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''Deinotherium'' was a large elephant-like proboscidean that appeared in the Middle
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent ...
and survived until the Early
Pleistocene The Pleistocene ( , often referred to as the '' Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed i ...
. Although superficially resembling modern elephants, they had notably more flexible necks, limbs adapted to a more
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often u ...
lifestyle as well as
tusks Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share c ...
that curved downwards and back. In addition, their tusks didn't emerge from the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The ...
as in elephants but from the
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
. ''Deinotherium'' was a widespread genus, ranging from
East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historica ...
to the south to Europe and east to the
Indian Subcontinent The Indian subcontinent is a physiographical region in Southern Asia. It is situated on the Indian Plate, projecting southwards into the Indian Ocean from the Himalayas. Geopolitically, it includes the countries of Bangladesh, Bhutan, Ind ...
. They were browsing animals with a diet mainly consisting of leaves, and they most likely died out as forested areas were gradually replaced by open grassland during the latter half of the
Neogene The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
.


History and naming

''Deinotherium'' has a long history, possibly dating back as early as the 17th century when a French surgeon named Matsorier found the bones of large animals in an area known as the "field of giants" near
Lyon Lyon,, ; Occitan: ''Lion'', hist. ''Lionés'' also spelled in English as Lyons, is the third-largest city and second-largest metropolitan area of France. It is located at the confluence of the rivers Rhône and Saône, to the northwest of th ...
. Matsorier is said to have exhibited these bones across France and Germany as the supposed bones of a French monarch, until he was exposed and the bones were handed over to the French National Museum of Natural History. In 1775 researchers recognized the bones as belonging to an animal "similar to a mammoth" and during the late 18th/early 19th century
George Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in nat ...
hypothesized that they actually belonged to a large
tapir Tapirs ( ) are large, herbivorous mammals belonging to the family Tapiridae. They are similar in shape to a pig, with a short, prehensile nose trunk. Tapirs inhabit jungle and forest regions of South and Central America, with one species inhabi ...
with upwards curving tusks which he named ''Tapir gigantesque''. Another early hypothesis suggested that ''Deinotherium'' was a
Sirenian The Sirenia (), commonly referred to as sea-cows or sirenians, are an order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The Sirenia currently comprise two distinct f ...
that used its tusks to anchor itself to the sea floor while sleeping. The genus ''Deinotherium'' was coined in 1829 by
Johann Jakob von Kaup Johann Jakob von Kaup (10 April 1803 – 4 July 1873) was a German naturalist. A proponent of natural philosophy, he believed in an innate mathematical order in nature and he attempted biological classifications based on the Quinarian system. Kaup ...
to describe a fossil skull and mandible discovered in Germany. The type specimen, ''D. giganteum'' was at the time thought to be an evolutionary link between sloths and mastodonts. Further remains were discovered and named, including many that would later come to be considered part of the genus ''
Prodeinotherium ''Prodeinotherium'' is an extinct representative of the family Deinotheriidae that lived in Africa, Europe, and Asia in the early and middle Miocene. ''Prodeinotherium'', meaning "before terrible beast", was first named in 1930, but soon after, t ...
''. These additional remains also helped solidify ''Deinotherium's'' position within
Proboscidea The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Fr ...
and finds in India described as ''D. indicum'' extended the range of the genus outside of Europe. Fossils of an exceptionally large specimen found in Manzati, Romania between the late 19th and early 20th century were described as ''D. gigantissimum''. In Bulgaria ''Deinotherium'' remains have been found from 1897 onward, with one particular fossil of an almost complete animal found in 1965. These remains were officially described in 2006 as ''D. thraceiensis'', making it the most recently named species. The name ''Deinotherium'' is derived from the Ancient Greek , ' meaning "terrible" and , ' meaning "beast"). Some authors have on occasion referred to ''Deinotherium'' as ''Dinotherium'', following latinization of the first element of the name. Although pronunciation remains unchanged, ''Deinotherium'' remains the valid spelling as it was coined first.


Description

''Deinotherium'' was a large bodied proboscidean displaying continued growth between species. Two adults of ''D. giganteum'' are around tall and weighed . This is similar to adult males of ''D. proavum'', one of which weighed and was tall. However, both these species are smaller than a 45-year-old male of ''D. "thraceiensis"'', at tall and . The general anatomy of ''Deinotherium'' is similar to that of modern elephants with pillar-like limbs, although proportionally longer and more slender than those of other proboscideans. The bones of the toes are longer and less robust than in elephants and the neck likewise differs notably in that it is relatively longer, however still quite short compared to other modern browsers like
giraffes The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, ''Giraffa camelopardalis ...
. The permanent tooth formula of ''D. giganteum'' was \frac (
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
\frac), with vertical cheek tooth replacement. Two sets of bilophodont and trilophodont teeth were present. The molars and rear premolars were vertical shearing teeth, and suggest that deinotheres became an independent evolutionary branch very early on; the other premolars were used for crushing. The
cranium The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, ...
was short, low, and flattened on the top, in contrast to more advanced proboscideans, which have a higher and more domed forehead, with very large, elevated
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
s. The largest skulls of ''Deinotherium'' reached a length of . The nasal opening was retracted and large, indicating a large trunk. The
rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit * Rostrum (anatomy), a beak, or anatomical structure resembling a beak, as in the mouthparts of many sucking insects * Rostrum (ship), a form of bow on naval ships * R ...
was long and the rostral fossa broad. The mandibular symphyses (the lower jaw-bone) were very long and curved downward, which, with the backward-curved tusks, is a distinguishing feature of the group. These tusks are without doubt the most immediately visible feature of ''Deinotherium''. Unlike in modern proboscideans, which possess tusks that grow from the upper
incisors Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
, the tusks of ''Deinotherium'' grow from the lower incisors, with upper incisors and upper and lower canines missing entirely. The curvature is initially formed by the mandible itself, with the teeth themselves erupting at only the halfway point of the curve. The degree to which the tusks follow the direction predetermined by the mandible varies between specimens, with some tusks following the curve and pointing backwards, forming an almost semicircular shape, while in other specimens the tusks continue down almost vertically. The tusks have a roughly oval cross-section and could reach a length of . The shape of these tusks has led some researchers to coin the name "hoe tusker" in reference to ''Deinotherium''. Although the presence of a elephant-like proboscis or trunk in ''Deinotherium'' is evident thanks to the size and shape of the external nares, the exact shape and size of this trunk is a matter that's long been debated. Historic depictions commonly portray it as very elephantine with a long trunk and tusks breaking through the skin below an elephantine lower lip. In the early 2000s Markov and colleagues published papers on the facial soft tissue of ''Deinotherium'' contesting these ideas, instead suggesting an alternative soft tissue reconstruction. In the first of these publications the authors argue that, due to the origin of these animal's tusks, the lower lip should be situated beneath them as they evolved their classic downturned appearance. They further suggest that, while a trunk would be present, it would likely not resemble that of modern elephants and instead be more robust and muscular, which they reason is evidenced by the lack of a proper insertion surface. Although later research concurs that the trunk or proboscis of ''Deinotherium'' was likely notably different from those of modern proboscideans, the idea of a short tapir-like trunk is questioned. In particular, it is pointed out that the tall stature and still relatively short neck of ''Deinotherium'' would render it very difficult for the animal to drink without assuming a more complex posture. Thus it is suggested that the trunk must have been at least long enough for the animal to effectively drink.


Species

Throughout the long history of deinotheriid research, 31 species have been described and assigned to the family, many on the basis of poorly sampled material, especially teeth of varying size. The amount of species recognized by authors differs depending on researchers, but the three species most commonly considered valid are listed below. ;''D. bozasi'' :Known from
East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historica ...
, ''Deinotherium bozasi'' was the last known species of ''Deinotherium'', surviving in the Kanjera Formation,
Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi ...
, until the early Pleistocene roughly 1 million years ago. It is characterized by a narrower rostral trough, a smaller but higher nasal aperture, a higher and narrower cranium, and a shorter mandibular symphysis than the other two species. In a 2013 publication
Martin Pickford Martin Pickford was lecturer in the Chair of Paleoanthropology and Prehistory at the Collège de France and honorary affiliate at the Département Histoire de la Terre in the Muséum national d'Histoire. In 2001, Martin Pickford together wi ...
notes that ''D. bozasi'' has mandibles anatomically similar to those of ''D. proavum'', however most specimens are smaller than those of the European species. To explain this, two hypothesis are suggested, one that they share a common ancestor and the other that ''D. bozasi'' may be an example of Allen's rule, which states that animals at lower latitudes are typically smaller than relatives at higher latitudes. However Markov and colleagues suggest that the similar mandibular anatomy may be a case of parallel evolution between late European species and ''D. bozasi'' in response to aridification and an increased need for effective
mastication Chewing or mastication is the process by which food is crushed and ground by teeth. It is the first step of digestion, and it increases the surface area of foods to allow a more efficient break down by enzymes. During the mastication process, ...
. ;''D. giganteum'' :The type species ''D. giganteum'' was found in Europe from the Middle Miocene to Early Pliocene. However, the exact extinction of ''D. giganteum'' in Europe is unknown. The last known occurrences in Central and Western Europe appear to be in MN13 (
Messinian The Messinian is in the geologic timescale the last age or uppermost stage of the Miocene. It spans the time between 7.246 ± 0.005 Ma and 5.333 ± 0.005 Ma (million years ago). It follows the Tortonian and is followed by the Zanclean, the fir ...
to
Zanclean The Zanclean is the lowest stage or earliest age on the geologic time scale of the Pliocene. It spans the time between 5.332 ± 0.005 Ma (million years ago) and 3.6 ± 0.005 Ma. It is preceded by the Messinian Age of the Miocene Epoch, an ...
), while material from Russia might extend the range of the species to MN15 (
Ruscinian The Ruscian age is a period of geologic time (5.3–3.4 Ma) within the Pliocene used more specifically with European Land Mammal Ages. It precedes the Villanyian age and follows the Turolian age. The Ruscian overlaps the early Piacenzian and Zanc ...
). Fossils of ''D. giganteum'' have also been found on the island of
Crete Crete ( el, Κρήτη, translit=, Modern: , Ancient: ) is the largest and most populous of the Greek islands, the 88th largest island in the world and the fifth largest island in the Mediterranean Sea, after Sicily, Sardinia, Cypru ...
in the upper Miocene Faneroméni Formation, during a time when the island was still connected to the mainland. ;''D. indicum'' :The Asian species, ''D. indicum'' is distinguished by a more robust dentition as well as p4-m3 intravalley tubercles and found across the Indian subcontinent (India and Pakistan) during the Middle and Late Miocene. It disappeared from the fossil record about 7 million years ago (Late Miocene). Although it is generally regarded as valid, some researchers argue that it is synonymous with ''D. proavum'' and that the later name would take precedence. Pickford, for instance, argues that fossils from
Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Tu ...
create a geographic link between European populations and the Indian specimens, concluding that they may be one single wide ranging species. One hypothesis opposing this three-species model suggests that, rather than being a single consistent species lasting throughout the Miocene, ''D. giganteum'' actually represents multiple chronospecies, with the type species only applying to the intermediate form. Other species that have been described include: ;''D. levius'' (Jourdan, 1861) :''D. levius'' is a European species of ''Deinotherium'' recovered from sediments dating to the late Astaracian to Aragonian. While it is considered a synonym of ''D. giganteum'' by some researchers, others propose that it is a stratigraphically distinct chronospecies and the earliest of European ''Deinotherium''. In accordance to this hypothesis, ''D. levius'' would eventually give rise to ''D. giganteum'' by the
Vallesian The Vallesian age is a period of geologic time (11.6–9.0 Ma) within the Miocene used more specifically with European Land Mammal Ages. It precedes the Turolian age and follows the Astaracian age. The so-called Vallesian Crisis resulted in the e ...
stage of the Miocene, after which the two species continued to coexist until the formers extinction. ;''D. proavum'' (Eichwald, 1831) :''D. proavum'' is a large bodied species of ''Deinotherium'' that may be a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linnae ...
of ''Deinotherium giganteum''. Other research meanwhile proposes that it, alongside ''D. giganteum'' and ''D. levius'', is part of a single anagenetic lineage of ''Deinotherium'' species. For this hypothesis it has been suggested that it evolved from ''D. giganteum'' during the late Vallesion to Turolian, with early members of the species still being similar in size to its ancestor before surpassing it later during its range. However, the assignment of specimens to ''D. proavum'' is largely based on stratigraphy and size, making the differentiation between species difficult, especially with some research suggesting that the two species continued to coexist. ;''D. gigantissimus'' :''D. gigantissimus'' from Romania is typically considered to be a larger specimen belonging either to ''D. giganteum'' or ''D. proavum'' (depending on how many species are recognized by the respective author). The situation is similar in ''D. thraceiensis'' from Bulgaria, another notably large deinothere, described in 2006 but usually lumped into other European species by subsequent publications. The state of Asian species is especially complex, with a multitude of specimens being described from poor remains. These include ''D. sindiense'' (Lydekker, 1880), ''D. orlovii'' (Sahni and Tripathi, 1957), ''D. naricum'' (Pilgrim, 1908), and ''D. anguistidens'' (Koch 1845), all of which are generally considered dubious by publications of the 21st century. Only one other species from Africa was described, ''D. hopwoodi'' (Osborn, 1936), based on teeth from the Omo Basin in
Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the Er ...
. However his research was published posthumously and was predated by ''D. bozasi'', described two years prior. Another matter that complicates the amount of ''Deinotherium'' species recognized by science is the state of the genus ''Prodeinotherium''. One prevailing theory is that ''Prodeinotherium'' is a distinct genus ancestral to the larger ''Deinotherium'' species. Other researchers, however, argue that the anatomical differences, the difference in size in particular, are not enough to properly distinguish the two, which would subsequently render species of ''Prodeinotherium'' as ''Deinotherium'' instead. This would create the combinations ''D. bavaricum'', ''D. cuvieri'' (both European), ''D. hobleyi'' (Africa), ''P. pentapotamiae'', and possibly ''D. sinense'' (Asia). ''Deinotherium'' was a widespread genus, found across vast areas of
East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historica ...
, Europe, the
Arabian Peninsula The Arabian Peninsula, (; ar, شِبْهُ الْجَزِيرَةِ الْعَرَبِيَّة, , "Arabian Peninsula" or , , "Island of the Arabs") or Arabia, is a peninsula of Western Asia, situated northeast of Africa on the Arabian Plate ...
and
South South is one of the cardinal directions or compass points. The direction is the opposite of north and is perpendicular to both east and west. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Proto-Germanic ''*sunþaz' ...
to
East Asia East Asia is the eastern region of Asia, which is defined in both geographical and ethno-cultural terms. The modern states of East Asia include China, Japan, Mongolia, North Korea, South Korea, and Taiwan. China, North Korea, South Kore ...
. In Europe fossils are especially common in the southeast, with up to half of known specimens in the region originating in
Bulgaria Bulgaria (; bg, България, Bǎlgariya), officially the Republic of Bulgaria,, ) is a country in Southeast Europe. It is situated on the eastern flank of the Balkans, and is bordered by Romania to the north, Serbia and North Macedon ...
. Especially significant specimens include those found in Ezerovo, Plovdiv Province (type specimen of ''D. thraceiensis'') and near Varna.
Romania Romania ( ; ro, România ) is a country located at the crossroads of Central, Eastern, and Southeastern Europe. It borders Bulgaria to the south, Ukraine to the north, Hungary to the west, Serbia to the southwest, Moldova to the east, and ...
likewise yielded significant remains, with one notably large specimen being found by Grigoriu Ștefănescu near Mânzați (type specimen of ''D. gigantissimum''). The fossils of the two now invalid species are displayed at the National Museum of Natural History, Bulgaria and the Grigore Antipa National Museum of Natural History respectively. Multiple specimens have also been found in
Greece Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders with ...
and even on the island of
Crete Crete ( el, Κρήτη, translit=, Modern: , Ancient: ) is the largest and most populous of the Greek islands, the 88th largest island in the world and the fifth largest island in the Mediterranean Sea, after Sicily, Sardinia, Cypru ...
, indicating that the large animal had traveled there over a potential landbridge. Towards the east ''Deinotherium'' is known from finds in Russia (
Rostov-on-Don Rostov-on-Don ( rus, Ростов-на-Дону, r=Rostov-na-Donu, p=rɐˈstof nə dɐˈnu) is a port city and the administrative centre of Rostov Oblast and the Southern Federal District of Russia. It lies in the southeastern part of the East ...
),
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
, and
Turkey Turkey ( tr, Türkiye ), officially the Republic of Türkiye ( tr, Türkiye Cumhuriyeti, links=no ), is a transcontinental country located mainly on the Anatolian Peninsula in Western Asia, with a small portion on the Balkan Peninsula in ...
. The range of ''Deinotherium'' furthermore extends over the
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabia (including the Arabian Peninsula and Bahrain), Asia Minor (Asian part of Turkey except Hatay Province), East Thrace (Europ ...
, with the holotype of ''D. indicum'' being found on the island of
Perim Perim ( ar, بريم 'Barīm'', also called Mayyun in Arabic, is a volcanic island in the Strait of Mandeb at the south entrance into the Red Sea, off the south-west coast of Yemen and belonging to Yemen. It administratively belongs to Dhub ...
(
Yemen Yemen (; ar, ٱلْيَمَن, al-Yaman), officially the Republic of Yemen,, ) is a country in Western Asia. It is situated on the southern end of the Arabian Peninsula, and borders Saudi Arabia to the north and Oman to the northeast and s ...
) in the
Red Sea The Red Sea ( ar, البحر الأحمر - بحر القلزم, translit=Modern: al-Baḥr al-ʾAḥmar, Medieval: Baḥr al-Qulzum; or ; Coptic: ⲫⲓⲟⲙ ⲛ̀ϩⲁϩ ''Phiom Enhah'' or ⲫⲓⲟⲙ ⲛ̀ϣⲁⲣⲓ ''Phiom ǹšari''; ...
. Fossils are also known from
Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Tu ...
and multiple localities on the
Indian Subcontinent The Indian subcontinent is a physiographical region in Southern Asia. It is situated on the Indian Plate, projecting southwards into the Indian Ocean from the Himalayas. Geopolitically, it includes the countries of Bangladesh, Bhutan, Ind ...
such as the
Siwalik Hills The Sivalik Hills, also known as the Shivalik Hills and Churia Hills, are a mountain range of the outer Himalayas that stretches over about from the Indus River eastwards close to the Brahmaputra River, spanning the northern parts of the Indian ...
. The easternmost occurrence of the genus appears to be in the province of
Gansu Gansu (, ; alternately romanized as Kansu) is a province in Northwest China. Its capital and largest city is Lanzhou, in the southeast part of the province. The seventh-largest administrative district by area at , Gansu lies between the Tibe ...
,
Northwest China Northwest China () is a statistical region of China which includes the autonomous regions of Xinjiang and Ningxia and the provinces of Shaanxi, Gansu and Qinghai. It has an area of 3,107,900 km2. The region is characterized by a (semi-)arid ...
. The western range of ''Deinotherium'' spans most of West and Central Europe including
Hungary Hungary ( hu, Magyarország ) is a landlocked country in Central Europe. Spanning of the Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croa ...
, the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The Cz ...
(Františkovy Lázně),
Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populou ...
(Gratkorn Locality),
Switzerland ). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...
(
Jura Mountains The Jura Mountains ( , , , ; french: Massif du Jura; german: Juragebirge; it, Massiccio del Giura, rm, Montagnas da Jura) are a sub-alpine mountain range a short distance north of the Western Alps and mainly demarcate a long part of the Fre ...
), France ("Field of Giants"),
Portugal Portugal, officially the Portuguese Republic ( pt, República Portuguesa, links=yes ), is a country whose mainland is located on the Iberian Peninsula of Southwestern Europe, and whose territory also includes the Atlantic archipelagos of the ...
,
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = '' Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , ...
and Germany. Some of the earliest and most significant finds in Germany have been made in the Dinotheriensande ( Eppelsheim Formation) of the
Mainz Basin The Mainz Basin (german: Mainzer Becken) or Rhine-Main BasinElkins, T H (1972). ''Germany'' (3rd ed.). London: Chatto & Windus, 1972, p. 218. . is the name given to a Cenozoic marine basin that covered the area of the present-day region of Rhenish ...
, named for their great abundance of deinothere remains. The holotype specimen of ''Deintherium'', described by Kaup in the early 1800s, stems from this part of Europe. Outside of
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago ...
''Deinotherium'' is found in
East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historica ...
, with specimens known from the
Olduvai Gorge The Olduvai Gorge or Oldupai Gorge in Tanzania is one of the most important paleoanthropological localities in the world; the many sites exposed by the gorge have proven invaluable in furthering understanding of early human evolution. A steep- ...
in
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands an ...
, the Omo Basin and
Middle Awash The Middle Awash is a paleoanthropological research area in the Afar Region along the Awash River in Ethiopia's Afar Depression. It is a unique natural laboratory for the study of human origins and evolution and a number of fossils of the earliest ...
of
Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the Er ...
, and multiple localities in
Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi ...
. ''Deinotherium bozasi'' remains have been found in the Kenyan Chemoigut Beds around Lake Baringo, as well as the Kubi Algi Formation and Koobi Fora Formation in East Rudolf. An additional tooth is known from
Sahabi The Companions of the Prophet ( ar, اَلصَّحَابَةُ; ''aṣ-ṣaḥāba'' meaning "the companions", from the verb meaning "accompany", "keep company with", "associate with") were the disciples and followers of Muhammad who saw or m ...
,
Libya Libya (; ar, ليبيا, Lībiyā), officially the State of Libya ( ar, دولة ليبيا, Dawlat Lībiyā), is a country in the Maghreb region in North Africa. It is bordered by the Mediterranean Sea to the north, Egypt to the east, Suda ...
and it's possible that both ''Deinotherium'' and ''Prodeinotherium'' coexisted in the Kenyan Ngorora Formation.


Evolution

The origin of Deinotheres can be found in the Oligocene of Africa with the relatively small bodied '' Chilgatherium''. Initially restricted to Africa, the continued northward movement of the African Plate eventually caused the Proboscidean Datum Event, during which proboscideans diversified and spread into Eurasia, among them the ancestral ''Prodeinotherium'', thought to be the direct predecessor of the larger ''Deinotherium''. Generally, ''Deinotherium'' displays relatively little change in morphology throughout its evolution, but a steady increase in body size from 2 meters shoulder height in ''Prodeinotherium'' to up to 4 meters in later ''Deinotherium'' species and a mass far exceeding even large African elephants. The reasons for this rapid increase in body size is interpreted to have had multiple factors influencing it. On the one hand, increased size is an effective predator deterrent, especially during the Miocene when carnivorans had reached a great diversity including hyaenodonts, amphicyonids and large cats. Secondly, continued aridification during the Miocene increasingly split up woodlands, with greater distances of open landscape stretching between the food sources of browsers such as ''Deinotherium''. This also accounts for the morphological adaptations seen in the limbs of ''Deinotherium'', better suited for long distance travel. Furthermore, the appearance of ''Deinotherium'' coincided with falling temperatures during the middle Miocene. According to
Bergmann's rule Bergmann's rule is an ecogeographical rule that states that within a broadly distributed taxonomic clade, populations and species of larger size are found in colder environments, while populations and species of smaller size are found in warmer ...
, these circumstances favor increased body mass for maintaining heat in cold temperatures. Despite the many key adaptations deinotheres developed for effective foraging, the continued aridification that progressed throughout the Miocene eventually led to the extinction of the group, which failed to survive without readily available food sources matching their diet. Populations in Western Europe were the first to disappear, followed later by those in Eastern Europe. While European lineages of ''Deinotherium'' had gone extinct with the onset of the Pliocene, the genus managed to survive notably longer in its southern range in Africa. The last known ''Deinotherium'' remains, assigned to ''D. bozasi'', were found in sediments dating to the Pleistocene, approximately 1 million years ago.


Paleoecology

Several key adaptations suggest that ''Deinotherium'' was a folivorous, browsing proboscidean that preferred open woodland habitats and fed on the leaves of the tree canopy. In Asia ''D. indicum'' has been associated with wet and warm, low-energy woodland and in
Portugal Portugal, officially the Portuguese Republic ( pt, República Portuguesa, links=yes ), is a country whose mainland is located on the Iberian Peninsula of Southwestern Europe, and whose territory also includes the Atlantic archipelagos of the ...
deinotheriid remains were found in regions corresponding with moist, tropical to subtropical woodland conditions likened to modern
Senegal Senegal,; Wolof: ''Senegaal''; Pulaar: 𞤅𞤫𞤲𞤫𞤺𞤢𞥄𞤤𞤭 (Senegaali); Arabic: السنغال ''As-Sinighal'') officially the Republic of Senegal,; Wolof: ''Réewum Senegaal''; Pulaar : 𞤈𞤫𞤲𞤣𞤢𞥄𞤲𞤣� ...
. A browsing lifestyle is supported by the inclination of the occiput that gives ''Deinotherium'' a slightly more raised head posture, and their teeth, which strongly resemble those of modern tapirs, animals that predominantly feed on fruits, flowers, bark and leaves. Their limbs show some notable differences to ''Prodeinotherium'', allowing for a more agile mode of locomotion and allowing for easier travel across open landscapes in the search of food, which coincides with the widespread breakup of forests and expansion of grasslands during the time ''Deinotherium'' lived in Europe. Fossil finds from the Austrian Gratkorn locality and the Mainz Basin in Germany indicate that ''Deinotherium'' was not a permanent resident in some areas it inhabited. In Austria it has been suggested that they traversed areas on a regular basis, while in Germany there is evidence for the animals range shifting with changing climatic conditions, present during subtropical climate conditions and absent in sub-boreal conditions. One of the most enigmatic features of ''Deinotherium'' are their downturned tusks and their function. Research conducted on ''Deinotherium'' suggests that these tusks were likely not used for digging, nor are they sexually dimorphic, leaving use in feeding as their most likely function. These tusks exhibit patterns of wear, in particular on their medial and caudal sides. In a 2001 paper Markov and colleagues argue that ''Deinotherium'' could have used its tusks to remove branches that would have gotten in the way of feeding, while using the proboscis to transport leaf material into its mouth. From there ''Deinotherium'' would have used a powerful tongue (inferred based on a notable trough at the front of the symphysis) to further manipulate its food. Different tusk anatomy in young individuals would suggest altered feeding strategies in juveniles.


Cultural significance


Possible influence on Greek myths

''Deinotherium'' is among the fossil proboscideans represented in the Miocene-age deposits on the Greek island of Samos, alongside ''
Konobelodon ''Konobelodon'' is an extinct genus of amebelodont from southern Europe, China, and North America. Taxonomy ''Konobelodon'' was originally coined as a subgenus of '' Amebelodon'', and was subsequently elevated to full generic rank in a 2014 re ...
'' and ''
Choerolophodon ''Choerolophodon'' is an extinct genus of proboscidean that lived during the Miocene of Eurasia and Africa. Fossils of ''Choerolophodon'' have been found in Africa, Southeast Europe, Turkey, Iraq, Iran, the Indian subcontinent, and China. ''Choer ...
.''
Adrienne Mayor Adrienne Mayor (born 1946) is a historian of ancient science and a classical folklorist. Mayor specializes in ancient history and the study of " folk science", or how pre-scientific cultures interpreted data about the natural world, and how these ...
and
Nikos Solounias Nikos ( el, Νίκος, ''Níkos'') is a Greek given name. It originates from Greek ''Nikolaos'', which means "victory of the people".Liddell & Scott, Abridged Greek Lexicon Although used as a proper first name, Nikos is also a popular nickname of t ...
have speculated that these taxa may have influenced local legends of the island's deep history, serving as inspiration for gigantic mythical monsters called
Neades The Neades ( grc, Νηάς, Νηάδες) were legendary creatures of gigantic size said to inhabit the Greek island of Samos. Their voices were said to be capable of causing earthquakes, and ancient sources report that their bones were put on disp ...
, creatures whose voices were believed to cause earthquakes. Mayor and Solounias base their speculation on the fact that the fossils on Samos are found near a major
fault zone In geology, a fault is a planar fracture or discontinuity in a volume of rock across which there has been significant displacement as a result of rock-mass movements. Large faults within Earth's crust result from the action of plate tectoni ...
, suggesting that ancient Greeks may have interpreted the presence of their skeletal remains as being associated with past seismic activity in the region. Ancient sources attest that the bones of these creatures were put on display and that their stories inspired local expressions such as "They shout louder than the Neades!" Mayor and Solounias also suggest that these fossils may have inspired another story of the god
Dionysus In ancient Greek religion and myth, Dionysus (; grc, Διόνυσος ) is the god of the grape-harvest, winemaking, orchards and fruit, vegetation, fertility, insanity, ritual madness, religious ecstasy, festivity, and theatre. The Roma ...
waging war with the
Amazons In Greek mythology, the Amazons ( Ancient Greek: Ἀμαζόνες ''Amazónes'', singular Ἀμαζών ''Amazōn'', via Latin ''Amāzon, -ŏnis'') are portrayed in a number of ancient epic poems and legends, such as the Labours of Hercules ...
on Samos, drawing similar comparisons between the island's geology and a description by
Plutarch Plutarch (; grc-gre, Πλούταρχος, ''Ploútarchos''; ; – after AD 119) was a Greek Middle Platonist philosopher, historian, biographer, essayist, and priest at the Temple of Apollo in Delphi. He is known primarily for h ...
of this mythic fight. In this story, their skeletal remains are interpreted as belonging to the victims of this ancient war, who like the Neades were able to rend the earth when they cried out in death. They argue that ancient residents of Samos were aware of both the island's fossil record and its geology, and that these stories were early attempts to make sense of the two.


References


Further reading

* Carroll, R.L. (1988), ''Vertebrate Paleontology and Evolution'', WH Freeman & Co. * Colbert, E. H. (1969), ''Evolution of the Vertebrates'', John Wiley & Sons Inc (2nd ed.) * Harris, J.M. (1976) Evolution of feeding mechanisms in the family Deinotheriidae (Mammalia: Proboscidea). ''Zool. J. Linn. Soc.'' 56: 331-362 {{Authority control Deinotheriids Miocene proboscideans Pliocene proboscideans Pleistocene proboscideans Miocene genus first appearances Pleistocene genus extinctions Miocene mammals of Africa Miocene mammals of Europe Miocene mammals of Asia Pliocene mammals of Africa Pliocene mammals of Europe Pliocene mammals of Asia Pleistocene mammals of Asia Pleistocene mammals of Africa Pleistocene mammals of Europe Prehistoric elephants Prehistoric placental genera Taxa named by Johann Jakob Kaup Fossil taxa described in 1829