kovachev, 2006 - deinotherium thraceiensis sp nov
DESCRIPTION
mammuth descriptionTRANSCRIPT
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GEOLOGICA BALCANICA, 35. 3—4. Sofia, Dec. 2006, p. 5—40.
Introduction
The fossil materials presented herewith to the scien-tific community were found by Dimitar Kovachev in1965. The late Ivan Nikolov has worked for a longtime on their restoration, with minor participation ofD. Kovachev. In 1972 Nikolov reported to the De-partment of Palaeontology at the Geological Insti-
tute, Bulgarian Academy of Sciences, that the resto-ration process was finished. The assembled skeletonhas been mounted and permanently exhibited in theMuseum of Geology and Palaeontology, Sofia Uni-versity “St. Kliment Ohridski”. A diminished (3/4 ofnormal size) copy has been prepared and is now onexhibit at the Asenovgrad Palaeontological Branchof the National Natural History Museum. Both Ivan
Deinotherium thraceiensis sp. nov.from the Miocene near Ezerovo, Plovdiv District
Dimitar Kovachev1, Ivan Nikolov2
1 Asenovgrad Palaeontological Branch, National Natural History Museum, Asenovgrad2 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 îòëîæåíèéáëèçè 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, èîïèñàííîãî, 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 microtatonNicolov è ÷àñòü ÷åëþñòè Mastodon sp. Ïðè âñêðûòèè ñêåëåòà òîëüêî íåìíîãèå êîñòè íåõâàòàëî, à èìåííî, çàäíàÿ ëåâàÿ áåäðåííàÿ êîñòü, çàäíàÿ ëåâàÿ ôèáóëà, áîëüøèíñòâî ðåáð èíåêîòîðûå èç êàóäàëüíûõ ïîçâîíêîâ. Îíè ðåñòàâðèðîâàííûå íà îñíîâå ñóùåñòâóþùèõ ñèì-ìåòðè÷åñêèõ àíàëîãîâ. Ðàçìåðû êîñòåé è ñðàâíåíèå ñ äðóãèìè äåéíîòåðèÿìè îïèñàííûìèâ ëèòåðàòóðå ïîêàçàëî íåêîòîðûå ñóùåñòâåííûå ðàçëè÷èÿ. Íåêîòîðûå ðàçëè÷èÿ ïî îòíî-øåíèè ÷åðåïà èìåþò áåç ñîìíåíèÿ òàêñîíîìè÷åñêîå çíà÷åíèå.  ñâÿçè ñ ýòèì, îïèñàííûéäåéíîòåðèé îòíîñèòñÿ ê íîâîìó âèäó Deinotherium thraceiensis sp. n.
Abstract. The paper presents to the international palaeontological community a description ofthe 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 remainsare 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 wholerear left thighbone, the right rear fibula, most of the ribs as well as some caudal vertebrae. Theyare restored on the basis of symmetrical analogues existing. The measurements on the bonesfound and the comparisons with other deinotheres described in the literature showed certaindifferences. 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 nearEzerovo, Plovdiv District. – Geologica Balc., 35, 3—4; 5—40.
Key words: Deinotherium thraceiensis, Miocene, Maeotian, Bulgaria.
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Nikolov and Dimitar Kovachev were fully aware ofthe importance of this very rare case of finding awhole skeleton of such a huge extinct animal, andthey estimated it to be a representative of a new spe-cies of genus Deinotherium. Ivan Nikolov, well-knownfor his thoroughness, wanted to make an exemplarypublication but his early death stopped for a longtime the work on it. The results of his enormous la-bour put into the description of the bones, the resto-ration and mounting of this unique skeleton, re-mained in the archives of the Geological Institute.After the sudden untimely death of Nikolov in 1984,the work on the publication remained unfinished.Now, D. Kovachev decided to prepare the materialsfor publication with the clear idea about the respon-sibility undertaken, and that the whole blame for even-tual flaws and errors should be addressed to himself.Such publication would be of importance for thefuture studies on genus Deinotherium.
Many scientists have helped the authors in dif-ferent time. Prof. Dr. R. Dehm (Director of Univer-sität-Institut für Paläontologie und historische Geolo-gie – München) and Prof. Dr. H. Tobien (Directorof Paläontologisches Institut der Johanes-GutenbergUniversität – Meinz) helped I. Nikolov during thefirst studies and restoration. Academician T. Nikolov(Geological Institute, Bulgarian Academy of Scienc-es) gave valuable advice with the material. D. Ko-vachev wishes to express his gratitude to Dr. MarinIvanov from the Sofia University for his responsive-ness to the problems of this research. Dr. N. Spassov(National Natural History Museum, Sofia) kindlyhelped with the literature I. Nikolov had used at thelibrary of the Museum.
Studies on the genus Deinotherium –a historical review
As early as the 17 century a place near Lyon, France,was known as “the field of the giants” because of thelarge animal bones often found there. Some of thosebones came to Matsorier – a surgeon, who used toshow them for years in France and Germany as thebones of king Töteboch. Much later, the real tomb ofthe king was found, and the deceit was exposed. Thehuge bones were transferred to the Natural HistoryMuseum in Paris. Probably these are the first Deinothe-rium remains found.
One century later, in 1715, Réaumure admittedhe could not refer the bones to any known animal.Kenedy supposed in 1775 that the materials were re-lated to mammoths. G. Cuvier thought in 1779—1836that the animal had been a large tapir coexisting withmastodons. He called it Tapir gigantesque andthought that its tusks were curved upwards.
Kaup created in 1829 the genus Deinotherium withthe species D. giganteum upon the skull and mandi-ble found near Eppelsheim, Germany. He thoughtthis was an intermediary form between the sloth andthe mastodons and referred them to the larger taxo-nomic unit Curtognati. In 1841 and 1857 he de-
scribed further remains of that genus found nearEppelsheim and Westhofen. According to him, theadult animals had five teeth in their upper and low-er jaws. In 1833 he described the new species D. cu-vieri, but because it was often found in Bavaria, H. v.Meyer called it D. bavaricum. This name is still usedtoday. However, arguments about the exact place andthe life environment of deinotheres still continued.
P. Gervais accepted in 1848 the opinion of Kenedyand Koch that the genera Deinotherium and Masto-don should be referred to Proboscidea. He recognizedthe existence of three species: D. giganteum Kaup =Tapir gigantesque Cuvier; D. intermedium and D. cu-vieri Kaup. S. Pictet reported in 1853 about the findingnear the village of Absdorf of a whole skeleton of D.giganteum Kaup. Only the head, the first two vertebrae,the corpus of a thoracic and a caudal vertebra andfragments of the limbs were preserved. These remainsshowed that it is close to the mastodon and belongs toPachydermae. Four deinothere species were recognizedin 1858—1859 by M. Lartet, and namely, Deinotheriumbavaricum H. v. Meyer, D. giganteum Kaup, D. sp. (anintermediary form between the first two), and D. cu-vieri Kaup. The first species is known from the Mi-ocene of France and Bavaria, the second from the LateMiocene of France, Germany and Greece, the thirdand fourth from the Miocene of France. After Lartet,the dental formula of genus Deinotherium is: I 0-0/1-1;Pm 2-2/2-2 and M 3-3/3-3 for the permanent teeth,and for the deciduous – I 0-0/1-1; deciduous molars3-3/3-3. New fossil finds in Greece (Pikermi), India (Si-walik), Europe (France, Switzerland, Austria), etc. havebeen reported during the last quarter of the 19th centu-ry by A. Gaudry, H. Hensel, B. A. Lydekker, E. Chantre,M. Vacek, V. Biber, C. Deperet, and other authors. Gr.Stefanescu first reported in 1891, and then described(1895–1910) a deinothere skeleton found near Man-zati, Romania. Because of the large dimensions of allits bones and teeth, he named it D. gigantissimum. Ehik(1930) published upper and lower molars and somefinger bones, found in the Pliocene sediments of Hun-gary, as a new species – Prodinotherium hungaricum.Osborn (1936) changed the name of the genus (theLatinized form Dinotherium) to the ancient Greekform Deinotherium. He revised all then knowndeinothere materials, discussing in details each spe-cies, and illustrating skulls/mandibles and teeth fromD. giganteum Kaup, D. bavaricum H. v. Meyer, D.cuvieri Kaup, D. pentapotamiae Lydekker, D. indi-cum Falconer, etc. (Fig. 1).
Numerous new finds of deinotheres have been re-ported during the whole 20th century from Austria,Bohemia, Bulgaria, France, Germany, Greece, Hun-gary, Macedonia, Moldova, Pakistan, Romania, Rus-sia, Serbia, Ukraina, etc.
Studies on the genus Deinotheriumin Bulgaria
The first Deinotherium remains in Bulgaria have beenfound by G. Bonchev (Áîí÷åâ, 1897) who made ex-
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Fig. 1. Distribution of genus Deinotherium in Southeastern Europe and Asia Minor
cavations in the Middle Sarmatian limestones nearNessebar, studying the hipparion fauna. Three yearslater he reported new finds from the same locality.Áàêàëîâ (1911/1913) published the two deinotheremandibles excavated by G. Bonchev, together with
other single teeth from the village Sovolyano, nearPleven, the village Archar and the village Mazgosh.They were all brought to the Institute of Geology bydifferent people. From these materials Bakalov de-termined two species – D. giganteum Kaup race mi-
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nor and D. giganteum Kaup race major. To the firsthe referred the smaller teeth, and to the second –the larger ones. Later on (Áàêàëîâ, 1949/1950)Bakalov described Deinotherium teeth found at the“Meander of the Konska river” near the village No-evtsi; the village (now – a town; for some time calledBatanovtsi) Temelkovo (Pernik region) and Aksako-vo, near Varna, as D. giganteum Kaup race major(from the first two localities, Pliocene) and D. gigan-teum Kaup race minor (from the Sarmatian lime-stones near Aksakovo). Íèêîëîâ (1960) reported amolar fragment of D. giganteum Kaup race majorfrom the lignite coals near the village Hrabarsko (So-fia region), assuming a Pontian age. The whole fossilmaterial belonging to genus Deinotherium then knownfrom Bulgaria has been published by Áàêàëîâ,Íèêîëîâ (1962). They recognize two races – minorand major, and refer the finds as follows: two man-dibles, an upper jaw and three separate teeth to raceminor, and a mandible, seven complete upper andlower teeth and three tooth fragments to race major.The age of both races varies from Sarmatian to Le-vantian. Íèêîëîâ (1962) referred the molar and frag-ment of a scapula found in the Levantian sedimentsnear the village Popitsa, Vratsa region to D. gigan-teum Kaup. Íèêîëîâ, Êîâà÷åâ (1966) described alower molar found by the latter author in the sandsin the valley of Cherkezitsa river, near the villageAhmatovo. They assume the age to be Levantian be-cause of the Anancus arvernensis Croizet et Jobertteeth found there, referring the find to D. giganteumKaup. Êîâà÷åâ (1966) reported the finding of analmost complete skeleton of Deinotherium near thevillage Ezerovo, Plovdiv region. This same skeleton isthe subject of the present paper. Öàíêîâ, Íèêîëîâ(1966) published a brief review of the proboscideansin Bulgaria, and gave also some preliminary infor-mation about the finding of Kovachev reportingmetrical data on some bones.
Distribution of genus Deinotheriumin Southeast Europe and parts of AsiaMinor
Until now, more than 80 localities (partly shown onFig. 1) within the region have yielded deinothere re-mains, and their number is constantly growing.
There are 16 localities from former Yugoslavia:1. Veles, Mokranje, Smederevska palanka, Resnik,Meduhana, Vracevic, Duboko, Tsiganski potok, andbetween the villages Viteževo and Porodin – D. gi-ganteum Kaup. Near Veles and Ravanica, togetherwith D. giganteum Kaup, D. bavaricum H. v. Meyeris also present. The sediments near the villages Medu-hana, Ravanica and those between Viteževo andPorodin are of Lower Sarmatian, and the rest – ofPontian age.2. Drenovica, Chukovica, Bresnica, Prebrezka, Zdar-ski and Kriva reka – D. bavaricum H. v. Meyer. Theage cited is Helvetian-Tortonian, Lower Sarmatian,Sarmatian and Middle Miocene.
In Greece D. giganteum Kaup is known from fourlocalities: Pikermi, Thessaloniki and Tanagra (Pon-tian), and Samos (Middle Pontian). From the Sar-matian of Chios D. bavaricum H. v. Meyer has beendescribed.
From Turkey (Asia Minor), D. giganteum Kauphas been described from Küçükçekmece (Maeotian).
From the territory of Romania and part of Bessa-rabia, two taxa are known – D. giganteum Kaup fromthe Pontian and Dacian sediments near Vernesti,Culm, Lichtental, Pelineu, Telenesti, Visterniceni,Baimaclia, Cimislia, Comanesti, Curtea and Solcus-tria; and D. gigantissimum Stefanescu, – from the Plio-cene sediments near Manzati, Breaza and Cimislia.
From the territory of Moldova and Bessarabia, twotaxa are known too: 1. D. giganteum Kaup near Tara-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 312/22. 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
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PLATE I PLATE II
PLATE III PLATE IV
PLATE V
PLATE VI
PLATE VII
PLATE VIII PLATE IX
PLATE X PLATE XI
PLATE XII PLATE XIII
PLATE XIV PLATE XV
PLATE XVI PLATE XVII
PLATE XVIII
PLATE XX
PLATE XIX
PLATE XXI PLATE XXII
PLATE XXIII
PLATE XXIV
92 Geologica Balcanica, 3-4/2006
Fig. 2. Distribution of genus Deinotherium in Bulgaria: 1. Ezerovo; 2. Nesebar; 3. Izgrev; 4. Parvomai; 5. Ahmatovo; 6. Lyubovishte;7. Sovolyano; 8. Batanovtsi (Temelkovo); 9. Noevtsi; 10. Sofia; 11. Kremikovtsi; 12. Hrabarsko; 13. Aldomirovtsi; 14. Katina; 15.Kula; 16. Archar; 17. Mihaylovo; 18. Glozhene; 19. Popitsa; 20. Pleven; 21. Novo selo; 22. Aksakovo; 23. Yarebichna; 24. Breznik;25. Hadjidimovo; 26. Varna – Galata; 27. Kalimantsi; 28. Konyovo; 29. Rogozen; 30. Gabra – former open pit “Bolshevik”; 31.Mazgosh (now on Serbian territory)
PLATE V
Deinotherium thraceiensis sp. n.1. Mandible. Scale ca. 8.3% coll. SU.M. No. SU Pl 312/2; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE VI
Deinotherium thraceiensis sp. n.1. Lower cheek teeth – P3 to M3. Scale ca. 25% coll. SU.M. No. SU Pl 312/2; Locality: Ezerovo, near Plovdiv; Level: Maeotian
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rasspol, Donau, Velikovo and Krivoy Rog. The sedi-ments are of Maeotian, even Upper Sarmatian age.2. D. gigantissimum Stefanescu near Volchinec, Rah-ny-Lessovye and Pripecheni. The host sediments arealso of Maeotian and Upper Sarmatian age.
A considerable number of deinothere localitiesare known from the territory of Bulgaria (Fig. 2).Some of the first Deinotherium finds in Bulgariacame from the locality Mazgosh (now in Serbia).The larger number of localities in the western partof the country is a result from the fact that thispart is better studied. Some of the more importantfinds come from Nessebar (by G. Bonchev), No-evtsi, locality “Meander of the Konska river” (by P.Bakalov), and the village Ezerovo, “Kolnu dere”(by D. Kovachev).
Some of the materials have been found andbrought to the specialists by common people (farm-ers, workers, etc.). Therefore, the exact location hasbeen mentioned only approximately, and there is nostratigraphic profile. In such cases, a probable age
has been assumed after the age of the sediments inthe noted areas.
The vertical distribution of deinotheres in Bul-garia starts in the Sarmatian, includes the Maeotianand Pontian and ends in the Pliocene. The remainsof D. bavaricum H. v. Meyer are found only in Sar-matian sediments, while D. giganteum Kaup is presentfrom the Middle Sarmatian up to the Romanian.
Stratigraphic notes on the sedimentsnear Ezerovo, Plovdiv region
The village of Ezerovo is situated ca. 30 km east ofPlovdiv (Fig. 3). Sediments of Eocene, Miocene,Pliocene and Pleistocene Age crop out in the area(Fig. 4).
The Upper Eocene (Priabonian) is represented bylight organogenic limestones, rich of nummulites andcorals. South of the village the dam wall is built ontothem. The thickness exposed is about 50 m. In a bore-
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Fig. 4. Stratigraphic profile of the locality Ezerovo
Fig. 3. Position of the locality of Ezerovo in the Upper Thracian plain
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hole (No 16; Áðúíêèí, Ñòàí÷åâà, 1965) situatednear the village of Ezerovo, these rocks are coveredby massive limestones, marls, and well-cementedquartz sandstones. They are referred to the Oligoceneon the basis of rich foraminifer and ostracod fauna.These rocks pass into the Lower Miocene dark grayaleurolites and marls with foraminifer and ostracodfauna similar to the Aquitanian-Burdigalian faunasreported in France (Áðúíêèí, Ñòàí÷åâà, 1965). Themaximum thickness of the Lower Miocene is about180 m.
The Palaeogene and Lower Miocene marine sed-iments are unconformably covered by fluvial sedi-ments referred to the Ahmatovo Formation (Êîþì-äæèåâà, Äðàãîìàíîâ, 1979). They crop out at nu-merous places in the valley of the river Cherkezitsa.At the basis there are aleurolites and shales coveredby sands of light rusty colour, alternating with con-glomerate lenses. At places the sand layers are cross-bedded. Petrified trees occur in the sands. The thick-ness of the sand layers varies from 1.5 m to 7.5 m;and that of the conglomerates is up to 1.5 m. Thewhole thickness of the sands has been revealed onlyby drillings and is up to 515 m. Ahmatovo Forma-tion is referred to the Maeotian, Pontian and Da-cian regional Paratethys stages. Three macrocycleshave been distinguished (Êîþìäæèåâà, Äðàãîìà-íîâ, 1979). The first macrocycle is referred to theMaeotian (possibly also upper parts of the Sarma-tian) on the basis of Lower to Middle Turolian (“Pik-ermian”) mammal fauna. The Pontian age of thelower parts of the second cycle is determined by UpperPikermian (= Upper Turolian) fauna, and the up-per parts of the second cycle are referred to the Da-cian Regional Stage by Ruscinian fossil remains. Thefossil locality of Deinotherium thraceiensis sp. n. be-longs to the first cycle of the Ahmatovo Formation(Nikolov, 1985, p. 58).
About 800 m west of the village of Ezerovo, rightfrom the road to the dam, at the fork with the roadfor the pumping station, in light medium-grainedsands, D. Kovachev found in 1965 the skeleton ofDeinotherium thraceiensis sp. n. described in thepresent paper. It was lying at a depth of ca. 40 mrelatively to the level of borehole No 16. At the samelevel, but 50 m to the east, V. Tsankov and I. Vaptzarovhave found in 1956 a part of a skull, determined byP. Bakalov as Trilophodon angustidens Cuvier. In 1962St. Stoykov and I. Nikolov found 4—5 m below thefind described here, teeth of Hipparion microtatonNikolov and a part of a mandible of Mastodon sp.They all indicate a Maeotian age. Above thedeinothere have been found teeth of Hipparion med-iterraneum Gervais and Tetralophodon longirostrisKaup. They unequivocally prove a Pontian age.
After 1965, D. Kovachev collected the followingfossils during numerous field trips:1. A skull, a mandible and bones from the limbs ofMastodon sp. In the structure of its molars it resem-bles T. angustidens Cuvier, but the tusks and the skullare different from this species.
2. A whole mandible with a preserved symphysis andM3 dext et sin of Choerolophodon sp.3. Two maxillary fragments with M3 also belongingto Choerolophodon sp.4. M3 sin et M2 dext – Choerolophodon sp.5. Right semimandible with M3 – Tetralophodon lon-girostris Kaup.6. M3 sin – Anancus sp.7. M2 and M3 dext and M3 sin – Zygolophodon bor-soni Hays.8. M3 dext – Zygolophodon borsoni Hays.9. M3 sin Deinotherium giganteum Kaup.10. Right semimandible – Microstonyx major.
They will be subject of other publications.The Neogene sands are covered by a layer of soil
1.0 to 1.5 m thick including isolated rock fragmentsof different size. These soils are referred to the Pleis-tocene.
Palaeontological part
Order Proboscidea Illiger, 1821Suborder Deinotherioidea Osborn, 1921Family Deinotheriidae Bonaparte, 1845Genus Deinotherium Kaup, 1829Type species of the genus Deinotherium giganteumKaup, 1829
Deinotherium thraceiensis sp. n.
(Pl. I – XXII)Derivatio nominis: thraceiensis from Greek “Thra-ceia” – Thrace, the region of the find.Holotype. As such we design all the bones of the skel-eton – No. 312/1 to No. 312/23-30, collection of theMuseum of Geology and Palaeontology of the SofiaUniversity “St. Kliment Ohridski”.
Locality. Light, medium-grained sands (AhmatovoFormation, first cycle) near Ezerovo, Plovdiv region,just at the turn for the pumping station from theroad to the dam.
Age. Maeotian.Diagnosis. Large animals, skull short and high. Na-sal bones short, fusing at the anterior ends, curvedand slightly flattened laterally. Large external nares.Ear area also large and with trapezoid shape. Eyesockets almost separated from the ear areas. Fore-head high and, compared to the external nares –short. The occipital bone is high and wide. Symphy-sis of the mandible curved downwards and back-wards. Incisors of moderate length, slightly curvingoutwards. Their tips end exactly beneath processusangularis. Neck short, with seven thin cervical verte-brae. First ones lack processus spinosus, in the lastones it is weakly developed. Skull and mandible, rel-atively to the size of the whole skeleton, are muchsmaller than in the other proboscideans. Fingers high,with large phalanxes – hoofs, strongly developed onthe third finger.
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Fig. 5. Deinotherium thraceiensis sp. n.
Description and comparison
Skull. (Pl. I, fig. 1 and 2; Pl. II, fig. 1 and 2; pl. III, fig.2; Fig. 5 – A, B, C and D).The skull described here is entirely preserved. It isvery short and its posterior part – high (Fig. 5).The premaxillary bones (os premaxilarae), which arestrongly developed in Proboscidea, do not merge attheir anterior ends and are slightly curved downwards.There are no upper tusks and alveoli for them inthese bones. They are widening in front of the eyesockets. All their surface is smooth, without any rug-osity.Nasal bones (os nasale) are short, high in their ante-rior part, rounded and laterally flattened. On theinner side, in the region of the maxillary bones, thenasals are slightly concave, so that together with thepremaxillary bones they have a rounded, wedge-likeshape (Fig. 5, A). They are connected by a strong su-ture. The nasal canal has an inner width of 70.0 mmand length to the end of the external nares 710.0 mm.External nares (Fig. 5, ex. n., and Pl. II, figs 1 and 2)pear-shaped and rather large. They start almost atthe beginning of os nasale, gradually becoming wid-er and deeper. By the beginning of the eye socketthey narrow, then becoming straight and ending witha regular, arch-shaped curve just at the beginningof the ear area. They penetrate for 70–80 mm moreand end in the oscular area with the dorsal nasalopening, which is small and oval, parallel to the fore-head. On the bottom, parallel to the nasal cavity, cris-
ta conchalis dorsalis is outlined. (Measurements ofthe external nares in Table 1).The calvaria – os frontale and os parietale (fronto-parietal part) starts against the beginning of the eararea ends with os occipitale. In the middle it is slightlyconcave, and in the posterior part – strongly convex(measurements in Table 1).Occipital bone (os occipitale). (Fig. 5 A, C, D ocs;Pl. III, Fig. 2). High and wide, laterally slightly round-ed, under the condyles and by the parietal bone al-most straight and parallel. On both sides its wingsare curved backwards, and in the region of thecondyles and above them it is slightly convex. In re-lation to the forehead this bone is almost perpendic-ularly situated, at an angle of 80° (measurements inTable 1).
Occipital condyles. (Fig. 5; Pl. I, Fig. 1 and 2).They are situated in the middle and in the lower partof the occipital bone. Their shape is irregular andoval. Wider and convex in their upper part. Theysurround foramen magnum – fm, which has an ovalshape and is situated perpendicularly to the occipi-tal bone (measurements in Table 1).Eye socket (orbita) (Fig. 5 A orb.) is preserved onlyon the right side of the skull. In front it is limited bythe massive and laterally strongly protruding os lac-rimale, and in the back – by the well pronouncedprocessus postorbitale, which is long and thin. Frombelow it is limited by os zygomaticum. Thus the eyesocket is almost entirely separated from the ear area(measurements in Table 1).
13
Table
1
Skulls – Crania
№
Dim
ensi
ons,
mm
D
.thra
ceie
nsis
E
zero
vo
D.g
igan
teum
Kau
p E
ppel
shei
m
D.g
igan
teum
Kau
p M
unic
h D
.gig
anti
ssim
um
Stef
anes
cu M
anza
ti
D.b
avar
icum
H
. v. M
eyer
B
avar
ia
1.
Tot
al le
ngth
13
20.0
12
1.0
1035
.0
1260
.0
860.
0 2.
L
engt
h fr
om o
s na
sale
to c
ond.
occ
ipita
lis
1290
.0
900.
0 95
0.0
1118
.0
780.
0 3.
L
engt
h fr
om o
s na
sale
to o
s oc
cipi
tale
10
12.0
84
0.0
- -
- 4.
W
idth
at t
he b
asis
of
os o
ccip
itale
16
5.0
240.
0 25
2.0
270.
0 -
5.
Wid
th a
t the
bas
is o
f th
e ex
tern
al n
ares
(na
sale
) 57
0.0
- 10
8.0
- -
6.
Wid
th in
the
mid
dle
of th
e ex
tern
al n
ares
(na
sale
) 68
0.0
- 17
0.0
-
7.
Wid
th in
eye
soc
kets
73
0.0
- 50
0.0
- -
8.
Wid
th in
ear
are
a
540.
0 57
0.0
- -
- 9.
H
eigh
t of
the
skul
l at M
1 71
5.0
- 36
0.0
- -
10.
Hei
ght o
f th
e sk
ull a
t M3
805.
0 55
0.0
380.
0 63
0.0
385.
0 11
. L
engt
h of
ear
are
a 38
0.0
530.
0 -
540.
0 26
0.0
12.
Hei
ght o
f ey
e so
cket
18
5.0
13.
Wid
th o
f ey
e so
cket
12
0.0
14.
Dep
th o
f ey
e so
cket
16
0.0
-
15.
Len
gth
of o
s na
sale
to th
e an
teri
or e
nd o
f P3
350.
0 32
4.0
330.
0 43
0.0
300.
0 16
. W
idth
of
os n
asal
e in
pre
max
illar
ia
260.
0
17
. W
idth
of
os n
asal
e to
the
ante
rior
end
of
P3 13
0.0
18.
Wid
th o
f os
nas
ale
in th
e m
iddl
e be
twee
n th
ese
two
poin
ts
185.
0
19
. M
inim
um h
eigh
t of
os n
asal
e 25
0.0
160.
0 10
8.0
210.
0 -
20.
Beg
inni
ng o
f ex
tern
al n
ares
fro
m th
e be
ginn
ing
of o
s na
sale
15
0.0
21.
Wid
th o
f ex
tern
al n
ares
at t
he f
irst
cur
ve
160.
0
22
. W
idth
of
exte
rnal
nar
es a
t the
sec
ond
curv
e 30
0.0
23.
Wid
th o
f ex
tern
al n
ares
at t
he e
ye s
ocke
t 36
0.0
24.
Dep
th o
f th
e ex
tern
al n
ares
at t
he f
irst
cur
ve
154.
0
25
. D
epth
of
the
exte
rnal
nar
es a
t the
sec
ond
curv
e 16
5.0
26.
Dep
th o
f th
e ex
tern
al n
ares
in th
e m
iddl
e 50
0.0
27.
Len
gth
of o
s fr
onta
le
308.
0 -
- -
- 28
. M
inim
um w
idth
of
os f
ront
ale
500.
0 -
- -
- 29
. M
axim
um w
idth
of
os f
ront
ale
540.
0 -
- -
- 30
. L
engt
h of
os
occi
pita
le
850.
0 -
- -
- 31
. H
eigh
t of
os o
ccip
itale
70
0.0
450.
0 -
420.
0 45
0.0
32.
Hei
ght o
f co
ndyl
us o
ccip
italis
19
6-si
n 17
6-de
xt
33.
Max
imum
wid
th o
f co
ndyl
us o
ccip
italis
12
6-si
n 13
4-de
xt
34.
Max
imum
hei
ght o
f fo
ram
en m
agnu
m
155.
0
35
. M
axim
um w
idth
of
fora
men
mag
num
75
.0
36.
Len
gth
of a
rcus
zyg
omat
icus
44
0.0
37.
Wid
th o
f ar
cus
zygo
mat
icus
95
-100
38
. M
axim
um d
epth
of
pala
tum
dur
um
176.
0
39
. W
idth
of
pala
tum
dur
um a
t P3
550.
0
40
. L
engt
h of
toot
h ro
w P
3 -M3
510.
0 47
0.0
390.
0 54
0.0
252.
0
14
Ear area. (Fig. 5 A, C or. r and Pl. I, Fig. 2). It is high,wide and deep. Begins immediately behind the eyesocket. In the posteriror part it borders os occipitale.Maxillary bone (Maxilla) (Fig. 5 B m and Pl. I, Fig. 2).Wide, slightly concave at the palatum. Length400.0 mm, height at M1 – 250 mm. Facial crest andinfraorbital canal not pronounced.Palatum durum. (Fig. 5 B; Pl. I, Fig. 2). Long andvery narrow. Strongly concave in the middle (mea-surements in Table 1).Upper cheek teeth (Pl. I, Fig. 2 and Pl. IV, Fig. 1).Both toothrows are entirely preserved. Only the firstmolars are slightly worn out, so that part of the den-tin is seen. There is no difference in the structure ofthe left and the right ones. Their sizes differ with 1 to3 mm (see Table 2) but this is assumed to be normal.We shall discuss only the morphological differenc-es, where there are such.
The third premolar (P3 dext) is large, of almostregular trapezoid shape. It has one large outer ridgeand two inner, situated diagonally to it. Their innerends touch each other, and the outer ends – theanterior and the posterior part of the outer ridge cor-respondingly. Thus the three ridges surround a deeptriangular valley. The anterior inner ridge is wider,with a triangular shape, slightly worn out at the tip.Its inner side is concave and, together with the cin-gulum on this side of the crown, surrounds a smalland shallow triangular valley. On the inner side it isslightly more worn out. The cingulum is well devel-oped on the whole anterior and the posterior outerpart of the crown.
The fourth premolar (P4 dext) has an almost tet-ragonal shape. It consists of one outer and two innerridges, situated perpendicularly to the outer. Theanterior inner ridge is wider than the posterior andless worn out. It has the shape of a isosceles triangle,the base of which touches the inner anterior part ofthe outer ridge. At this place the two ridges surrounda shallow valley. The posterior ridge is narrower andless worn out. By touching the outer ridge it is divid-ed in two parts, and V-shaped. Thus the ridges sur-round a deep triangular valley with steep walls. Onthe inner and outer posterior side and on the anteri-or side of the crown there is a slightly developed cin-gulum. The left P4 has a weaker cingulum on its an-terior side.
The first molar (M1 dext) is long and narrow. Itconsists of three ridges. First two are more worn out.The three ridges on the inner side are wider. Theygradually become thinner toward the outer wall ofthe crown. At the very end they are slightly curveddownwards but they don’t touch each other. Theyare separated by small valleys narrowing in their mid-dles. The second and third ridges are slightly con-cave on the posterior side, while the anterior side ofthe third ridge is more convex. A tubercle at the baseof each valley, but only from the inner side, closesthem. The valleys are open outwards. In front, thecrown has a cingulum which, together with the an-terior wall of the first ridge, surrounds a long andshallow valley.
The second molar (M2 dext) is large and almostsquare. It is built by two thick and high ridges whichare slightly curved backwards on the outer side. Thuson the posterior wall of the ridge there is a smallconcavity, while in front they are straight. A deepvalley with steep walls separates them. Both teeth (leftand right) have a clear cingulum on the anteriorand the posterior sides. In the anterior part of thecrown the cingulum is something like an anteriortalon and has a worn-out tip.
The last molar (M3 dext) is the largest. It has twothick ridges, slightly convex forwardly. A deep andwide valley separates them. The ridges are wider onthe inner side and slightly curved backwards on theouter side. They form a small concavity on the poste-rior side. The valley between the two ridges is free.The cingulum is well developed both at the anteriorand the posterior part, shaping an anterior and pos-terior talon.
Comparison
According to the literature, there are only two com-pletely preserved skulls found up till now. The firsthas been found near Eppelsheim in 1835, describedby Andrews (1921) as D. giganteum Kaup and isstored at the British Museum. The second has beenfound near the village Gussiatin, Tirasspol region,by a palaeontological expedition from the Instituteof Zoology, Academy of Sciences of the UkrainianSSR in 1963, described by Svistun in 1974 and re-ferred by him to D. levius Jourdan. In Vienna thereis a mounted skeleton of D. bavaricum H. v. Meyerbut its skull is entirely reconstructed (see Pl. XXII).Similar is the case with D. gigantissimum Stefanescuin Bucharest. From the skull of the deinothere skele-ton recently mounted in Kishinev only part of themaxilla with the upper teeth is preserved.
In conclusion, it seems that the only possible com-parisons are with the skulls from Eppelsheim andGussiatin. The main differences between them andthe skull described by us (cf. Figs. 5, 6 and 7) are asfollows:1. Relatively to the body size, the skull described hereis short, but high. Compared to the other two, theEzerovo skull is highest, and the one from Gussiatin– the lowest.2. The premaxillary bones have interesting differences(Figs. 5, 6 and 7 pmx). In the skull described by usthey are curved downwards as in the Eppelsheim skull,but are not so strongly flattened, the part not knittogether is much smaller and the incisure betweenthem is narrower and shallower. In the Gussetin skullthe curving is weaker. In this species, in their pre-orbital part the lateral processes are much stronger thanin both our and the Eppelsheim skull. Besides, in theGussiatin skull the premaxillary bones are coarsely rug-ged with a tuberosity protruding forward. On it, ac-cording to Svistun 1974, there could be horn-like struc-tures substituting for the upper tusks. There is no suchthing in our material, neither in the Eppelsheim skull.
15
D. g
igan
teum
Kau
p D
. thr
acee
nsis
sp
.n. E
zero
vo, B
ulga
ria
D. g
igan
tiss
imum
St
efan
escu
, 190
9, R
oman
ia
race
mjo
r B
akal
ov, N
ikol
ov,
1962
, Bul
gari
a ra
cem
inor
, Bak
alov
, N
ikol
ov, 1
962,
Bul
gari
a
Mea
sure
men
ts in
mm
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
Len
gth
of th
e to
oth
100.
0 10
0.0
- -
102.
0 -
- -
- 36
.0
W
idth
of
the
toot
h 10
5.0
106.
5 -
- 10
0.0
- -
- -
30.0
P3
Wid
th o
f fi
rst r
idge
90
.5
91.8
-
- 97
.0
- -
- -
17.0
Wid
th o
f se
cond
rid
ge
106.
5 10
6.0
- -
106.
0 -
- -
- 36
.0
E
nam
el th
ickn
ess
4.0
4.2
- -
- -
- -
- -
L
engt
h of
the
toot
h 75
.0
81.5
87
.0
- 80
.0
- -
- -
51.0
Wid
th o
f th
e to
oth
100.
8 98
.5
- -
- -
- -
- 50
.0
P4 W
idth
of
firs
t rid
ge
96.3
87
.0
96.0
-
68.0
-
- -
- 42
.5
W
idth
of
seco
nd r
idge
10
0.5
102.
5 -
- 62
.0
- -
- -
50.0
Ena
mel
thic
knes
s 4.
2 4.
3 -
- -
- -
- -
-
Len
gth
of th
e to
oth
110.
0 11
0.0
119.
0 11
5.0
99.0
10
3.0
92.6
-
- 74
.0
W
idth
of
the
firs
t rid
ge a
t the
bas
e 94
.6
100.
0 10
0.0
93.0
82
.0
73.0
75
.0
- -
52.0
Wid
th o
f th
e fi
rst r
idge
at t
he to
p 70
.0
73.0
-
- -
- -
- -
29.5
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 94
.0
97.0
98
.0
92.0
65
.0
70.0
61
.0
- -
57.0
M
1 W
idth
of
the
seco
nd r
idge
at t
he to
p 69
.0
71.0
-
- -
- -
- -
33.0
Wid
th o
f th
e th
ird
ridg
e at
the
base
83
.5
84.5
86
.0
83.0
-
- 53
.0
- -
48.5
Wid
th o
f th
e th
ird
ridg
e at
the
top
63.5
62
.0
- -
- -
- -
- 31
.0
W
idth
of
post
erio
r ta
lon
- -
- -
- -
- -
- -
E
nam
el th
ickn
ess
4.3
4.3
- -
- -
- -
- -
L
engt
h of
the
toot
h 10
7.7
106.
5 11
6.0
114.
0 10
3.0
92.0
86
.0
91.0
70
.0
71.0
Wid
th o
f th
e fi
rst r
idge
at t
he b
ase
113.
6 11
5.0
129.
0 11
1.0
103.
0 98
.0
96.0
10
1.0
62.0
70
.5
W
idth
of
the
firs
t rid
ge a
t the
top
88.8
87
.5
- -
- -
- -
44.0
52
.0
M2
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 11
0.0
110.
0 11
7.0
114.
0 10
2.0
90.0
84
.5
83.0
59
.0
73.0
Wid
th o
f th
e se
cond
rid
ge a
t the
top
85.3
84
.5
- -
- -
- -
38.0
51
.5
W
idth
of
post
erio
r ta
lon
- -
- -
- -
- -
- -
E
nam
el th
ickn
ess
4.7
4.6
- -
- -
- -
- -
L
engt
h of
the
toot
h 11
0.0
108.
0 11
5.0
107.
0 96
.6
105.
0 89
.0
96.0
-
-
Wid
th o
f th
e fi
rst r
idge
at t
he b
ase
118.
0 11
5.0
114.
0 11
4.0
103.
0 10
7.0
95.0
10
5.6
- -
W
idth
of
the
firs
t rid
ge a
t the
top
100.
0 98
.0
- -
- -
- -
- -
M3
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 10
7.2
107.
0 10
7.0
104.
0 98
.0
101.
0 89
.0
89.0
-
-
Wid
th o
f th
e se
cond
rid
ge a
t the
top
87.0
89
.0
- -
- -
- -
- -
W
idth
of
post
erio
r ta
lon
78.0
78
.0
- -
- -
- -
- -
E
nam
el th
ickn
ess
5.1
5.2
- -
- -
- -
- -
Table
2U
pper
cheek-teeth
18
Fig. 6. Deinotherium levius Jourdan
Fig. 7. Deinotherium giganteum Kaup
3. The external nares (Figs. 5, 6, and 7 – exn) in D.thraceiensis sp. n. is large, deep and pear-shaped. Itstarts from the very beginning of the nasal bones,gradually widening and taking two thirds of thelength of the whole skull. In the other species it isshorter and almost of the same width in its anteriorand posterior parts.4. The nasal bones in D. thraceiensis sp. n. are short,narrow, frontally rounded and fused. In the otherspecies they are longer, wider, almost flat for most oftheir length, and separated. In the Gussiatin speciesthey protrude above the external nares. This is notthe case with the two other skulls (Figs. 5, 6, and 7).5. The calvaria of the skull (fronto-parietal part, Figs.5, 6, and 7 – Ic in D. thraceiensis sp. n.) is high andwide). In the other two species it is lower and shorter.
In the Gussiatin species it is even strongly concavein its anterior part.6. The eye socket (Figs. 5, 6, and 7 – orb) in theEzerovo skull is large and separated from the eararea. In D. giganteum Kaup they are almost fused,and in D. levius Jourdan from Gussiatin the situa-tion is closer to our case.7. The zygomatic arch (Figs. 5, 6, and 7 – zyg) of D.thraceiensis sp. n. with its developed proc. postorbit-alis resembles the one of D. levius Jourdan, but in itsposterior part, behind its connection with the tem-poral bone, it lacks the processus that Svistun notes.He regards this processus as related to the develop-ment of the lower tusks. No comparison with the zy-gomatic arch of the Eppelsheim skull is possible,because it is not preserved.
19
8. There are significant differences between the threeskulls in the occipital region. Os occipitale in D. thra-ceiensis sp. n. is high, wide and at an angle of 80°toward the forehead, and of ca. 70° to the plane ofthe teeth. In D. giganteum Kaup these angles are 70°and 50° correspondingly; in D. levius Jourdan thedeclination of the occipital bone is even larger, sothe angle is only 60°. As a whole, the skull of D. leviusJourdan is very flat and low. Os occipitale is visiblyconcave and very wide, forming two lateral wings(Fig. 6 C and D – OS). Those wings are almost lack-ing in D. thraceiensis sp. n. and in D. giganteumKaup they are much less developed.9. The position of the occipital condyles is ratherdifferent (Figs. 5, 6, and 7 – oc). In D. thraceiensissp. n. they are situated in the middle and in the low-er part of the occipital bone, almost the same is theirposition in D. giganteum Kaup, and in D. levius Jour-dan they are on the upper half of the noted bone.According to Svistun, this permitted the animal toraise its head almost at right angle to its neck, thusfully using its back curved lower tusks.10. Worth mentioning is a characteristic peculiarityof the Gussiatin skull lacking in all the others,including the skull from Ezerovo. This skull has adouble articulation with the mandible. Once by anarticular surface on the zygomatic process of thetemporal bone and then by a second surface on thepetromastoideum. Between those two surfaces is theexternal meatus of the auditory canal. How andwhen this second articular surface was used, isunclear.
After comparing their anatomy we can see thatdeinotheres could be divided in two groups. One in-cludes D. giganteum Kaup, D. gigantissimum Stefa-nescu, D. thraceiensis sp. n., D. levius Jourdan andD. indicus Falconer, which have larger teeth, and thesecond – D. bavaricum H. v. Meyer, D. pentapota-miae Lydekker and others, all with smaller teeth.
As a whole, the teeth are rather similar and differ-ences are seen mainly in the two premolars P3 and P4.
In D. giganteum Kaup P3 is more rounded on theinner side. The two inner ridges are more perpendic-ular to the outer one. The anterior inner ridge is notconnected with the outer and is at an acuter angle to itthan in other taxa. They almost don’t touch each otheron the inner side. If anything like that is observed inthese species, than it is down at the very basis. Theridges merge when they are worn. The cingulum ismore clearly seen here. It consists of numerous largeand small tubercles on all sides of the crown.
No comparison with the third premolars of D. gi-gantissimum Stefanescu is possible, because they havebeen restored after D. giganteum Kaup.
In D. levius Jourdan the anterior inner ridge of P3
is situated diagonally to the outer one and touchesit. The posterior one is wider and V-shaped in itsupper part, with a shorter anterior part. It is also di-agonal to the outer ridge, but does not touch it.
In D. indicum Falconer the inner ridges are somemore diagonally situated, compared to D. giganteumKaup, but their tips do not contact.
D. bavaricum H. v. Meyer has small P3s. Theirinner ridges are some more perpendicular to the outerone. This is rather clear with the anterior inner ridge.The valley between them is wider and deeper.
In D. pentapotamiae Lydekker, D. levius Jourdanand D. indicum Falconer the differences are almostthe same, concerning the position of the two innerridges and the size of the tooth and its cingulum.
The fourth premolar of D. thraceiensis sp. n. hasinner ridges perpendicular to the outer one. Theposterior is narrower and longer. The triangular val-ley surrounded by the posterior inner wall of the out-er ridge and the inner wall of the posterior ridge isclearer than in all the other species.
D. gigantissimum Stefanescu has a longer P4. Itstransverse ridges are at a larger distance from eachother. Besides, it has a strong cingulum on the ante-rior and the posterior side, lacking in our species.
In D. bavaricum H. v. Meyer P4s are smaller andof a more regular square shape. The transverse ridg-es are more distant at the inner side of the crown,and the valley surrounded by them and the outerridge is wider and larger.
In D. levius Jourdan the posterior inner ridge isalso attached to the outer one with its longer posteri-or branch.
In D. indicum Falconer and D. pentapotamiaeLydekker the last premolars have a square shape andare smaller. Their transverse ridges are parallel andof equal size. The valley between them is larger andeverywhere equally wide.
In the M1s of D. giganteum Kaup, unlike D. thra-ceiensis sp. n., the three ridges are almost of equalsize, with an equally developed cingulum on bothsides of the valleys, blocking them entirely on theouter side. The latter is of significant taxonomicalvalue.
In D. gigantissimum Stefanescu has a developedcingulum on the outer anterior and on the posteriorside of the crown, resembling a chain of large tuber-cles, closely arranged along the end of the tooth.
There are no significant differences in the struc-ture of the first molars of D. bavaricum H. v. Meyer,except in their size.
The other molars (M2 and M3) of D. thraceiensisare not very different from those of the other species.
Mandible. (Pl. III, Fig. 1 and Pl. V).Like the skull, it is entirely preserved. Compared tothe size of the body, it is not large (measurements inTable 3). Ramus horizontalis long, high, wide andlaterally slightly flattened. Below P3 there is a wellshaped double opening – foramen mentale. Thehighest point of the mandible is at the symphysis, itbecomes lower towards the posterior end (see PlateV). Because of this, the last molar is not horizontal,but inclined backwards. Processus angularis is strong-ly developed, and fossa masseterica is wide but notvery deep. The most typical feature of the animalsbelonging to this group is the shape of the symphys-is. Together with the tusks, it is curved down andback. In D. thraceiensis sp. n. the symphysis is solid,
16
Table
2 A
Upper
cheek-teeth
D. g
igan
teum
Kau
p D
. gig
ante
um
Dep
eret
, 188
7 D
. thr
acei
ensi
s sp
.n. E
zero
vo,
Bul
gari
a
afte
r Петрон
jеви
h,
1954
, Y
ugos
lavi
a
afte
r
Ath
anas
ia, 1
907,
R
oman
ia
afte
r
А. Б
елокрые,
19
60, K
rivo
i R
og, M
oldo
va
Stro
mer
, M
ünhe
n
from
Se
int
Yea
n
from
E
ppel
sh
eim
D. b
avar
icum
H. v
. Mey
er,
Ger
man
y
Mea
sure
men
ts in
mm
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
Len
gth
of th
e to
oth
100.
0 10
0.0
- 73
.5
87.0
-
98.0
96
.0
72.0
71
.0
85.0
83
.0
66.0
66
.0
52.0
53
.0
Wid
th o
f th
e to
oth
105.
0 10
6.5
- 69
.0
100.
0 -
- -
- -
- -
61.0
61
.0
33.0
-
Wid
th o
f se
cond
rid
ge
90.5
91
.8
- 68
.2
98.0
-
97.0
95
.0
69.0
71
.0
- -
- -
47.5
-
Wid
th o
f se
cond
rid
ge
106.
5 10
6.0
- 70
.4
70.0
-
- -
- -
- -
- -
- -
P3
Ena
mel
thic
knes
s 4.
0 4.
2 -
- 5.
0 -
- -
- -
- -
- -
- -
Len
gth
of th
e to
oth
75.0
81
.5
- 67
.5
62.0
-
88.0
86
.0
68.0
63
.0
77.0
78
.0
61.0
61
.0
50.0
-
Wid
th o
f th
e to
oth
100.
8 98
.5
- 79
.0
50.0
-
- -
- -
- -
66.0
66
.0
51.0
-
Wid
th o
f fi
rst r
idge
96
.3
87.0
-
- 55
.0
- 99
.0
98.0
74
.0
73.0
-
- -
- -
- W
idth
of
seco
nd r
idge
10
0.5
102.
5 -
75.0
37
.0
- -
- -
- -
- -
- -
- P4
Ena
mel
thic
knes
s 4.
2 4.
3 -
- -
- -
- -
- -
- -
- -
- L
engt
h of
the
toot
h 11
0.0
110.
0 61
.2
- 11
9.0
- 10
4.0
- 81
.0
82.0
-
95.0
75
.0
75.0
72
.0
- W
idth
of
the
firs
t rid
ge a
t the
bas
e 94
.6
100.
0 50
.5
- 10
0.0
- 93
.0
- 68
.0
68.0
-
- 62
.0
62.0
54
.0
- W
idth
of
the
firs
t rid
ge a
t the
top
70
.0
73.0
-
- 48
.0
- -
- -
- -
- -
- -
- W
idth
of
the
seco
nd r
idge
at t
he b
ase
94.0
97
.0
70.8
-
98.0
-
- -
- -
- -
- -
54.0
-
Wid
th o
f th
e se
cond
rid
ge a
t the
top
69.0
71
.0
- -
45.0
-
- -
- -
- -
- -
- -
Wid
th o
f th
e th
ird
ridg
e at
the
base
83
.5
84.5
-
- 86
.0
- -
- -
- -
- -
- 44
.0
- W
idth
of
the
thir
d ri
dge
at th
e to
p 63
.5
62.0
-
- -
- -
- -
- -
- -
- -
- W
idth
of
post
erio
r ta
lon
- -
- -
- -
- -
- -
- -
- -
- -
M1
Ena
mel
thic
knes
s 4.
3 4.
3 -
- -
- -
- -
- -
- -
- -
- L
engt
h of
the
toot
h 10
7.7
106.
5 79
.5
- -
115.
0 99
.0
- 80
.0
78.0
-
91.0
68
.0
68.0
60
.0
64.0
W
idth
of
the
firs
t rid
ge a
t the
bas
e 11
3.6
115.
0 80
.5
- -
88.0
-
- 82
.0
81.0
-
- 66
.0
66.0
-
61.5
W
idth
of
the
firs
t rid
ge a
t the
top
88.8
87
.5
- -
- -
- -
- -
- -
- -
- -
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 11
0.0
110.
0 79
.0
- -
89.0
10
8.0
- -
- -
- -
- -
61.0
W
idth
of
the
seco
nd r
idge
at t
he to
p 85
.3
84.5
-
- -
- -
- -
- -
- -
- -
- W
idth
of
post
erio
r ta
lon
- -
- -
- 75
.0
- -
- -
- -
- -
- -
M2
Ena
mel
thic
knes
s 4.
7 4.
6 -
- -
- -
- -
- -
- -
- -
- L
engt
h of
the
toot
h 11
0.0
108.
0 -
70.0
-
- -
- 81
.0
82.0
-
91.0
67
.0
63.0
62
.0
63.0
W
idth
of
the
firs
t rid
ge a
t the
bas
e 11
8.0
115.
0 -
73.0
-
- -
- 89
.0
89.0
-
- 70
.0
70.0
-
63.0
W
idth
of
the
firs
t rid
ge a
t the
top
100.
0 98
.0
- -
- -
- -
- -
- -
- -
- -
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 10
7.2
107.
0 -
69.0
-
- -
- -
- -
- -
- -
56.0
W
idth
of
the
seco
nd r
idge
at t
he to
p 87
.0
89.0
-
- -
- -
- -
- -
- -
- -
- W
idth
of
post
erio
r ta
lon
78.0
78
.0
- -
- -
- -
- -
- -
- -
- -
M3
Ena
mel
thic
knes
s 5.
1 5.
2 -
- -
- -
- -
- -
- -
- -
-
173 Geologica Balcanica, 3-4/2006
Note
: In
the
colu
mns
aft
er L
ydek
ker
, 1880,
the
size
is
in i
nches
.
Table
2 B
Upper
cheek-teeth
Rho
ne –
Dep
eret
, 188
7 af
ter
Lyd
ekke
r, 1
880
D. t
hrac
eien
sis
sp.n
. E
zero
vo, B
ulga
ria
D. p
enta
pota
mia
e af
ter
Deh
m 1
963,
Pa
kist
an
M
easu
rem
ents
in m
m
sin
dext
D. l
eviu
s Jo
rdan
D
. gig
ante
um
Kau
p D
. cuv
ieri
K
aup
D.
pent
apot
amia
e D
. ind
icum
Fal
. N
arba
da, I
ndia
si
n de
xt
sin
L
engt
h of
the
toot
h 10
0.0
100.
0 66
.0
70.0
53
.0
- -
50.0
51
.0
58.4
Wid
th o
f th
e to
oth
105.
0 10
6.5
- -
- -
- 45
.0
51.0
58
.2
P3 W
idth
of
seco
nd r
idge
90
.5
91.8
-
- -
- -
51.0
52
.0
60.0
Wid
th o
f se
cond
rid
ge
106.
5 10
6.0
- -
- -
- -
- -
E
nam
el th
ickn
ess
4.0
4.2
- -
- -
- -
- -
L
engt
h of
the
toot
h 75
.0
81.5
78.0
-
- -
- -
52.4
Wid
th o
f th
e to
oth
100.
8 98
.5
-
- -
- -
- 64
.4
P4 W
idth
of
firs
t rid
ge
96.3
87
.0
-
- -
- -
- 62
.2
W
idth
of
seco
nd r
idge
10
0.5
102.
5
- -
- -
- -
-
Ena
mel
thic
knes
s 4.
2 4.
3
- -
- -
- -
-
Len
gth
of th
e to
oth
110.
0 11
0.0
95
.0
72.0
2.
3 3.
9 72
.0
70.0
68
.4
W
idth
of
the
firs
t rid
ge a
t the
bas
e 94
.6
100.
0
- -
2.1
2.5
62.0
52
.2
62.2
Wid
th o
f th
e fi
rst r
idge
at t
he to
p
70.0
73
.0
-
- -
- -
- -
W
idth
of
the
seco
nd r
idge
at t
he b
ase
94.0
97
.0
-
- -
- 60
.3
- 62
.2
M1
Wid
th o
f th
e se
cond
rid
ge a
t the
top
69.0
71
.0
-
- -
- -
- -
W
idth
of
the
thir
d ri
dge
at th
e ba
se
83.5
84
.5
-
- -
- -
- -
W
idth
of
the
thir
d ri
dge
at th
e to
p 63
.5
62.0
- -
- -
- -
-
Wid
th o
f po
ster
ior
talo
n -
-
- -
- -
- -
-
Ena
mel
thic
knes
s 4.
3 4.
3
- -
- -
- -
-
Len
gth
of th
e to
oth
107.
7 10
6.5
91
.0
60.0
2.
1 2.
5 -
- 71
.3
W
idth
of
the
firs
t rid
ge a
t the
bas
e 11
3.6
115.
0
- -
2.5
2.4
- -
73.2
Wid
th o
f th
e fi
rst r
idge
at t
he to
p 88
.8
87.5
- -
- -
- -
- M
2 W
idth
of
the
seco
nd r
idge
at t
he b
ase
110.
0 11
0.0
-
- -
- -
- 71
.8
W
idth
of
the
seco
nd r
idge
at t
he to
p 85
.3
84.5
- -
- -
- -
-
Wid
th o
f po
ster
ior
talo
n -
-
- -
- -
- -
-
Ena
mel
thic
knes
s 4.
7 4.
6
- -
- -
- -
-
Len
gth
of th
e to
oth
110.
0 10
8.0
91
.0
62.0
2.
75
- 67
.0
66.0
68
.4
W
idth
of
the
firs
t rid
ge a
t the
bas
e 11
8.0
115.
0
- -
2.4
- -
70.0
74
.0
W
idth
of
the
firs
t rid
ge a
t the
top
100.
0 98
.0
-
- -
- -
- -
M3
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 10
7.2
107.
0
- -
- -
69.0
59
.6
64.3
Wid
th o
f th
e se
cond
rid
ge a
t the
top
87.0
89
.0
-
- -
- -
- -
W
idth
of
post
erio
r ta
lon
78.0
78
.0
-
- -
- -
- -
E
nam
el th
ickn
ess
5.1
5.2
-
- -
- -
- -
18
Fig. 6. Deinotherium levius Jourdan
Fig. 7. Deinotherium giganteum Kaup
3. The external nares (Figs. 5, 6, and 7 – exn) in D.thraceiensis sp. n. is large, deep and pear-shaped. Itstarts from the very beginning of the nasal bones,gradually widening and taking two thirds of thelength of the whole skull. In the other species it isshorter and almost of the same width in its anteriorand posterior parts.4. The nasal bones in D. thraceiensis sp. n. are short,narrow, frontally rounded and fused. In the otherspecies they are longer, wider, almost flat for most oftheir length, and separated. In the Gussiatin speciesthey protrude above the external nares. This is notthe case with the two other skulls (Figs. 5, 6, and 7).5. The calvaria of the skull (fronto-parietal part, Figs.5, 6, and 7 – Ic in D. thraceiensis sp. n.) is high andwide). In the other two species it is lower and shorter.
In the Gussiatin species it is even strongly concavein its anterior part.6. The eye socket (Figs. 5, 6, and 7 – orb) in theEzerovo skull is large and separated from the eararea. In D. giganteum Kaup they are almost fused,and in D. levius Jourdan from Gussiatin the situa-tion is closer to our case.7. The zygomatic arch (Figs. 5, 6, and 7 – zyg) of D.thraceiensis sp. n. with its developed proc. postorbit-alis resembles the one of D. levius Jourdan, but in itsposterior part, behind its connection with the tem-poral bone, it lacks the processus that Svistun notes.He regards this processus as related to the develop-ment of the lower tusks. No comparison with the zy-gomatic arch of the Eppelsheim skull is possible,because it is not preserved.
19
8. There are significant differences between the threeskulls in the occipital region. Os occipitale in D. thra-ceiensis sp. n. is high, wide and at an angle of 80°toward the forehead, and of ca. 70° to the plane ofthe teeth. In D. giganteum Kaup these angles are 70°and 50° correspondingly; in D. levius Jourdan thedeclination of the occipital bone is even larger, sothe angle is only 60°. As a whole, the skull of D. leviusJourdan is very flat and low. Os occipitale is visiblyconcave and very wide, forming two lateral wings(Fig. 6 C and D – OS). Those wings are almost lack-ing in D. thraceiensis sp. n. and in D. giganteumKaup they are much less developed.9. The position of the occipital condyles is ratherdifferent (Figs. 5, 6, and 7 – oc). In D. thraceiensissp. n. they are situated in the middle and in the low-er part of the occipital bone, almost the same is theirposition in D. giganteum Kaup, and in D. levius Jour-dan they are on the upper half of the noted bone.According to Svistun, this permitted the animal toraise its head almost at right angle to its neck, thusfully using its back curved lower tusks.10. Worth mentioning is a characteristic peculiarityof the Gussiatin skull lacking in all the others,including the skull from Ezerovo. This skull has adouble articulation with the mandible. Once by anarticular surface on the zygomatic process of thetemporal bone and then by a second surface on thepetromastoideum. Between those two surfaces is theexternal meatus of the auditory canal. How andwhen this second articular surface was used, isunclear.
After comparing their anatomy we can see thatdeinotheres could be divided in two groups. One in-cludes D. giganteum Kaup, D. gigantissimum Stefa-nescu, D. thraceiensis sp. n., D. levius Jourdan andD. indicus Falconer, which have larger teeth, and thesecond – D. bavaricum H. v. Meyer, D. pentapota-miae Lydekker and others, all with smaller teeth.
As a whole, the teeth are rather similar and differ-ences are seen mainly in the two premolars P3 and P4.
In D. giganteum Kaup P3 is more rounded on theinner side. The two inner ridges are more perpendic-ular to the outer one. The anterior inner ridge is notconnected with the outer and is at an acuter angle to itthan in other taxa. They almost don’t touch each otheron the inner side. If anything like that is observed inthese species, than it is down at the very basis. Theridges merge when they are worn. The cingulum ismore clearly seen here. It consists of numerous largeand small tubercles on all sides of the crown.
No comparison with the third premolars of D. gi-gantissimum Stefanescu is possible, because they havebeen restored after D. giganteum Kaup.
In D. levius Jourdan the anterior inner ridge of P3
is situated diagonally to the outer one and touchesit. The posterior one is wider and V-shaped in itsupper part, with a shorter anterior part. It is also di-agonal to the outer ridge, but does not touch it.
In D. indicum Falconer the inner ridges are somemore diagonally situated, compared to D. giganteumKaup, but their tips do not contact.
D. bavaricum H. v. Meyer has small P3s. Theirinner ridges are some more perpendicular to the outerone. This is rather clear with the anterior inner ridge.The valley between them is wider and deeper.
In D. pentapotamiae Lydekker, D. levius Jourdanand D. indicum Falconer the differences are almostthe same, concerning the position of the two innerridges and the size of the tooth and its cingulum.
The fourth premolar of D. thraceiensis sp. n. hasinner ridges perpendicular to the outer one. Theposterior is narrower and longer. The triangular val-ley surrounded by the posterior inner wall of the out-er ridge and the inner wall of the posterior ridge isclearer than in all the other species.
D. gigantissimum Stefanescu has a longer P4. Itstransverse ridges are at a larger distance from eachother. Besides, it has a strong cingulum on the ante-rior and the posterior side, lacking in our species.
In D. bavaricum H. v. Meyer P4s are smaller andof a more regular square shape. The transverse ridg-es are more distant at the inner side of the crown,and the valley surrounded by them and the outerridge is wider and larger.
In D. levius Jourdan the posterior inner ridge isalso attached to the outer one with its longer posteri-or branch.
In D. indicum Falconer and D. pentapotamiaeLydekker the last premolars have a square shape andare smaller. Their transverse ridges are parallel andof equal size. The valley between them is larger andeverywhere equally wide.
In the M1s of D. giganteum Kaup, unlike D. thra-ceiensis sp. n., the three ridges are almost of equalsize, with an equally developed cingulum on bothsides of the valleys, blocking them entirely on theouter side. The latter is of significant taxonomicalvalue.
In D. gigantissimum Stefanescu has a developedcingulum on the outer anterior and on the posteriorside of the crown, resembling a chain of large tuber-cles, closely arranged along the end of the tooth.
There are no significant differences in the struc-ture of the first molars of D. bavaricum H. v. Meyer,except in their size.
The other molars (M2 and M3) of D. thraceiensisare not very different from those of the other species.
Mandible. (Pl. III, Fig. 1 and Pl. V).Like the skull, it is entirely preserved. Compared tothe size of the body, it is not large (measurements inTable 3). Ramus horizontalis long, high, wide andlaterally slightly flattened. Below P3 there is a wellshaped double opening – foramen mentale. Thehighest point of the mandible is at the symphysis, itbecomes lower towards the posterior end (see PlateV). Because of this, the last molar is not horizontal,but inclined backwards. Processus angularis is strong-ly developed, and fossa masseterica is wide but notvery deep. The most typical feature of the animalsbelonging to this group is the shape of the symphys-is. Together with the tusks, it is curved down andback. In D. thraceiensis sp. n. the symphysis is solid,
20
Table
3M
andib
les – m
andib
ula
e
D. g
igan
teum
Kau
p D
. bav
aric
um H
. v. M
ayer
№
D
imen
sion
s, m
m
D. t
hrac
eien
sis
sp.n
. E
zero
vo
D. g
igan
tiss
imum
R
oman
ia
Mol
dova
M
ünch
en
H. v
. Mey
er
Mün
chen
1.
T
otal
leng
th –
fro
m th
e be
ginn
ing
to th
e en
d of
ram
us a
scen
dens
11
20.0
12
00.0
10
00.0
11
20.0
82
0.0
840.
0 2.
L
engt
h of
sym
phys
is –
hor
izon
tal
360.
0 42
0.0
-
320.
0 23
0.0
235.
0 3.
L
engt
h of
sym
phys
is –
ver
tica
l 43
5.0
- -
- -
- 4.
D
iam
eter
of
the
sym
phys
is a
t the
ang
le w
ith
the
man
dibl
e 31
0.0
- -
5.
D
iam
eter
of
the
sym
phys
is a
t the
beg
inni
ng o
f I
280.
0 -
-
6.
Wid
th o
f th
e th
roug
h of
the
sym
phys
is
145.
0 -
-
7.
Len
gth
of s
ymph
ysis
and
ram
us a
scen
dens
11
00.0
11
50.0
-
8.
H
eigh
t of
the
man
dibl
e at
P3
235.
0 -
-
9.
Hei
ght o
f th
e m
andi
ble
at M
1 23
0.0
190.
0 23
0.0
210.
0 16
0.0
154.
0 10
. H
eigh
t of
the
man
dibl
e at
M3
185.
0 -
192.
0
11.
Hei
ght o
f ra
mus
asc
ende
ns a
t pro
c. a
rtic
ular
is
685.
0 68
0.0
455.
0
12.
Hei
ght o
f ra
mus
asc
ende
ns a
t pro
c. c
oron
oide
us
505.
0 -
-
13.
Wid
th –
mid
dle
of r
amus
asc
ende
ns
470.
0 48
2.0
-
14.
Dis
tanc
e be
twee
n th
e tw
o se
mim
andi
bles
at P
3 10
5.0
- -
15
. D
ista
nce
betw
een
the
two
sem
iman
dibl
es a
t M1
150.
0 -
-
16.
Dis
tanc
e be
twee
n th
e tw
o se
mim
andi
bles
at M
3 12
0.0
- -
17
. L
engt
h of
the
inci
sors
– I
855.
0 60
8.0
900.
0 79
5.0
530.
0 52
5.0
18.
Dia
met
er o
f I –
at t
he b
asis
13
2.0
172.
0 14
2.0
19
. D
iam
eter
of
I – in
the
mid
dle
125.
0 12
0.0
125.
0
20.
Dis
tanc
e be
twee
n th
e po
ints
of
I 47
0.0
- -
21
. D
ista
nce
betw
een
the
poin
t of
I and
ram
us a
scen
dens
89
5.0
- -
22
. L
engt
h of
the
enti
re to
oth
row
P3-
M1
510.
0 54
0.0
470.
0 48
5.0
370.
0 36
5.0
21
Table
4Lower cheek te
eth
D. g
igan
teum
Kau
p D
. thr
acei
ensi
s
sp.n
. Eze
rovo
, Bul
gari
a D
. gig
anti
ssim
um
Stef
anes
cu, 1
909,
Rom
ania
B
akal
ov, N
ikol
ov, 1
962,
B
ulga
ria
race
min
or, B
akal
ov,
Nik
olov
, 196
2, B
ulga
ria
M
easu
rem
ents
in m
m
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
Len
gth
of th
e to
oth
80.5
75
.5
- -
- 80
.0
- -
- -
W
idth
of
the
toot
h 68
.0
67.0
-
- -
- -
- -
- P3
Wid
th o
f fi
rst r
idge
58
.0
55.0
-
- -
68.0
-
- -
-
Wid
th o
f se
cond
rid
ge
70.0
73
.0
- -
- 62
.0
- -
- -
E
nam
el th
ickn
ess
4.0
3.8
- -
- -
- -
- -
L
engt
h of
the
toot
h 89
.0
89.0
90
.0
- -
92.0
-
- -
-
Wid
th o
f th
e to
oth
80.0
81
.0
- -
- -
- -
- -
P4 W
idth
of
firs
t rid
ge
76.0
78
.0
72.0
-
- 88
.0
- -
- -
W
idth
of
seco
nd r
idge
81
.1
83.0
73
.0
- -
86.0
-
- -
-
Ena
mel
thic
knes
s 4.
0 4.
0 -
- -
- -
- -
-
Len
gth
of th
e to
oth
107.
5 10
7.0
111.
0 10
9.0
- 11
1.0
- -
109.
0 73
.0
W
idth
of
the
firs
t rid
ge a
t the
bas
e 82
.0
81.5
72
.0
77.0
-
85.0
-
- 81
.0
73.0
Wid
th o
f th
e fi
rst r
idge
at t
he to
p -
- -
- -
- -
- -
-
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 81
.0
81.5
72
.0
- -
82.0
-
- 76
.0
37.0
M
1 W
idth
of
the
seco
nd r
idge
at t
he to
p -
- -
- -
- -
- -
-
Wid
th o
f th
e th
ird
ridg
e at
the
base
75
.0
75.5
71
.0
70.0
-
75.0
-
- -
-
Wid
th o
f th
e th
ird
ridg
e at
the
top
- -
- -
- -
- -
- -
W
idth
of
post
erio
r ta
lon
- -
- -
- -
- -
- -
E
nam
el th
ickn
ess
4.3
4.2
- -
- -
- -
- -
L
engt
h of
the
toot
h 11
5.5
112.
5 11
1.0
- 10
6.0
104.
0 -
- 71
.0
72.0
Wid
th o
f th
e fi
rst r
idge
at t
he b
ase
103.
0 11
0.5
99.0
-
84.0
10
3.0
- -
70.5
67
.0
W
idth
of
the
firs
t rid
ge a
t the
top
80.0
81
.0
- -
- -
- -
- -
M2
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 97
.5
107.
0 97
.0
- 82
.0
99.5
-
- 73
.0
76.0
Wid
th o
f th
e se
cond
rid
ge a
t the
top
83.0
75
.5
- -
- -
- -
- -
W
idth
of
post
erio
r ta
lon
- -
- -
- -
- -
- -
E
nam
el th
ickn
ess
4.6
4.5
- -
- -
- -
- -
L
engt
h of
the
toot
h 12
0.0
118.
0 11
5.0
- 10
0.5
110.
0 -
115.
5 -
-
Wid
th o
f th
e fi
rst r
idge
at t
he b
ase
106.
0 10
5.0
114.
0 -
84.0
11
0.0
- 92
.0
- -
W
idth
of
the
firs
t rid
ge a
t the
top
85.0
82
.0
- -
68.0
-
- -
- -
M3
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 95
.0
100.
0 10
7.0
- 83
.0
93.0
-
96.0
-
-
Wid
th o
f th
e se
cond
rid
ge a
t the
top
78.5
80
.0
- -
66.5
-
- -
- -
W
idth
of
post
erio
r ta
lon
64.0
67
.0
- -
65.0
-
- 56
.0
- -
E
nam
el th
ickn
ess
5.2
5.3
- -
- -
- -
- -
22
Table
4 A
Lower cheek te
eth
D. g
igan
teum
Kau
p D
. thr
acei
ensi
s sp
.n. E
zero
vo,
Bul
gari
a
Петрон
jеви
h, 1
954,
Y
ugos
lavi
a А
. Белокрые,
196
0,
Kri
voi R
og, M
oldo
va
Stro
mer
, 193
8,
Mün
chen
R
. Dei
m, 1
949,
Sou
th B
avar
ia,
Mus
eum
of
Mün
chen
Sim
ione
scu.
19
39,
Rom
ania
Mea
sure
men
ts in
mm
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
L
engt
h of
the
toot
h 80
.5
75.5
58
.2
40.0
81
.0
- 60
.0
- 62
.0
- 58
.0
- -
W
idth
of
the
toot
h 68
.0
67.0
-
- 64
.0
- -
- 55
.0
- 47
.0
- -
P3 W
idth
of
firs
t rid
ge
58.0
55
.0
38.5
28
.0
- -
- -
- -
- -
-
Wid
th o
f se
cond
rid
ge
70.0
73
.0
49.1
-
- -
55.0
-
- -
- -
-
Ena
mel
thic
knes
s 4.
0 3.
8 -
- -
- -
- -
- -
- -
L
engt
h of
the
toot
h 89
.0
89.0
64
.4
50.5
-
88.0
68
.0
- 68
.0
57.0
69
.0
67.0
-
W
idth
of
the
toot
h 80
.0
81.0
-
- -
99.0
-
- 60
.0
47.0
62
.0
61.0
-
P4 W
idth
of
firs
t rid
ge
76.0
78
.0
57.5
39
.0
- -
55.0
-
- -
- -
21.5
Wid
th o
f se
cond
rid
ge
81.1
83
.0
- 40
.0
- -
56.0
-
- -
- -
27.5
Ena
mel
thic
knes
s 4.
0 4.
0 -
- -
- -
- -
- -
- -
L
engt
h of
the
toot
h 10
7.5
107.
0 -
- -
- 75
.0
- 80
.0
- 83
.0
82.0
-
W
idth
of
the
firs
t rid
ge a
t the
bas
e 82
.0
81.5
-
- -
- 55
.0
- -
- -
- 22
.0
W
idth
of
the
firs
t rid
ge a
t the
top
- -
- -
- -
- -
- -
- -
-
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 81
.0
81.5
59
.7
- -
- -
- -
- 62
.0
- 28
.0
M1
Wid
th o
f th
e se
cond
rid
ge a
t the
top
- -
- -
- -
- -
- -
- -
-
Wid
th o
f th
e th
ird
ridg
e at
the
base
75
.0
75.5
-
-
- 52
.0
- 60
.0
- -
- -
W
idth
of
the
thir
d ri
dge
at th
e to
p -
- -
-
- -
- -
- -
- -
W
idth
of
post
erio
r ta
lon
- -
-
- -
- -
- -
- -
-
Ena
mel
thic
knes
s 4.
3 4.
2 -
-
- -
- -
- -
- -
L
engt
h of
the
toot
h 11
5.5
112.
5 76
.7
10
2.0
- 79
.0
- 82
.0
76.0
80
.0
78.0
-
W
idth
of
the
firs
t rid
ge a
t the
bas
e 10
3.0
110.
5 68
.7
-
86.0
71
.0
- 76
.0
68.0
76
.0
74.0
-
W
idth
of
the
firs
t rid
ge a
t the
top
80.0
81
.0
-
- -
- -
- -
- -
- M
2 W
idth
of
the
seco
nd r
idge
at t
he b
ase
97.5
10
7.0
66.0
- -
70.0
-
- -
76.0
76
.0
-
Wid
th o
f th
e se
cond
rid
ge a
t the
top
83.0
75
.5
-
- -
- -
- -
- -
-
Wid
th o
f po
ster
ior
talo
n -
- -
-
- -
- -
- -
- -
E
nam
el th
ickn
ess
4.6
4.5
-
- -
- -
- -
- -
-
Len
gth
of th
e to
oth
120.
0 11
8.0
-
105.
0 10
7.0
100.
0 -
93.0
81
.0
86.0
83
.0
-
Wid
th o
f th
e fi
rst r
idge
at t
he b
ase
106.
0 10
5.0
-
93.0
10
0.0
? -
78.0
68
.0
76.0
76
.0
-
Wid
th o
f th
e fi
rst r
idge
at t
he to
p 85
.0
82.0
-
-
- -
- -
- -
- -
M3
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 95
.0
100.
0 62
.0
-
- -
- -
- 76
.0
76.0
-
W
idth
of
the
seco
nd r
idge
at t
he to
p 78
.5
80.0
-
-
- -
- -
- -
- -
W
idth
of
post
erio
r ta
lon
64.0
67
.0
-
- -
- -
- -
- -
-
Ena
mel
thic
knes
s 5.
2 5.
3 -
-
- -
- -
- -
- -
23
Table
4 B
Lower cheek te
eth
Note
: In
the
colu
mns
aft
er L
ydek
ker
, 1880,
the
size
is
in i
nches
.
Rho
ne
Dep
eret
, 188
7 fr
om S
iwal
iku
Nar
bada
, af
ter
Lyd
ekke
r, 1
889,
Indi
a D
. thr
acei
ensi
s sp
.n. E
zero
vo,
Bul
gari
a
Mea
sure
men
ts in
mm
sin
dext
D. b
avar
icum
H.v
. M
eyer
, R.D
ehm
, 194
9,
Sout
h B
avar
ia, M
useu
m o
f M
ünhe
n
D.
levi
us
Jord
an
D.
giga
nteu
m
Kau
p
D. c
uvie
ri
Kau
p D
. pe
ntap
otam
iae
Lyd
ekke
r D
. in
dicu
m
Falc
oner
D. p
enta
pota
mia
e L
ydek
ker,
aft
er
R. D
ehm
, 196
3,
Paki
stan
L
engt
h of
the
toot
h 80
.5
75.5
44
.0
43.0
-
55.0
63
.0
43.0
1.
8 1.
47
- -
54.0
Wid
th o
f th
e to
oth
68.0
67
.0
- 36
.0
- -
- -
2.1
2.05
-
- 33
.0
P3 W
idth
of
firs
t rid
ge
58.0
55
.0
- -
- -
- -
- -
- -
42.5
Wid
th o
f se
cond
rid
ge
70.0
73
.0
- -
- -
- -
- -
- -
-
Ena
mel
thic
knes
s 4.
0 3.
8 -
- -
- -
- -
- -
- -
L
engt
h of
the
toot
h 89
.0
89.0
43
.0
50.0
57
.0
70.0
70
.0
48.0
1.
8 -
2.9
56.0
-
W
idth
of
the
toot
h 80
.0
81.0
-
42.0
-
- -
- 1.
7 -
2.6
42.0
-
P4 W
idth
of
firs
t rid
ge
76.0
78
.0
- -
47.0
-
- -
- -
- -
-
Wid
th o
f se
cond
rid
ge
81.1
83
.0
- -
- -
- -
- -
- 45
.8
-
Ena
mel
thic
knes
s 4.
0 4.
0 -
- -
- -
- -
- -
- -
L
engt
h of
the
toot
h 10
7.5
107.
0 72
.0
67.0
68
.0
60.0
2.
34
1.8
4.0
67.0
-
W
idth
of
the
firs
t rid
ge a
t the
bas
82
.0
81.5
47
.0
- -
- 2.
4 2.
1 2.
8 47
.0
-
Wid
th o
f th
e fi
rst r
idge
at t
he to
p
- -
- -
-
-
- -
- -
-
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 81
.0
81.5
50
.0
41.0
45
.0
- -
- -
- -
M1
Wid
th o
f th
e se
cond
rid
ge a
t the
top
- -
- -
-
-
- -
- -
-
Wid
th o
f th
e th
ird
ridg
e at
the
base
75
.0
75.5
-
- -
- -
- -
- -
W
idth
of
the
thir
d ri
dge
at th
e to
p -
- -
- -
- -
- -
- -
W
idth
of
post
erio
r ta
lon
- -
- -
-
-
- -
- -
-
Ena
mel
thic
knes
s 4.
3 4.
2 -
- -
- -
- -
- -
L
engt
h of
the
toot
h 11
5.5
112.
5 67
.0
64.0
71
.0
59.0
2.
4 2.
90
3.9
73.0
-
W
idth
of
the
firs
t rid
ge a
t the
bas
e 10
3.0
110.
5 57
.0
54.0
-
- 2.
15
2.15
3.
5 65
.0
-
Wid
th o
f th
e fi
rst r
idge
at t
he to
p 80
.0
81.0
-
- 56
.0
- -
- -
- -
M2
Wid
th o
f th
e se
cond
rid
ge a
t the
bas
e 97
.5
107.
0 58
.5
53.0
-
- -
- -
58.0
-
W
idth
of
the
seco
nd r
idge
at t
he to
p 83
.0
75.5
-
- -
- -
- -
- -
W
idth
of
post
erio
r ta
lon
- -
- -
-
-
- -
- -
-
Ena
mel
thic
knes
s 4.
6 4.
5 -
- -
- -
- -
- -
L
engt
h of
the
toot
h 12
0.0
118.
0 68
.0
70.0
77
.0
72.0
2.
9 3.
18
- 65
.0
73.0
Wid
th o
f th
e fi
rst r
idge
at t
he b
ase
106.
0 10
5.0
59.5
55
.0
-
-
2.25
2.
45
- -
-
Wid
th o
f th
e fi
rst r
idge
at t
he to
p
85.0
82
.0
- -
-
-
- -
- -
- M
3 W
idth
of
the
seco
nd r
idge
at t
he b
ase
95.0
10
0.0
- 51
.0
-
-
- -
- -
-
Wid
th o
f th
e se
cond
rid
ge a
t the
top
78.5
80
.0
- -
-
-
- -
- -
-
Wid
th o
f po
ster
ior
talo
n 64
.0
67.0
-
- -
- -
- -
- -
E
nam
el th
ickn
ess
5.2
5.3
- -
-
-
- -
- -
-
24
PLATE VII
Deinotherium thraceiensis sp. n.1. Part of vertebral column (vertebrae, ribs). Scale ca. 5%Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE VIII
Deinotherium thraceiensis sp. n.1—3. Atlas. Scale ca. 13.6%. Coll. SU.M. No. SU Pl 312/3-14, 5. Axis. Scale ca. 13.6%. Coll. SU.M. No. SU Pl 312/3-2Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE IX
Deinotherium thraceiensis sp. n.1. Third cervical vertebra. Coll. SU.M. No. SU Pl 312/3-32. Fourth cervical vertebra. Coll. SU.M. No. SU Pl 312/ 3-43. Fifth cervical vertebra. Coll. SU.M. No. SU Pl 312/3-54. Second thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-95. Sixth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-13Scale 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/ 3-162. Tenth thoracic vertebra. Coll. SU.M. No. SU Pl 312/ 3-173. Eleventh thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-184. Twelfth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-195. Thirteenth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-206. Fourteenth thoracic vertebra. Coll. SU.M. No. SU Pl 312/3-21Scale 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 312/3-324. First lumbar vertebra. Coll. SU.M. No. SU Pl 312/3-22Scale 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 312/62, 2a. Left scapula in medial and lateral view. Coll. SU.M. No. SU Pl 312/5Scale 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 312/82. Left humerus in cranial view. Coll. SU.M. No. SU Pl 312/7Scale 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 312/102, 2a. Right radius in lateral and medial view. Coll. SU.M. No. SU Pl 312/12Scale ca. 6.2%; Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XV
Deinotherium thraceiensis sp. n.Carpals1. Cuneiforme (ulnare) sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-12. Lunare sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-23. Magnum sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-74. Unciforme sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-85. Pisiforme sin. Scale ca. 15.6%. Coll. SU.M. No. SU Pl 312/13-4Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE VII
PLATE VIII PLATE IX
PLATE X PLATE XI
PLATE XII PLATE XIII
PLATE XIV PLATE XV
PLATE XVI PLATE XVII
PLATE XVIII
PLATE XX
PLATE XIX
PLATE XXI PLATE XXII
PLATE XXIII
PLATE XXIV
25
PLATE XVI
Deinotherium thraceiensis sp. n.1, 1a. Metacarpus I of the first digit in frontal and lateral view. Scale ca. 18.6%. Coll. SU.M. No. SU Pl 312/14-12, 2a. Metacarpus II of the second digit in frontal and lateral view. Scale ca. 18.6%. Coll. SU.M. No. SU Pl 312/14-23, 3a. Second phalanx of the second digit in frontal and lateral view. Scale ca. 14.1%. Coll. SU.M. No. SU Pl 312/15-2Locality: 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 312/14-52, 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 312/15-84. Patella. Scale ca. 22.2%. Coll. SU.M. No. SU Pl 312/18-2Locality: Ezerovo, near Plovdiv; Level: Maeotian
PLATE XVIII
Deinotherium thraceiensis sp. n.1. Left hand. Scale ca. 7.7%. Coll. SU.M. No. SU Pl 312/13Locality: 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 312/3-32Locality: 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 312/16-1Locality: 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 312/17-1Scale 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 312/19-13. Right tibia (restored) in dorsal view. Coll. SU.M. No SU Pl 312/19-2Scale 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 312/142. Calcaneus sin. Scale ca 20.3%. Coll. SU.M. No SU Pl 312/21-1Locality: Ezerovo, near Plovdiv; Level: Maeotian.
PLATE XXIV
Deinotherium thraceiensis sp. n.Restored skeletonColl. SU.M. No SU Pl 312Locality: Ezerovo, near Plovdiv; Level: Maeotian
4 Geologica Balcanica, 3-4/2006
26
wide and regularly curved. In its upper part there isa wide deep groove, gradually becoming narrowerand shallower with the curving of the sympnysis.Ramus ascendens wide and thick. All the area ofproc. angularis is thick at the base and strongly pro-truding backwards. This thickness reaches as far asproc. articularis. The anterior part of ramus ascen-dens is significantly thinner, however, incl. proc. coro-noideus itself. All the ramus ascendens in this part islaterally slightly concave. Foramen mandibulae wideand deep. Proc. coronoideus almost vertical in theanteriror part. Tips of the processes curved back-wards; at the posterior ends there is a moderate con-cavity (incisure). Processus articularis thick and high.Perpendicularly positioned, with well shaped artic-ular surfaces. Two rami of the mandible not parallel.At the level of M1 a widening begins, rami comingcloser 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 sym-physis, shaping the curve together with it. Their basisstarts deep inside the symphysis. There they are hollow,their alveoli are of semi-circular shape (Fig. 8).
At the same time they are becoming thicker in-side. At the end of the symphysis the tusks are al-ready wholly solid. They gradually become thinnerand, curving in two directions – outwards and back-wards, go apart from each other. Their tips are point-ed, slightly smoothed only at the foremost part of theinner side, but there is no clear flat surface. Theirtips end just below the end of processus angulare.
Lower cheek-teeth. (Pl. VI). The toothrows also havefive teeth each – two premolars and three molars.Unlike the upper premolars, the lower are muchnarrower, and their structure is very different,especially P3.
P3 has a long and narrow crown, pointed at theanterior end. One large longitudinal ridge is situat-ed along its axis, and another, transversal – in theposterior part. They form the occlusial surface ofthe tooth. The longitudinal ridge is wider at its basis,gradually narrowing towards the apex of the crown.Its widest part is in the middle of the crown. There itis more worn out and part of the dentine is seen. Onthe inner part of the tooth, at the place where thelongitudinal ridge contacts the inner one, the firstnarrows abruptly thus shaping a large triangularvalley. Outside the crown is smooth.
P4 dext has a larger crown, elongated and wide.The occlusal surface is moderately worn. It is formedby two transversal ridges, contacting at the outer partof the crown. At this place they divide the valley be-tween them in two not quite equal parts. The anteriorwall of the first ridge is strongly concave. The poste-rior part of the crown has a weak cingulum.
M1 dext has three transversal ridges. First two aremore worn. Their structure is generally the same asin M1. All three ridges are wider on the outer side.The valleys dividing them become gradually narrowertowards the outer part. In this part the anterior wallsof the second and third ridge are slightly convex but
Fig. 8. Shape of the tusks of Deinotherium thraceisensis sp. n.A – shape of the tusks at the basis of the alveolus; B – shape ofthe tusks at 115.0 mm from their basis in the symphysis; C –cross-section of I dext at 600.0 mm from the symphysis –natural size
they don’t touch the ridge in front of them. On theinner side, at the bottom of each valley there is a weaktubercle, and on the anterior side – a small cingulum.Left M1 resembles the right but is more worn.
M2 dext is large and tetragonal. It has two thicktransversal ridges, wider on the outer side. There bothridges form a small concavity. The valley betweenthem is deep and free but on the inner side there is asmall tubercle. On its anterior and posterior sides thecrown has a cingulum.
M3 dext is large too but with an irregular tetrago-nal shape. Built by two transversal ridges. The first iswider. Both are widening outwards. At the ends –inner and outer – they are slightly curved to the front,forming with their anterior walls shallow valleys. Thevalley between them is deep and unblocked. Theposterior talon on the inner side of the crown is formedby numerous tubercles of different size.
Comparison. The structure of the described mandi-ble is close to the other species (Fig. 9).
The symphysis is moderately large. There are dif-ferences in the shape of ramus horizontalis, proc.articularis, proc. coronoideus and the tusks.1. The symphysis of D. bavaricum H. v. Meyer is moreprotruding. In D. levius Jourdan it is more roundedand curved inwards and in D. giganteum Kaup, aswell as in D. gigantissimum Stefanescu and D. thra-ceiensis sp. n. the curving is even stronger and moregradual.
27
2. In D. thraceiensis sp. n. ramus horizontalis (in theregion of proc. angualris) strongly protrudes back-wards with a pronounced curve. It almost reachesthe level of incisura mandibulae.3. Ramus ascendens is almost perpendicular to ra-mus horizontalis in its anterior part.4. Processus articularis is high and abruptly separat-ed from ramus ascendens, forming with it an obtuseangle in its posterior part. Anteriorly it descendssteeply toward incisura mandibulae.5. Processus coronoideus is high too and anteriorlyalmost vertical, nearly at the same level as proc. ar-ticularis.6. Incisura mandibulae is deep and almost horizon-tal in relation to the mandible, perpendicular to ra-mus ascendens.
In D. gigantissimum Stefanescu the curve at proc.angularis is weaker. Poc. articularis is not so abrupt-ly separated from it. Proc. coronoideus posteriolyturns into incisura mandibulae. The latter is very
shallow. The symphysis is some more elongated, butit has been restored. No P3 is known for this species.7. The third premolar – P3 in D. thraceiensis sp. n. isbuilt mainly by one longitudinal ridge and one weaktransversal ridge on the posterior inner side. The lon-gitudinal crest touches the inner one. No cingulum.
In D. giganteum Kaup there is a bending tubercleon the anterior inner side of P3 which surrounds adeep valley together with the longitudinal ridge. Ahigh cingulum encircles the crown.
In D. bavaricum H. v. Meyer P3 is smaller andbuilt by two smaller transversal ridges and a largelongitudinal one, positioned on the outer side of thecrown.8. P4 in D. thraceiensis sp. n. has two transversal ridg-es connected on the outer side. Cingulum is weakand only on the posterior side.
In D. giganteum Kaup the structure of the tooth isthe same, but with well developed cingulum on theinner side of the crown.
In D. gigantissimum Stefanescu P4 has the samestructure as D. giganteum Kaup but without the largecingulum.
P4 in D. bavaricum H. v. Meyer has two ridges.The posterior is larger, with two parts at a right angleto each other. It is elongated and touches the poste-rior wall of the anterior ridge. The latter is “C-shaped”. It surrounds a tetragonal valley anteriorly.Strong cingulum.
As a whole, the fourth premolars in the studiedtaxa are rather close in their structure.9. M1, M2 and M3 in the species described show nosignificant differences when compared to the otherstudied species. A small exception is the fact thatour species lacks a developed cingulum. The dis-placement of the third ridge in M3 is not regarded asa distinct taxonomic peculiarity by us.
We had no opportunity to compare the materialto the species D. cuvieri Kaup, D. levius Jourdan, D.indicum Falconer, D. pentapotamiae Lydekker.
Vertebral column (Pl. VII).In D. thraceiensis sp. n. it consists of 7 cervical, prob-ably 14 thoracic, probably 14 lumbar, 3 sacral andprobably 6 caudal vertebrae.Atlas (Pl. VIII, Figs. 1, 2 and 3).Almost entirely preserved. Processes are only partial-ly broken, without losing the shape of the vertebra. Itconsists of two arches – dorsal and ventral, limitinga large foramen vertebrae. The foramen widens down-wards and is slightly constricted in the middle. There’sno corpus vertebrae. In fact the atlas is a wide bonering. The transverse processes have become large boneplates – wings of the atlas. There is no spinous pro-cess. The atlas articulates with the skull by wide ovalconcavities, cranially positioned on the arches. Cau-dally, on the ventral arch, there is an articular sur-face for the dens of the axis. Foramen ovale and for.ventrale lateralis on both sides of the vertebra andlarge (measurements in Table 5).Axis (Pl. VIII, Fig. 4 and 5). Body narrow and oval,spinous process high, wide and long. Two strong
Fig. 9. Comparison of deinothere mandibles: a – Deinotheri-um bavaricum H. v. Meyer; b – Deinotherium levius Jourdan;c – Deinotherium giganteum Kaup; d – Deinotherium thra-ciensis sp. n.
28
wards the spinous process. All other thoracic verte-brae resemble the second. There are differences onlyin the size of proc. transversus and proc. spinosus,which are smaller in each next vertebra. From thefirst to the twelfth the size of the corpus and foram-ina decreases, then increases to the last. In the fif-teenth and all the following there is no fovea costal-is. So, there were no ribs attached to these vertebrae(measurements in Table 6).
Lumbar vertebrae (Pl. XI, Fig. 4)Resemble the last thoracic vertebrae. Here, too, foveacostalis is missing. The diameter and thickness ofcorpus vertebrae gradually decreases from the firstto the last. Spinous processes become shorter. As inthe thoracic vertebrae, they are inclined backwards.In the other vertebrates these processes on the lum-bars are pointing anteriorly. This peculiarity shouldn’tbe of taxonomic importance for genus Deinotheriumbecause it is present in the other proboscideans aswell. The transverse processes are not long as shouldbe expected for lumbars so we cannot speak of proc.costarius. These peculiarities in the development ofthe processes of the lumbars and the lack of foveacostalis in the last thoracic vertebrae make it diffi-cult for us to decide unequivocally where one groupof vertebrae ends and the other begins. This is whywe cannot say for sure how many the thoracic andcorrespondingly the limbar vertebrae are. The diam-eters of foramen vertebrae increase as in the last tho-racic vertebrae. They are largest in the last vertebra(Table 7). This character too creates difficulties inthe determination of the two groups of vertebrae but
crests on tip with a deep furrow between them. Densepistrophei connects the axis with the atlas. Fora-men vertebrae much smaller than in atlas, tetragonal(measurements in Table 5).Third, fourth and fifth cervicals (Pl. IX, Figs. 1, 2and 3). All have similar structure. Narrow bodies.Cranially slightly convex, caudally slightly concave.Transverse processes small and rounded in the ends.Transverse foramina limited by the bodies and thetransverse processes. Well pronounced articular sur-faces on the cranial and caudal processes. Foramenvertebrae wide and high. Spinous processes not pro-nounced.
Sixth and seventh cervicals are built in the sameway as the previous three, but they have spinous pro-cesses, largest on the seventh. Below the transverseprocesses the seventh cervical has a well shaped foveacostalis – a concavity for the head of the first ribwhich is between the seventh and the eighth vertebra(measurements of all cervicals in Table 5).
Thoracic vertebrae (Pl. IX, Fig. 4 and 5, Pl. X, Fig. 1,2, 3, 4, 5, and 6).The first thoracic vertebra is very similar to the lastcervical, only its spinous process is larger. Thus itresembles the second thoracic vertebra, where thisprocess is most developed. The second thoracic ver-tebra (Pl. IX, Fig. 4) has a distinct structure. Its bodyis small, cranially convex, caudally – slightly round-ed. Its cranial articular processes are less develpedthan the caudal. Fovea costalis cranialis larger thanf. c. caudalis. Spinous process strong, widening atthe tip, tetragonal. Foramen vertebrae elongated to-
Table 5Cervicals
Table 6Thoracics
THORACICS № Dimensions 1 2 3 4 5 6 7 8 9 10 11 12 13 14 1. Diameter of corpus
vertebrae 250.0 240.0 230.0 220.0 210.0 200.0 196.0 192.0 188.0 186.0 182.0 180.5 180.0 179.0
2. Thickness of corpus vertebrae
102.0 96.0 92.0 89.0 85.0 86.0 87.0 88.0 89.0 90.0 90.5 91.0 92.0
3. Length at proc. spinosus 400.0 396.0 390.0 384.0 380.0 362.0 340.0 320.0 310.0 295.0 290.0 287.0 280.0 4. Length of proc. spinosus 205.0 610.0 580.0 550.0 532.0 510.0 500.0 460.0 400.0 340.0 290.0 289.0 288.0 285.0 5. Length of proc.
transversus 93.0 102.0 112.0 117.0 120.0 116.0 112.0 110.0 106.0 100.0 96.0 94.0 87.0
6. Length of foramen vertebrae
110.0 97.0 92.0 88.0 83.0 80.0 70.0 62.0 54.0 46.0 40.0 43.5 47.0 58.0
7. Width of foramen vertebrae
78.0 70.0 62.0 50.0 41.0 30.0 38.0 45.0 52.0 57.0 60.0 60.0 60.5 59.0
CERVICSLES № Dimensions, mm Atlas Axis 3 4 5 6 7
1. Height of the vertebra 300.0 340.0 330.0 340.0 343.0 350.0 352.0 2. Width of the vertebra 400.0 320.0 310.0 315.0 218.0 224.0 226.0 3. Length of corpus vertebrae 212.0 230.0 238.0 240.0 246.0 250.0 4. Width of corpus vertebrae 170.0 195.0 202.0 206.0 208.0 210.0 5. Length of articular surfaces 100.0 103.0 105.0 106.0 108.0 6. Height of processus spinosus 60.0 150.0 7. Length of foramen vertebrae 200.5 165.0 60.0 80.0 8. Width of foramen vertebrae 145.0 93.0
29
it probably has its functional meaning. It should berelated to the use of the hindlimbs. The accumula-tion of nerve tissue close to the pelvis probably facil-itated the movements of these huge “columns” – thehindlimbs.
Sacral vertebrae (Pl. XI, figs. 1, 2 and 3). Three, fused,building a single bone – the sacrum. Body of thesacrum large, with wide, slightly rounded wings. Twoforamina sacralia dorsalia on each of the two sidesof the wings. Spinous processes of the first twovertebrae smaller and narrower, on third – highestand widening in its upper part. Two foramina oneach side ventrally, by the attachment of thevertebrae. Of these four ventrally situated foramina,the first two are wider, penetrating inside to thesacral canal. Body of the sacrum oval, anteriorlyslightly concave and protruding in front of thewings.
Caudal vertebrae. Preserved are only six original cau-dals, but on the mounted skeleton there are 21. Thefirst, those after the sacrum, resemble the last lum-bars, but their processes are gradually decreasing insize. In the first, a chanal is still present, which grad-
ually disappears. The body of the caudals is elon-gated, narrow and convex on both sides.
Comparison. Many of the vertebrae in the skeletonswe compare our materials with are reconstructedwhich makes the comparison difficult. Nevertheless,one can see that in our specimen the spinous pro-cesses of the cervical vertebrae are weaker. These pro-cesses are strong and high in all known skeletons. Adifference in D. thraceiensis sp. n. is the large tetrag-onal widening of the tip of this process on the sec-ond thoracic. There are no other visible differences.The number of the vertebrae (mainly of the thoracicsand the caudals) varies. In fact their number is un-known. As far as we know, there are no finds of awhole tail.
Ribs (costae). Fragments of seven pairs of ribs werefound. As far as the fifteenth vertebra lacks foveacostalis, the ribs must have been 14. First rib is wideand thick. One half of caput costae articulating withthe last cervical is oval, the other half – smaller andspherical. Caput costae of the second vertebra hasthe same structure but its neck is narrow and short.Its body is wide and flat. The third rib is narrower,
Table 8Sacrum
Table 9Caudal vertebrae – vert. coccigiae
№ Dimensions mm 1. Total length 315.0 2. Width at the wings 405.0 3. Anterior width of the corpus (at first fused vertebra) 180.0 4. Posterior width of the corpus (at third fused vertebra) 140.0 5. Anterior height of the corpus (at first fused vertebra) 118.0 6. Posterior height of the corpus (at third fused vertebra) 50.0 7. Height of processus spinosus 155.0 8. Anterior width of canalis sacralis (at first fused vertebra) 162.0 9. Posterior height of canalis sacralis (at third fused vertebra) 64.0
№ Dimensions, mm 1 2 3 5 7 8 1. Height of the vertebra 180 175 155 150 130 130 2. Width of the vertebra 290 240 180 190 190 190 3. Diameter of corpus vertebrae 110 109 109 100 100 100 4. Thickness of corpus vertebrae 106 104 94 85 90 90 5. Height of processus spinosus 100 90 - - - - 6. Vertical diameter of foram. vertebrae 60 60 60 40 - - 7. Horizontal diameter of foram. vertebrae 100 100 60 50 - -
Table 7Lumbars
№ Dimensions 1 2 3 4 5 6 7 8 9 10
1. Diameter of corpus vertebrae 178.0 177.0 176.0 174.0 172.0 170.0 168.0 165.0 162.0 160.0 2. Thickness of corpus vertebrae 93.7 95.0 97.0 97.5 98.0 100.0 100.0 98.0 96.0 94.0 3. Length at proc. transversus 274.0 269.0 264.0 258.0 250.0 240.0 240.0 242.0 243.0 245.0 4. Length of proc. spinosus 282.7 280.5 277.8 275.5 273.0 270.0 268.0 266.5 262.0 258.0 5. Length of proc. transversus 82.0 77.0 73.0 69.8 66.5 62.0 63.0 63.0 64.5 66.0 6. Length of foramen vertebrae 64.0 75.0 81.0 90.0 102.0 110.0 120.0 130.0 140.0 150.0 7. Width of foramen vertebrae 58.5 60.0 61.0 59.5 60.0 60.0 61.0 61.5 62.0 63.0
30
Table
10
Rib
s – co
stae
№
Dim
ensi
ons,
mm
1
2 3
4 5
6 7
8 9
10
11
12
13
14
1.
Tot
al le
ngth
12
45.0
13
90.0
14
50.0
47
2.0
148.
0 14
30.0
13
76.0
13
22.0
12
74.0
12
30.0
11
95.0
11
02.0
95
0.0
840.
0 2.
D
iam
eter
of
capu
t cos
tae
80.0
82
.0
96.0
68
.0
69.9
70
.5
68.5
67
.0
64.0
62
.3
60.0
57
.2
54.0
50
.0
3.
Dia
met
er a
t the
th
icke
ning
98
.0
105.
0 81
.0
85.0
67
.0
54.5
52
.0
56.3
48
.1
46.5
45
.0
46.0
45
.5
45.0
4.
Max
imum
wid
th o
f th
e ri
b 80
.0
153.
0 11
4.0
91.0
75
.0
60.0
60
.5
60.9
61
.3
61.7
62
.0
64.5
67
.3
70.0
5.
Dis
tanc
e be
twee
n th
e be
ginn
ing
of c
aput
co
stae
to th
e th
icke
ning
50.0
14
0.0
120.
0 13
6.0
162.
0 19
4.0
198.
0 20
3.5
206.
0 21
3.0
220.
0 19
5.0
180.
0 16
0.0
Table
11
Sca
pula
Note
: T
he m
easurm
ents
in t
he t
able
are t
aken f
or t
he l
eft
rib
s
D. t
hrac
eien
sis
sp.n
. E
zero
vo, B
ulga
ria
№
Dim
ensi
ons
sin
dext
D. g
igan
tiss
imum
St
. Man
zati,
Rom
ania
St
efan
escu
D. g
igan
teum
Kau
p Pi
kerm
i D
. gig
ante
um
Kau
p
D. g
igan
teum
Kau
p M
unch
en -
Str
omer
D
. gig
ante
um K
aup
Mun
chen
- S
trom
er
1.
Tot
al le
ngth
11
35.0
11
35.0
97
0.0
730.
0 -
756.
0 -
2.
Wid
th c
-d
500.
0 50
0.0
300.
0 52
5.0
- 57
0.0
- 3.
M
axim
um w
idth
m-l
89
5.0
895.
0 65
0.0
- -
4.
Max
imum
wid
th o
f l
650.
0 65
0.0
-
- 5.
M
axim
um w
idth
of
m
245.
0 24
5.0
20
5.0
225.
0 6.
D
iam
eter
at t
he n
eck
254.
0 24
5.0
260.
0
-
- 7.
D
iam
eter
at t
he a
rtic
ular
sur
face
26
0.0
260.
0 28
0.0
170.
0 18
5.0
170.
0 25
0.0
8.
Hei
ght a
t the
art
icul
ar s
urfa
ce
96
.0
110.
0 11
0.0
160.
0 9.
L
engt
h of
tube
rosi
tas
spin
ae
635.
0 63
5.0
150.
0
-
- 10
. H
eigh
t of
acro
mio
n 21
5.0
215.
0 90
.0
- -
11.
Ang
le a
t l
85
o
31
also with a short and narrow neck as the first tworibs. The body of the fourth is long and almost round.Caput costae spherical, divided by a well pronouncedfurrow in the middle into two surfaces – facies ar-ticularis and facies articularis condilis, the first ofwhich is bigger. Near its distal end the rib graduallybecomes thinner. At the very end it becomes widerending with a rough surface for the attachment ofthe cartilage. The other ribs are similar to the fourth.Their length gradually decreases (measurements inTable 10).
Sternum. Only a fragment preserved, with a roughsurface and places for the attachment of the rib car-tilages.
Comparison. The most disputable problem concern-ing the ribs is their number. There is still no explicitopinion on that matter and we don’t know their pre-cise number. D. gigantissimum Stefanescu has 32 pairsof ribs restored, reaching up to the sacrum. Whatwas the reason for this number, we do not know. Bythe restoration of D. bavaricum H. v. Meyer 11 pairsof ribs were mounted. As we can see from Plate XXIV,they had 5th, 9th, 17th and 18th thoracic vertebrae.It seems they restricted themselves to the 12th.
Scapula (Pl. XII, Figs. 1 and 1a, 2 and 2a). Only theright is entirely preserved. There are separate frag-
ments from the left, and it was restored after the right.Dorsally strongly flattened, becoming thinner at theanterior and posterior edge. Spina scapulae and theedges surround two concavities of unequal size. Thecrest takes almost the entire length of the scapula. Itis divided in two near the tubersitas spinae, forminga wide long groove starting immediately behind theacromion and ending in the dorsal thickening. Theacromion itself is high and curved towards the neckof the scapula. Fossa infraspina shallower but widerthan fossa supraspina. Margo thoracales long andpointing outwards. It has a regular triangular shape.The articular surface for the attachment to the hu-merus is shallow.
Humerus (Pl. XIII, Figs. 1. 1a and 2). Both humerientirely preserved. Caput humeri large and curved,with no distinct neck. Tuberculum majus protrudesupwards, above caput humeri. Tuberculum minus ismuch smaller. Positioned cranio-medially. Betweenthe two tubercles there is a deep and wide furrow –sulcus intertubercularis. Shaft of the bone roundand smooth with weak sulcus spiralis. It starts on thedorsal surface above the caput and ends in front onthe distal part over the joint. In the distal part of thebone is trochlea humeri, with two large projections,the lateral being the higher. In front, above them, isthe wide and shallow fossa radialis, and on the pos-terior side – fossa olecrani. It is deeper and wider.
Table 12Humerus
D. thraceiensis sp.n. Ezerovo,
Bulgaria № Dimensions, mm
sin dext
D. gigantissimum St. Manzati,
Romania
D. giganteum Kaup Saint
Yean – Lartet
D. giganteum Kaup München –
Stromer
D. giganteum Kaup München – Stromer
№ 495
1. Total length 1142.0 1146.0 1100.0 740.0 1090.0 900.0 2. Width of caput humeri 295.0 292.0 170.0 120.0 - - 3. Width in proximal part 325.0 320.0 370.0 280.0 - - 4. Width in diaphysis 190.0 195.0 - 124.0 115.0 90.0 5. Width in distal part 355.0 350.0 310.0 240.0 - - 6. Width at epicondyles 290.0 296.0 300.0 316.0 290.0 200.0 7. Diameter of caput
humeri 200.0 200.0 - - 260.0 190.0
8. Diameter of trochanter major
202.0 207.0 260.0 108.0 260.0 210.0
9. Diameter of trochanter minor
185.0 190.0 140.0
Table 13Ulna
D. thraceiensic sp.n. Ezerovo,
Bulgaria № Dimensions, mm
sin dext
D. gigantissimum St. Manzati, Romania – Stefanescu
D. giganteum Kaup Saint Yean – Deperet, 1892
D. giganteum Kaup München № 495 – Stromer,
1938
D. giganteum Kaup München –
Stromer, 1938
1. Total length 1150.0 1130.0 1050.0 - 850.0 1020.0 2. Width at proximal end proc. olecrani 320.0 300.0 300.0 320.0 - - 3. Height from proc. olecrani to proc.
styloideus 150.0 140.0 - - - -
4. Height of inc. semilunaris 127.0 120.0 - 150.0 - - 5. Width at inc. semilunaris – prox. end 96.0 90.0 - 50.0 120.0 115.0 6. Width at inc. semilunaris – dist. end 265.0 60.0 - - 260.0 225.0 7. Width in diaphysis 88.0 86.0 - 130.0 115.0 90.0 8. Width at distal end 255.0 50.0 -
32
The lateral concavity for the sinews is larger thanthe medial.
Ulna (Pl. XIV, Fig. 1 and 1a). Both bones are entirelypreserved. They are wholly identical. Tuber olecranilarge and curved medially. Processus anconeus slight-ly twisted and situated posteriorly, incisura semilu-naris shallow and wide. Lateral coronoid processesstrongly protruding sideways. Their upper surfacesform the whole semilunar incisure. Between them isincisura radialis, with coarse surface.
According to V. M. Svistun, such a structure ofthe elbow joint permitted different movements whenthe forelimb was in flexion.
The diaphysis is flat, the bone ending distally withwell shaped processus styloideus ulnae and cicum-ferentia articularis.
Radius (Pl. XIV, Fig. 2 and 2a). This bone is muchsmaller than the ulna. Slightly widening in its prox-imal part, with well pronounced articular surface.Slightly rounded in the diaphysis. Ends with a clear-
ly outlined articular surface. With this part it articu-lates with the ulna. Slightly convex laterally, in thedistal end.
Comparison. There are no significant differencesbetween the taxa of genus Deinotherium concerningthe structure of the scapula, humerus, ulna and ra-dius. According to Svistun, the ulna of the skeletonhe describes has a triangular shape of the diaphysisand is fused with the radius at ca. 1/3 of its length.There is no such thing in D. thraceiensis sp. n.
Carpus (Pl. XV, Figs. 1–6; Fig. 10 A, B).Made up of nine bones, arranged in two rows – prox-imal and distal. Five bones in the proximal: scaphoi-deum, lunare, cuneiforme, pisiforme and trapezium;four in the distal: trapezoideum, magnum, unciformeand centrale. Trapezium and trapezoideum are miss-ing and have been reconstructed. More bones aremissing from the right wrist, and have been restoredafter the left. Here we’ll describe only the left bones(measurements in Table 15).
Table 14Radius
D.thraceiensis sp.n. Ezerovo, Bulgaria № Dimensions, mm
sin dext
D.gigantissimum St. Manzati,
Romania Stefanescu
D.giganteum Kaup Pikermi – Gaudry, 1862
1. Total length 928.0 925.0 680.0 920.0 2. Width at proximal end 144.0 142.0 130.0 150.0 3. Width in diaphysis 95.0 95.0 40.0 - 4. Width at distal end 210.0 214.0 - 210.0
5. Length of the articular surfaces for inc. semilunaris 160.0 160.0 90.0 -
6. Length of same surface 130.0 132.0 - -
Fig. 10. Manus of Deinotherium thraciensis sp. n. A – left manus; B – right manus
33
Table
15
Carp
als –
carp
us
sin e
t dext
Table
16
Meta
carp
als
Dei
noth
eriu
m th
race
iens
is s
p.n.
sc
apho
id
luna
re
pisi
form
e cu
neif
orm
e m
agnu
m
cent
rral
e un
cifo
rme
№
Dim
ensi
ons,
mm
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
1.
Len
gth
187.
0 18
8.0
166.
0 16
0.0
206.
0 20
6.0
162.
0 16
0.0
110.
0 11
0.0
115.
0 11
5.0
162.
0 16
0.0
2.
Wid
th a
t pro
xim
al e
nd
104.
0 10
2.0
3.
Wid
th o
f th
e fa
cet f
or th
e ra
dius
11
0.0
113.
0 11
5.0
140.
0
4.
W
idth
of
the
face
t for
McI
10
6.0
104.
0
5.
H
eigh
t of
the
face
t for
the
radi
us
120.
0 14
0.0
6.
Hei
ght o
f th
e tu
berc
le
114.
0 15
0.0
7.
Lat
eral
hei
ght
77.0
78
.0
77.0
78
.0
8.
Ant
erio
r w
idth
86
.0
86.0
9.
L
engt
h of
the
face
t for
the
ulna
45
.0
43.0
10
. L
engt
h of
the
face
t for
the
cune
ifor
m
66.0
65
.0
11.
Wid
th o
f th
e fa
cet f
or th
e cu
neif
orm
15
0.0
150.
0 12
. L
engt
h of
the
proc
essu
s
15
0.0
150.
0
13
. L
engt
h of
pro
xim
al a
rtic
ular
sur
face
14
0.0
140.
0
14
. W
idth
of
prox
imal
art
icul
ar s
urfa
ce
142.
0 14
0.0
15.
Wid
th o
f th
e bo
ne
100.
0 10
2.0
98.0
97
.0
100.
0 10
0.0
16.
Len
gth
of th
e fa
cet f
or o
s lu
nare
16
0.0
160.
0
17
. W
idth
of
the
face
t for
os
luna
re
98.0
98
.0
Dei
noth
eriu
m th
race
iens
is s
p.n.
Mc-
I M
c-II
M
c-II
I M
c-IV
M
c-V
№
D
imen
sion
s, m
m
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
1.
L
engt
h 22
6.0
225.
0 23
0.0
230.
0 24
8.0
250.
0 23
2.0
230.
0 20
5.0
205.
0 2.
W
idth
at p
roxi
mal
end
14
0.0
140.
0 12
5.0
125.
0 11
2.0
112.
0 98
.0
108.
0 14
6.0
145.
0 3.
W
idth
at d
ista
l end
11
5.0
115.
0 10
8.0
125.
0 11
4.0
113.
0 10
9.0
98.0
97
.0
97.0
4.
W
idth
at d
ista
l sin
ew p
roce
sses
-
- 10
4.0
108.
0 10
5.0
106.
0 10
3.5
103.
5 11
4.0
113.
0 5.
L
engt
h of
sm
all f
acet
-
- 12
6.0
108.
0 13
5.0
135.
0 11
5.0
115.
0 14
0.0
140.
0 6.
L
engt
h of
larg
e fa
cet
- -
110.
0 12
6.0
136.
0 13
8.0
- -
112.
0 14
0.0
5 Geologica Balcanica, 3-4/2006
34
Scaphoideum (radiale) (Fig. 10 A, B No.1). Long,narrow and almost flat with rough lateral surface.Two large facets on the medial surface. Proximalcontacts lunare, distal – centrale. A facet on thevery proximal end, articulating with the distal end ofthe radius. The lower end of the scaphoid articu-lates with the fifth metacarpal.Lunare (Fig. 10 A, B No. 2). Triangular. Anteriorsurace rounded. Two concave facets in the proximalpart, articulating with the distal parts of the ulnaand radius. Lateral larger. Two slightly concave fac-ets in the distal part articulate with magnum andcentrale. The lateral facet of the scaphoid and themedial of the cuneiform are divided in the middleby a crest for the attachment of sinews.Cuneiforme (ulnare) (Fig. 10 A, B No. 3). The bonehas a peculiar form. It is flat. Medially, a large pro-cessus turns down to the proximal part of McI. Onthe inner side of this processus is os unciforme, whichis part of the distal row. In the proximal posteriorpart there is a small facet for the pisiform. The ante-rior and the medial surfaces of the bone are rough.Pisiforme (Fig. 10 A, B No. 4). Smallest bone of thewrist. Its anterior part contacts the ulna and the cu-neiform by two pronounced facets. Rounded in theother parts. Slightly curving downward at the very end.Magnum (Fig. 10 A, B No. 7). Almost square, elon-gated anteriorly. Situated in the middle of the distalrow, between os unciforme and os centrale. Two fac-ets on the distal part. Medial larger, touching theproximal part of McIII, the other touches a smallpart of McIV. Surface rough in the anterior part,and the facets for articulation with os unciforme andos centrale are concave in the middle – this is theplace where sinews are attached.Unciforme (Fig. 10 A, B No. 8). Largest of the distalrow. Tetragonal. Slightly rounded anteriorly, elon-gated posteriorly. The proximal acet for the cunei-form is large and medially inclined. Two facets indistal end, medial larger and contacting McII, later-al – McIII.Centrale (Fig. 10 A, B No. 9). Although slightly elon-gated in the ends, it also has a square shape. Situat-ed between the magnum and the scaphoid. Distalsurface slightly concave, entirely lying on the proxi-mal articular surface of McIV. Facet for articulationwith the magnum concave in the middle with a placefor the attachment of sinews.
Metacarpus (Pl. XVII; Fig. 10 A, B; No. I, II, III, IVand V). Here, as with the wrist, more bones are pre-served from the left forelimb.First metacarpal McI. Long, wide and flat. Facet forarticulation with the cuneiform and the unciformnarrow and long. Slightly convex in the proximal end,slightly concave in the middle. Large rounded artic-ular surface on distal end. Strong sinew processes inposterior and anterior parts.Second metacarpal McII. Also long, but narrow.Rounded in the anteriror part, strongly elongated pos-teriorly, gradually becoming narrower. Facet for unci-form in the proximal end long and wide. Laterally to it
a facet for McIII, smaller than the first. Slightly con-cave diaphysis, distal end widening with a large artic-ular surface for the first phalanx of the second digit.Sinew processes strong and on both sides of the bone.McIII and Mc IV resemble McII. Facets in proximalparts for magnum, unciform and central.McV is a mirror image of McI. Facet for the scaphoidin anterior part.
Phalanxes (Pl. XVII and XVIII; Fig. 10 A and B No.1, 2 and 3).First phalanx of the first digit PH-I,1 is medium-sized,with arounded, elongated flat shaft. Deep articularsurface on proximal end for the distal surface of McI.Phalanx rounded in posterior part. Narrowing dia-physis, distal end thicker and wider. Articular facetfor the next phalanx rounded and convex. Above andparallel to it the sinew concavity, weakly pronounced.First phalanx of second digit PH-II,1 resembles McII.PH-III,1 and PH-IV,1 do not differ from PH-II,1, butPH-V,1 is a mirror image of PH-I,1.Second phalanx of the first digit PH-I,2 is smallerthan the other phalanxes. In its posterior part, by thesinew crests, it is more convex, and concave in themiddle. Proximal articular surface concave, distalrounded and also concave in the middle.
PH-II,2 is also smaller than the other phalanxes.It looks like a second phalanx of a horse, i.e. shortand wide. Very thick proximal end, thinner in themiddle, thick again distally at the articular surface.
PH-III,2 and PH-IV,2 look like PH-II,2. PH-V,2 isa mirror image of PH-I,2.
PH-I,3, PH-II,3, PH-III,3, PH-IV,3 and PH-V,3,third phalanxes of all digits have almost the samestructure. Only their size is different, so we’ll describejust one phalanx.
D. thraceiensis sp. n. has a long and large hoof.In its proximal part it is slightly convex, in the mid-dle and in the distal part – concave. Large articularsurface for the second phalanx on the posterior part.Slightly rounded in front. Measurements of the dig-its are given in Table 17.
Height of the entire manus (wrist and digits) is460.0 mm; diameter with spread digits – 902.0 mm.
ComparisonAll carpal and metacarpal bones, as well as the pha-lanxes are similar in D. giganteum Kaup, D. gigantis-simum Stefanescu and D. bavaricum H. v. Meyer.No taxonomically significant differences were found.Maybe only the hoofs show some. It was already saidthat in D. thraceiensis sp. nov. they are long. Theirwalking surface is flat and concave in the middlewhich means they were touching the ground with alltheir surface. In the other species the hoofs seem totouch the ground in most cases only with their ante-rior part.
Pelvis (Pl. XIX, Fig. 1; Pl. XX, Fig. 1). Both hipbonesare entirely preserved.The wing of the hipbone is a wide S-shaped curvedplate, anteriorly rounded, with a pronounced pubic
35
Table
18
Pel
vis
D. t
hrac
eien
sis
sp.n
. E
zero
vo, B
ulga
ria
№
Dim
ensi
ons,
mm
sin
dext
D. g
igan
tissi
mum
St
. Man
zati,
Rom
ania
–
Stef
anes
cu
D. g
igan
teum
Kau
p M
ünch
en –
St
rom
er, 1
938
D. g
igan
teum
Kau
p M
ünch
en –
Str
omer
, 19
38
D. g
igan
teum
, Kri
voi
Rog
, Mol
dova
–
Белокрые,
196
0
1.
Len
gth
of w
ing
d-c
1110
.0
1165
.0
1130
.0
1000
.0
780.
0
2.
Tot
al le
ngth
of
the
win
g h-
d 11
60.0
11
75.0
11
80.0
-
-
3.
Len
gth
of w
ing
to f
or. o
btur
atum
82
0.0
740.
0 90
0.0
830.
0 56
0.0
4.
L
engt
h of
for
. obt
urat
um
266.
0 28
0.0
- -
- 24
5.0
5.
Wid
th o
f fo
r. o
btur
atum
12
6.0
150.
0 -
- -
6.
D
iam
eter
at t
he a
ceta
bulu
m
190.
0 18
8.0
- -
- 23
0.0
7.
Hei
ght o
f sa
cral
pro
cess
32
0.0
320.
0 -
- -
250.
0 8.
L
engt
h of
pel
vic
sym
phys
is
462.
0 46
0.0
- 47
0.0
340.
0
9.
Hei
ght o
f sa
me
385.
0 27
5.0
- -
- -
Table
17
Bones
of
the
dig
its
– f
ore
lim
b
Dei
noth
eriu
m th
race
iens
is s
p. n
. 1-
st d
igit
2-
nd d
igit
3-rd
dig
it
1 2
3 1
2 3
1 2
3 №
D
imen
sion
s, m
m
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
1.
L
engt
h 11
0 11
0 10
0 10
0 98
96
15
0 15
0 80
80
13
0 13
0 17
8 18
0 10
5 10
2 16
7 16
7 2.
W
idth
at p
roxi
mal
end
12
5 12
5 91
91
66
68
10
4 10
5 10
6 10
7 95
95
.5
115
116
108
108
111
112
3.
Wid
th a
t dis
tal e
nd
88
88
82.6
83
-
- 94
95
97
97
-
- 10
7 10
7 10
2 10
3 -
-
Table
17
Bones
of
the
dig
its
– f
ore
lim
b (
continued
)
Dei
noth
eriu
m th
race
iens
is s
p. n
. 4-
th d
igit
5-th
dig
it 1
2 3
1 2
3 №
D
imen
sion
s, m
m
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
1.
Len
gth
178
180
101
102
120.
5 12
3 14
0 14
0 10
4 10
2 11
6 11
7.5
2.
Wid
th a
t pro
xim
al e
nd
110
111
96
98
87
88
95
96
96
96
96
97.5
3.
W
idth
at d
ista
l end
96
96
90
89
-
- 90
90
80
80
-
-
36
D. thraceiensis sp.n. Ezerovo, Bulgaria № Dimensions, mm
sin dext
D. gigantissimum St. Manzati, Romania –
Stefanescu
D. giganteum Kaup München –
Stromer, 1938
D. giganteum Kaup München –
Stromer, 1938 1. Total length 1650.0 1650.0 1440.0 188.0 1060.0 2. Diameter of caput femuri 195.0 195.0 - 210.0 140.0 3. Height of caput femuri from [???] 160.0 160.0 - 4. Width at proximal end 375.0 375.0 350.0 400.0 295.0 5. Width in diaphysis 225.0 225.0 - 6. Width at distal end 330.0 340.0 440.0 315.0 175.0 7. Thickness at proximal end 150.0 150.0
Table 19Femur
Table 21Fibula
D. thraceiensis sp.n. Ezerovo, Bulgaria № Dimensions, mm
sin dext
D. gigantissimum St. Manzati, Romania –
Stefanescu
D. giganteum Kaup Pikermi – Gaudry, 1862
1. Total length 1125.0 565.0 830.0 870.0 2. Width at proximal end 90.0 72.0 70.0 70.0 3. Width in diaphysis 126.0 103.0 - 100.0 4. Width at distal end 275.0 250.0 170.0 140.0
5. Length of the articular surface for os calcaneus 80.0 72.0 - 70.0
6. Length of the small facet 76.0 60.0 - 55.0
Table 20Tibia
D. thraceiensis sp.n. Ezerovo, Bulgaria № Dimensions, mm
sin dext
D. gigantissimum St. Manzati, Romania –
Stefanescu
D. giganteum Kaup Pikermi – Gaudry, 1862
1. Total length 1200.0 1250.0 920.0 950.0 2. Width at proximal end 370.0 365.0 330.0 310.0 3. Thickness at proximal end 320.0 320.0 – – 4. Width in diaphysis 160.0 165.0 260.0 140.0 5. Width at distal end 300.0 330.0 260.0 340.0
crest. The wing significantly narrows posteriorly.Dorso-lateral surface smooth and slightly concave,medial almost flat. In the region of facies articularisthe surface is rough. Acetabulum well shaped. Fora-men obturatorium oval.
Femur (Pl. XXI, Fig. 1 and 1a). We found only theright thighbone. Left was restored after it. The boneis long and solid. Caput femuri large, almost paral-lel to the bone’s axis. A small neck separates it fromtrochanter major, which is laterally situated. Fossatrochanterica strongly pronounced. Shaft flat, slightlycurved medially and laterally. Two condyles in thedistal part of the bone. Condylus medialis larger andsome higher than condylus lateralis. Fossa inter-condylaris which separates them is wide and deep.Fossa muscularis cranialis and F. m. caudalis deepand parallel to the condyles. Longitudinal crests sur-rounding trochlea patellaris high and rounded intheir uppermost part.
Patella (Pl. XVIII, Fig. 4). Pear shaped. Concaveanteriorly. In the middle it is convex like a hemi-sphere surrounded by a deep and wide ring. Flat-
tened posteriorly. The larger part of the bone wascovered by a cartilage with the shape of the thigh-bone’s surface it articulated with.
From the lower hindlimbs, only distal part of the lefttibia and fragments of the proximal parts of the twofibulae were found.
Tibia (Pl. XXII, Fig. 2 and 2a). Although the bonewas restored to its full size, we’ll discuss only its dis-tal part which is original. Down of the diaphysis thebone becomes triangular. It gradually widens, form-ing at the distal end the cochlea tibiae. Maleolusmedialis strongly developed, incisura fibularis deepand wide.
Fibula. The original fragments used for the restora-tion are from the proximal part. This is a long andnarrow bone. Almost flat in its proximal part. Thickat the beginning, articular surfaces unclear.
Comparison. There are no differences of taxonomi-cal value in the hipbones and the bones of the lowerleg.
37
Tarsus (Pl. XXIII, Fig. 1 and 1a; Fig. 11 A and B)Consists of seven bones. Two in the proximal row:calcaneus and astragalus; five in the distal: cuboid,three cuneiforms and, between them, navicular. Onlythe calcaneus was found. The rest were restored af-ter D. gigantissimum Stefanescu. The toes were re-stored, too.
Comparison. There are no differences in the struc-ture of the tarsals in the various deinothere species.There are such however in their digits. In D. thra-ceiensis sp. n. the size of the first and second pha-lanxes gradually decreases. In D. gigantissimum Ste-fanescu all digits have large first phalanxes, the sec-ond are much smaller and the third – even more.There is a big difference in their size. D. bavaricumH. v. Meyer and D. giganteum demonstrate a muchmore gradual transition from the first to the last pha-
Table 22Tarsals – tarsus sin et dext
Deinotherium traceiensis sp.n.
Astragalus Calcaneus Naviculare Cuneiforme mediale
Cuneiforme intermedium
Cuneiforme laterale № Dimensions, mm
sin dext sin dext sin dext sin dext sin dext sin dext 1. Diameter of the corpus 313.0 310.0 2. Proximal height 90.0 90.0
3. Average thickness of the corpus 130.0 130.0 190.0 200.0
4. Proximal width 150.0 150.0 5. Total width 245.0 250.0 6. Anterior length of the bone 170.0 170.0 222.0 220.0 114.0 113.0 118.0 117.0 7. Posterior length of the bone 180.0 180.0 8. Total length of the bone 350.0 350.0 9. Diameter of the bone 335.0 337.0
10. Thickness of the bone 78.0 80.0 11. Height of the bone 81.0 80.0 79.0 78.0 82.0 80.0
Fig. 11. Pes of Deinotherium thraciensis sp. n. A – left pes; B – right pes
lanx. In D. thraceiensis sp. n. the last phalanx - thehoof – in the hindlimbs is large, as with the fore-limbs. In the compared species they are much small-er. They are anteriorly situated like nails and touchthe ground only with the anterior surface, or with asmall part of the lower surface.
It can be seen from the description that D. thraceiensissp. n. has numerous characters distinguishing it fromall the other species. Here are some of them:1. Skull – short, high and, compared to the size ofthe body – very small.2. External nares large and deep, with a peculiarshape.3. Nasal bones short, narrow and high, frontally fused.4. Forehead high and wide.5. Occipital bones high and wide, at an angle of 80°to the forehead.
38
Table
23
Met
ata
rsals –
met
ata
rsus
sin e
t dex
t
Dei
noth
eriu
m th
race
iens
is s
p.n.
M
t-I
Mt-
II
Mt-
III
Mt-
IV
Mt-
V
№
Dim
ensi
ons,
mm
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
1.
Len
gth
212.
0 21
2.0
220.
0 22
0.0
225.
0 22
5.0
238.
0 24
0.0
222.
0 22
2.0
2.
Wid
th a
t pro
xim
al e
nd
137.
0 13
8.0
114.
0 11
4.0
120.
0 12
5.0
120.
0 12
0.0
146.
0 14
6.0
3.
Wid
th a
t dis
tal e
nd
120.
0 12
0.0
115.
0 11
5.0
115.
0 11
5.0
114.
0 11
5.0
116.
0 11
6.0
4.
Wid
th a
t dis
tal s
inew
pro
cess
es
135.
0 13
6.0
112.
0 11
2.0
102.
0 10
3.0
112.
0 11
2.0
117.
0 11
7.0
5.
Len
gth
of s
mal
l fac
et
131.
0 13
0.0
- -
- -
- -
- -
6.
Len
gth
of la
rge
face
t 12
6.0
126.
0 -
- -
- -
- -
-
Table
24
Bones
of
the
dig
its
– h
indlim
b –
Phala
ngi sin e
t dex
t
Dei
noth
eriu
m th
race
iens
is s
p. n
. 1-
st d
igit
2-
nd d
igit
3-rd
dig
it
1 2
3 1
2 3
1 2
3 №
D
imen
sion
s, m
m
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
1.
L
engt
h 13
6 13
6 10
7 10
7 13
0 13
0 16
0 16
0 10
0 10
0 14
2 14
2 18
0 18
0 12
5 12
5 16
5 16
7 2.
W
idth
at p
roxi
mal
end
12
12
2 10
5 10
5 94
.5
94
112
112
104
105
98
98
117
116
110
111
107
107
3.
Wid
th a
t dis
tal e
nd
100
100
92
92
- -
101
101.
5 87
87
.5
- -
104
104
95
94
- -
Table
24 A
Bones
of
the
dig
its
– h
indlim
b –
Phala
ngi sin e
t dex
t
Dei
noth
eriu
m th
race
iens
is s
p. n
. 4-
th d
igit
5-th
dig
it 1
2 3
1 2
3 №
D
imen
sion
s, m
m
sin
dext
si
n de
xt
sin
dext
si
n de
xt
sin
dext
si
n de
xt
1.
Len
gth
169
170
105
105
125
125
146
147
104
104
103
103
2.
Wid
th a
t pro
xim
al e
nd
117
117
110
110
87.5
89
11
0 11
1.5
103
104
85
86
3.
Wid
th a
t dis
tal e
nd
106
108
93
94
- -
105
103
92
92
- -
39
6. Eye socket differentiated and large.7. Occipital condyles low.8. P3 and P4 differently structured.9. Ramus horizontalis strongly protruding backwardsat angulare.10. Processus articularis and p. coronoideus high.11. Incisura mandibulae deep.12. Tusks differently structured.13. P3 differently structured.14. Curve of the symphysis much more regular.15. Spinous processes on the cervicals shorter andweaker; first cervicals lack processes.16. Spinal canal widening at the lumbar region.17. Spinous process on second thoracic vertebrastrongly developed.18. 14 ribs.19. Manus and pes high and spread.20. Hoofs larger and entirely ‘lying’ on the ground.
On the basis of these characters, we believe thatthe specific status of D. thraceiensis n. sp. is entirelyproved. To this conclusion we shall add some shortnotes on its anatomy.
We are acquainted with the reconstructions byProf. Abel of D. bavaricum H. v. Meyer, based on the
skeleton in Vienna, and another of D. giganteumKaup. We know also the paintings by Augusta andBurian of D. giganteum Kaup, and the painting ofD. gigantissimum Stefanescu exhibited in the Bucha-rest Museum in front of the skeleton itself. It is madefollowing the first two authors. Everywhere the skullwith the mandible takes about 1/3 of the whole skel-eton. The neck is thick, merging with the vertebralcolumn because, according to them, the last cervi-cals have high spinous processes. Frontally, the chestis narrow, and the body – solid, with high legs. Thelatter are narrower near the fingers because of thesmall hoofs.
In D. thraceiensis sp. n. the skull and the mandi-ble are much less than 1/3 of the whole skeleton. Theneck is short and only the last cervicals have spinousprocesses. They are very high on the first thoracicvertebrae, widening at the tip. This permits us to sup-pose it had a high hump like Elephas.
On the other hand, as far as the first rib is wideand solid, its chest was probably broad. The digitswere more spread.
For the time being, we cannot comment on theproblems of the phylogeny of genus Deinotherium.They could become the subject of a future work.
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