above caput humeri. Tuberculum minus is cise number. D. gigantissimum Stefanescu has 32 pairs much smaller. Positioned cranio-medially. Between of ribs restored, reaching up to the sacrum. What the two tubercles there is a deep and wide furrow – was the reason for this number, we do not know. By sulcus intertubercularis. Shaft of the bone round the restoration of D. bavaricum H. v. Meyer 11 pairs and smooth with weak sulcus spiralis. It starts on the of ribs were mounted. As we can see from Plate XXIV, dorsal surface above the caput and ends in front on they had 5th, 9th, 17th and 18th thoracic vertebrae. the distal part over the joint. In the distal part of the It seems they restricted themselves to the 12th. bone is trochlea humeri, with two large projections, the lateral being the higher. In front, above them, is Scapula (Pl. XII, Figs. 1 and 1a, 2 and 2a). Only the the wide and shallow fossa radialis, and on the pos- right is entirely preserved. There are separate frag- terior side – fossa olecrani. It is deeper and wider.

Table 12 Humerus D. thraceiensis sp.n. Ezerovo, D. gigantissimum D. giganteum D. giganteum D. giganteum Kaup № Dimensions, mm Bulgaria St. Manzati, Kaup Saint Kaup München – München – Stromer Romania Yean – Lartet Stromer № 495 sin dext 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 200.0 200.0 - - 260.0 190.0 humeri 8. Diameter of trochanter 202.0 207.0 260.0 108.0 260.0 210.0 major 9. Diameter of trochanter 185.0 190.0 140.0 minor

Table 13 Ulna

D. thraceiensic D. gigantissimum D. giganteum sp.n. Ezerovo, St. Manzati, D. giganteum Kaup München D. giganteum № Dimensions, mm Kaup Saint Yean Kaup München – Bulgaria Romania – – Deperet, 1892 № 495 – Stromer, Stromer, 1938 sin dext Stefanescu 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. 150.0 140.0 - - - - styloideus 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

D.thraceiensis sp.n. D.gigantissimum D.giganteum Kaup Pikermi – № Dimensions, mm Ezerovo, Bulgaria St. Manzati, Gaudry, 1862 sin dext Romania Stefanescu 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 Length of the articular 5. 160.0 160.0 90.0 - surfaces for inc. semilunaris 6. Length of same surface 130.0 132.0 - -

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

Fig. 10. Manus of Deinotherium thraciensis sp. n. A – left manus; B – right manus

32 sp.n. sp.n. sp.n. sp.n. Deinotherium thraceiensis Deinotherium thraceiensis Mc-I Mc-II Mc-III Mc-IV Mc-V Mc-IV Mc-III Mc-II Mc-I sin dext sin dext sin dext sin dext sin dext dext sin dext sin sin dext dext sin sin dext scaphoid lunare pisiforme cuneiforme magnum centrrale unciforme unciforme centrrale magnum cuneiforme pisiforme lunare scaphoid sin dext sin dext sin dext sin dext sin dext sin dext sin dext dext sin dext sin dext sin dext sin sin dext dext sin dext sin Dimensions, mm Dimensions, Dimensions, mm Dimensions, 2. Width at proximal end end 102.0 proximal 104.0 at Width Length height 1. width 2. 3. 4. radius the for facet the of Width Lateral 5. McI for facet the of Width 78.0 Anterior 6. radius the for facet the of Height 7. 77.0 tubercle the of Height 8. 78.0 9. 86.0 77.0 110.0 ulna the for facet the of Length 86.0 113.0 106.0 115.0 115.0 160.0 110.0 162.0 160.0 160.0 166.0 206.0 188.0 187.0 162.0 115.0 206.0 110.0 104.0 140.0 120.0 140.0 114.0 150.0 45.0 43.0 2. Width at proximal end end end proximal distal 112.0 112.0 125.0 125.0 140.0 140.0 145.0 146.0 98.0 108.0 at 109.0 113.0 114.0 125.0 108.0 115.0 115.0 98.0 at 97.0 1. 97.0 Length 2. Width 3. Width 4. 5. 6. processes sinew distal at Width facet small of Length facet large of Length 205.0 205.0 232.0 230.0 250.0 248.0 230.0 230.0 225.0 226.0 - - - - 104.0 - - 108.0 126.0 110.0 105.0 108.0 126.0 106.0 135.0 136.0 103.5 135.0 138.0 103.5 115.0 114.0 - 115.0 113.0 140.0 - 140.0 112.0 140.0 № № 16. Length of the facet for os lunare 160.0 160.0 lunare 160.0 lunare 160.0 98.0 os 98.0 140.0 os surface 142.0 processus for 100.0 for articular bone 100.0 150.0 the facet 97.0 facet 150.0 proximal the 98.0 of the 102.0 the of 100.0 of Length of 10. of 11. cuneiform the for facet the of Length Width 12. cuneiform the for facet the of Width Width 13. Length 14. surface articular proximal of Length Width 15. 16. 17. 66.0 65.0 140.0 140.0 150.0 150.0

Table 15 Carpals – carpus sin et dext Table 16 Metacarpals

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

, Krivoi , 1960 Rog, Moldova– Белокрые D. giganteum Kaup 1938 D. giganteum München – Stromer, Stromer, – München Kaup sp. n. sp. n. München – – München Stromer, 1938 D. giganteum

Deinotherium thraceiensis Deinotherium thraceiensis Stefanescu D. gigantissimum St. Manzati, Romania – Romania Manzati, St.

4-th digit digit 4-th digit 5-th sp.n. sin dext dext sin Ezerovo, Bulgaria D. thraceiensis thraceiensis D. 1-st digit digit 1-st digit 2-nd digit 3-rd 1 2 3 1 2 3 sin dext sin dext sin dext sin dext sin dext sin dext dext sin dext sin dext sin dext sin dext sin dext sin 1 2 3 1 2 3 1 1 2 3 3 2 sin dext sin dext sin dext sin dext sin dext sin dext sin dext sin dext sin sin dext sin dext sin dext sin dext dext sin dext sin dext sin dext sin sin dext Dimensions, mm Dimensions, Dimensions, mm Dimensions, Dimensions, mm Dimensions, 1. Length Length 1. 2. 117.5 102 116 3. end proximal 104 at Width 140 end distal at 140 Width 123 120.5 102 101 180 178 110 96 111 96 96 90 98 89 87 88 - 95 - 96 90 96 90 96 80 96 80 97.5 - - 1. 2. d-c wing of Length 3. h-d wing the of length Total 4. obturatum for. to wing of Length 5. obturatum for. of Length 6. obturatum for. of Width 7. acetabulum the at Diameter 8. process sacral of Height 820.0 1160.0 9. symphysis pelvic of Length same of Height 1110.0 1175.0 740.0 266.0 1165.0 190.0 126.0 280.0 462.0 320.0 1180.0 188.0 900.0 150.0 1130.0 460.0 320.0 385.0 - - - 275.0 - 830.0 - - 1000.0 - - 560.0 - - 780.0 - 470.0 - - - - 340.0 - - 245.0 - 230.0 250.0 - № № 3. Width at distal end end 125 125 end 91 83 - 82.6 88 - distal 91 66 proximal 94 95 97 103 68 - - 102 105 104 107 106 107 - - 107 115 95 at 116 108 at 95.5 108 111 112 Length 1. Width 2. Width 3. 100 110 110 100 98 96 150 150 80 130 80 130 178 180 105 102 167 167 №

Table 17 Bones of the digits – forelimb (continued) Table 17 Bones of the digits – forelimb Table 18 Pelvis

35 crest. The wing significantly narrows posteriorly. tened posteriorly. The larger part of the bone was Dorso-lateral surface smooth and slightly concave, covered by a cartilage with the shape of the thigh- medial almost flat. In the region of facies articularis bone’s surface it articulated with. the surface is rough. Acetabulum well shaped. Fora- men obturatorium oval. From the lower hindlimbs, only distal part of the left tibia and fragments of the proximal parts of the two Femur (Pl. XXI, Fig. 1 and 1a). We found only the fibulae were found. right thighbone. Left was restored after it. The bone is long and solid. Caput femuri large, almost paral- Tibia (Pl. XXII, Fig. 2 and 2a). Although the bone lel to the bone’s axis. A small neck separates it from was restored to its full size, we’ll discuss only its dis- trochanter major, which is laterally situated. Fossa tal part which is original. Down of the diaphysis the trochanterica strongly pronounced. Shaft flat, slightly bone becomes triangular. It gradually widens, form- curved medially and laterally. Two condyles in the ing at the distal end the cochlea tibiae. Maleolus distal part of the bone. Condylus medialis larger and medialis strongly developed, incisura fibularis deep some higher than condylus lateralis. Fossa inter- and wide. condylaris which separates them is wide and deep. Fossa muscularis cranialis and F. m. caudalis deep Fibula. The original fragments used for the restora- and parallel to the condyles. Longitudinal crests sur- tion are from the proximal part. This is a long and rounding trochlea patellaris high and rounded in narrow bone. Almost flat in its proximal part. Thick their uppermost part. at the beginning, articular surfaces unclear.

Patella (Pl. XVIII, Fig. 4). Pear shaped. Concave Comparison. There are no differences of taxonomi- anteriorly. In the middle it is convex like a hemi- cal value in the hipbones and the bones of the lower sphere surrounded by a deep and wide ring. Flat- leg.

Table 19 Femur

D. thraceiensis sp.n. D. gigantissimum St. D. giganteum Kaup D. giganteum Kaup № Dimensions, mm Ezerovo, Bulgaria Manzati, Romania – München – München – sin dext Stefanescu Stromer, 1938 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, D. gigantissimum St. D. giganteum Kaup № Dimensions, mm Bulgaria Manzati, Romania – Pikermi – Gaudry, 1862 sin dext Stefanescu 1. Total length 1200.0 1250.0 920.0 950.0 2. Width at proximal end 370.0 365.0 330.0 310.0 3. Thickness at proximal end 320.0 320.0 – – 4. Width in diaphysis 160.0 165.0 260.0 140.0 5. Width at distal end 300.0 330.0 260.0 340.0

Table 21 Fibula

D. thraceiensis sp.n. D. gigantissimum St. D. giganteum Kaup № Dimensions, mm Ezerovo, Bulgaria Manzati, Romania – Pikermi – Gaudry, 1862 Stefanescu sin dext 1. Total length 1125.0 565.0 830.0 870.0 2. Width at proximal end 90.0 72.0 70.0 70.0 3. Width in diaphysis 126.0 103.0 - 100.0 4. Width at distal end 275.0 250.0 170.0 140.0 Length of the articular surface for os 5. 80.0 72.0 - 70.0 calcaneus 6. Length of the small facet 76.0 60.0 - 55.0

36 Table 22 Tarsals – tarsus sin et dext

Deinotherium traceiensis sp.n. Cuneiforme Cuneiforme Cuneiforme № Dimensions, mm Astragalus Calcaneus Naviculare mediale intermedium laterale sin dext sin dext sin dext sin dext sin dext sin dext 1. Diameter of the corpus 313.0 310.0 2. Proximal height 90.0 90.0 Average thickness of the 3. 130.0 130.0 190.0 200.0 corpus 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) lanx. In D. thraceiensis sp. n. the last phalanx - the Consists of seven bones. Two in the proximal row: hoof – in the hindlimbs is large, as with the fore- calcaneus and astragalus; five in the distal: cuboid, limbs. In the compared species they are much small- three cuneiforms and, between them, navicular. Only er. They are anteriorly situated like nails and touch the calcaneus was found. The rest were restored af- the ground only with the anterior surface, or with a ter D. gigantissimum Stefanescu. The toes were re- small part of the lower surface. stored, too. It can be seen from the description that D. thraceiensis Comparison. There are no differences in the struc- sp. n. has numerous characters distinguishing it from ture of the tarsals in the various deinothere species. all the other species. Here are some of them: There are such however in their digits. In D. thra- 1. Skull – short, high and, compared to the size of ceiensis sp. n. the size of the first and second pha- the body – very small. lanxes gradually decreases. In D. gigantissimum Ste- 2. External nares large and deep, with a peculiar fanescu all digits have large first phalanxes, the sec- shape. ond are much smaller and the third – even more. 3. Nasal bones short, narrow and high, frontally fused. There is a big difference in their size. D. bavaricum 4. Forehead high and wide. H. v. Meyer and D. giganteum demonstrate a much 5. Occipital bones high and wide, at an angle of 80° more gradual transition from the first to the last pha- to the forehead.

Fig. 11. Pes of Deinotherium thraciensis sp. n. A – left pes; B – right pes

37 sp.n. sp.n. sp. n. sp. n. Deinotherium thraceiensis Deinotherium thraceiensis Deinotherium thraceiensis 4-th digit digit 4-th digit 5-th Mt-I Mt-II Mt-III Mt-IV Mt-V Mt-IV Mt-III Mt-II Mt-I 1 2 3 1 2 3 2 1 3 1 2 sin sin dext dext sin dext sin dext sin sin dext 1-st digit digit 1-st digit 2-nd digit 3-rd sin dext sin dext sin dext sin dext sin dext sin dext dext sin dext sin dext sin dext sin dext sin sin dext 1 2 3 1 2 3 3 1 2 sin dext sin dext dext sin dext sin sin sin dext dext sin dext dext sin dext sin dext sin sin dext Dimensions, mm Dimensions, Dimensions, mm mm Dimensions, Dimensions, mm Dimensions, 3. Width at distal end end end 104 85 86 - - 103 92 103 92 111.5 distal 110 proximal 93 94 - 108 105 89 87.5 110 110 117 117 - 106 at at Length Width 1. Width 2. 3. 103 103 104 147 104 146 125 125 105 105 169 170 № 2. Width at proximal end end 12 105 105 122 94.5 94 proximal 112 112 105 104 98 at 98 Length 111 110 1. 117 116 107 107 Width 2. 3. end distal at Width 136 107 107 136 130 130 100 160 160 100 100 142 142 125 125 180 180 165 167 100 92 92 - - 101 101.5 87 87.5 - - 104 104 95 94 - - № 2. Width at proximal end end end proximal distal at 137.0 at 138.0 120.0 120.0 125.0 120.0 Length 114.0 146.0 114.0 146.0 Width 1. 120.0 120.0 Width 115.0 114.0 115.0 115.0 115.0 116.0 115.0 2. 116.0 3. 4. processes 5. sinew distal at Width 6. facet small of Length facet large of Length 135.0 212.0 212.0 240.0 238.0 225.0 225.0 220.0 222.0 220.0 222.0 136.0 131.0 126.0 112.0 130.0 112.0 126.0 102.0 - - 103.0 - 112.0 - 112.0 - - 117.0 - 117.0 ------№

Table 24 Bones of the digits – hindlimb Phalangi sin et dext Table 24 A Bones of the digits – hindlimb Phalangi sin et dext Table 23 Metatarsals – metatarsus sin et dext

38 6. Eye socket differentiated and large. skeleton in Vienna, and another of D. giganteum 7. Occipital condyles low. Kaup. We know also the paintings by Augusta and 8. P3 and P4 differently structured. Burian of D. giganteum Kaup, and the painting of 9. Ramus horizontalis strongly protruding backwards D. gigantissimum Stefanescu exhibited in the Bucha- at angulare. rest Museum in front of the skeleton itself. It is made 10. Processus articularis and p. coronoideus high. following the first two authors. Everywhere the skull 11. Incisura mandibulae deep. with the mandible takes about 1/3 of the whole skel- 12. Tusks differently structured. eton. The neck is thick, merging with the vertebral 13. P3 differently structured. column because, according to them, the last cervi- 14. Curve of the symphysis much more regular. cals have high spinous processes. Frontally, the chest 15. Spinous processes on the cervicals shorter and is narrow, and the body – solid, with high legs. The weaker; first cervicals lack processes. latter are narrower near the fingers because of the 16. Spinal canal widening at the lumbar region. small hoofs. 17. Spinous process on second thoracic vertebra In D. thraceiensis sp. n. the skull and the mandi- strongly developed. ble are much less than 1/3 of the whole skeleton. The 18. 14 ribs. neck is short and only the last cervicals have spinous 19. Manus and pes high and spread. processes. They are very high on the first thoracic 20. Hoofs larger and entirely ‘lying’ on the ground. vertebrae, widening at the tip. This permits us to sup- pose it had a high hump like Elephas. On the basis of these characters, we believe that On the other hand, as far as the first rib is wide the specific status of D. thraceiensis n. sp. is entirely and solid, its chest was probably broad. The digits proved. To this conclusion we shall add some short were more spread. notes on its anatomy. For the time being, we cannot comment on the We are acquainted with the reconstructions by problems of the phylogeny of genus Deinotherium. Prof. Abel of D. bavaricum H. v. Meyer, based on the They could become the subject of a future work.

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