Chalicotherium
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''Chalicotherium'', from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
(''khálix''), meaning "gravel", and (''theríon''), meaning "beast", is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of extinct
odd-toed ungulate Perissodactyla (, ), or odd-toed ungulates, is an order of ungulates. The order includes about 17 living species divided into three families: Equidae (horses, asses, and zebras), Rhinocerotidae (rhinoceroses), and Tapiridae (tapirs). They t ...
s of the order
Perissodactyla Perissodactyla (, ), or odd-toed ungulates, is an order of ungulates. The order includes about 17 living species divided into three families: Equidae (horses, asses, and zebras), Rhinocerotidae (rhinoceroses), and Tapiridae (tapirs). They t ...
and
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Chalicotheriidae Chalicotheriidae (from Ancient Greek ''khálix'', "gravel", and ''theríon'', "beast") is an extinct family of herbivorous, odd-toed ungulate (perissodactyl) mammals that lived in North America, Eurasia, and Africa from the Middle Eocene to the ...
. The genus is known from
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
and
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
, from the
Middle Miocene The Middle Miocene is a sub-epoch of the Miocene epoch (geology), epoch made up of two Stage (stratigraphy), stages: the Langhian and Serravallian stages. The Middle Miocene is preceded by the Early Miocene. The sub-epoch lasted from 15.97 ± 0. ...
to
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
, 15.9~5.3 million years ago. This animal would look much like other chalicotheriid species: an odd-looking
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
with long
claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or Arthro ...
ed
forelimb A forelimb or front limb is one of the paired articulated appendages ( limbs) attached on the cranial (anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used inst ...
s and stouter weight-bearing
hindlimb A hindlimb or back limb is one of the paired articulated appendages ( limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-H ...
s. The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
, ''Chalicotherium goldfussi'', from
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
Europe, was described by
Johann Jakob 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 ...
in 1833. When the French naturalist
George Cuvier Jean Léopold Nicolas Frédéric, baron Cuvier (23 August 1769 – 13 May 1832), known as Georges Cuvier (; ), was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuv ...
first received a cleft claw from Eppelheim, Germany, he identified it as the toe bone of a gigantic
pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smutsia''. ''Manis'' comprises four species found in Asia, while ' ...
.


Description

''Chalicotherium'', like many members of Perissodactyla, was adapted to browsing, though the chalicotheres were uniquely adapted to do so among ungulates. Its arms were long and heavily clawed, allowing them to walk on their knuckles only. The arms were used to reach for the branches of large trees and bring them close to its long head to strip them clean of leaves. The
horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 mi ...
-like head itself shows adaptation to a diet of soft vegetation, since, as the animal reached
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans, it is related to both puberty and adulthood. ''Puberty'' is the biological process of sexual maturation, while ''adulthood'', the condition of being socially recognized ...
, the
incisor 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 ...
s and upper canines were shed, suggesting that its muscular lips and the resulting gum pads were enough to crop fodder which was then processed by squarish, low-crowned molars. Callosities on the
ischium The ischium (; : is ...
imply that these animals would sit on their haunches for extended periods of time, probably while feeding. Pad-supporting bony growth on the dorsal side of the manual
phalanges The phalanges (: phalanx ) are digit (anatomy), digital bones in the hands and foot, feet of most vertebrates. In primates, the Thumb, thumbs and Hallux, big toes have two phalanges while the other Digit (anatomy), digits have three phalanges. ...
is interpreted as evidence of
knuckle-walking Knuckle-walking is a form of quadrupedal walking in which the forelimbs hold the fingers in a partially flexed posture that allows body weight to press down on the ground through the knuckles. Gorillas and chimpanzees use this style of locomoti ...
, which would probably be useful to avoid wearing down the claws, preserving them for use as either as a forage-collecting
rake Rake may refer to: Common meanings * Rake (tool), a horticultural implement, a long-handled tool with tines * Rake (stock character), a man habituated to immoral conduct * Rake (poker), the commission taken by the house when hosting a poker game ...
or as formidable defensive weapons, or both. All of these characteristics show some
convergence Convergence may refer to: Arts and media Literature *''Convergence'' (book series), edited by Ruth Nanda Anshen *Convergence (comics), "Convergence" (comics), two separate story lines published by DC Comics: **A four-part crossover storyline that ...
with such other creatures as
ground sloth Ground sloths are a diverse group of extinct sloths in the mammalian superorder Xenarthra. They varied widely in size with the largest, belonging to genera '' Lestodon'', ''Eremotherium'' and ''Megatherium'', being around the size of elephants. ...
s,
great ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); '' Gorilla'' (the ...
s,
bear Bears are carnivoran mammals of the family (biology), family Ursidae (). They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats ...
s (especially
giant panda The giant panda (''Ailuropoda melanoleuca''), also known as the panda bear or simply panda, is a bear species endemic to China. It is characterised by its white animal coat, coat with black patches around the eyes, ears, legs and shoulders. ...
s), and a group of theropod dinosaurs known as
therizinosaur Therizinosaurs (; once called segnosaurs) are an extinct group of large herbivorous theropod dinosaurs whose fossils have been mainly discovered from Cretaceous deposits in Asia and North America. Potential fragmentary remains have also been foun ...
s.


Classification


Taxonomic history

The type specimens for ''Chalicotherium goldfussi'' were found in the Upper Miocene strata of the ''Dinotherien-sande'' beds near Eppelsheim, in the
Grand Duchy of Hesse The Grand Duchy of Hesse and by Rhine () was a grand duchy in western Germany that existed from 1806 to 1918. The grand duchy originally formed from the Landgraviate of Hesse-Darmstadt in 1806 as the Grand Duchy of Hesse (). It assumed the name ...
, Germany.
Johann Jakob 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 ...
, when describing this new animal in 1833, found the teeth to be pebble-like and named the creature accordingly. Later on, limbs found in strata located at
Sansan Sansan may refer to: *Sansan, Gers, commune in the Gers department, France *Sansan, Iran Sansiz () is a village in Dastjerdeh Rural District of Chavarzaq District in Tarom County, Zanjan province, Iran Iran, officially the Islam ...
in the department of
Gers Gers (; or , ) is a departments of France, department in the regions of France, region of Occitania (administrative region), Occitania, Southwestern France. Gers is bordered by the departments of Hautes-Pyrénées and Pyrénées-Atlantiques to ...
, Southwestern France, were first described as ''Macrotherium'' by
Édouard Lartet Édouard Lartet (15 April 180128 January 1871) was a French geologist and paleontologist, and a pioneer of Paleolithic archaeology. He is also known for coining the prehistoric taxon '' Amphicyon'', making it one of the earliest-described fossil ...
in 1837. Further study of these fossil remains and subsequent finds by Filhol warranted a referral of the material described as ''Macrotherium'' to ''Chalicotherium''. Referral history for each species is detailed in the species list below along with morphological and geographical data where available.


Species

Valid: * ''Chalicotherium goldfussi'' J. J. Kaup, 1833. :The type species, it was found in Upper Miocene beds located in Germany. It weighed around 1500 kg and was 2.6 m high at the shoulder. * ''Chalicotherium brevirostris'' Colbert, 1934 :First described as ''Macrotherium brevirostris'', this species hails from the Upper Miocene Tung Gur Formation,
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an Autonomous regions of China, autonomous region of China. Its border includes two-thirds of the length of China's China–Mongolia border, border with the country of Mongolia. ...
, China. * ''Chalicotherium salinum'' Pickford, 1982 :First described as ''Macrotherium salinum'' by Forster Cooper, this species was first discovered at the Lower
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Siwaliks beds in India; its chronological and geographic range was later extended to the Middle and Upper
Miocene The Miocene ( ) is the first epoch (geology), 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 mea ...
, and to Pakistan and China, respectively. Invalid: * ''Chalicotherium antiquum'' J. J. Kaup, 1833. :Found at the same locality as the type species, it was later found wanting of diagnostic features and sunk into the type species. Misassigned specimens: * ''Chalicotherium'' cf. ''C. brevirostris'' Wang ''et al.'', 2001. :Hailing for the Tsaidam Basin, northern
Qinghai Qinghai is an inland Provinces of China, province in Northwestern China. It is the largest provinces of China, province of China (excluding autonomous regions) by area and has the third smallest population. Its capital and largest city is Xin ...
-
Tibetan Plateau The Tibetan Plateau, also known as the Qinghai–Tibet Plateau or Qingzang Plateau, is a vast elevated plateau located at the intersection of Central Asia, Central, South Asia, South, and East Asia. Geographically, it is located to the north of H ...
, China. * "Chalicotherium modicum" Stehlin, 1905. :A ''
nomen nudum In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...
'', actually a ''Schizotherium priscum'' tooth. * "''Chalicotherium''" ''bilobatum'' Cope. :Hailing from the Oligocene of
Saskatchewan Saskatchewan is a Provinces and territories of Canada, province in Western Canada. It is bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and to the south by the ...
, this very fragmentary specimen was the type on which Russel erected the genus '' Oreinotherium''. * ''Chalicotherium'' spp. :Specimens found in two
Tajikistan Tajikistan, officially the Republic of Tajikistan, is a landlocked country in Central Asia. Dushanbe is the capital city, capital and most populous city. Tajikistan borders Afghanistan to the Afghanistan–Tajikistan border, south, Uzbekistan to ...
localities, thought to pertain to at least two different species.


See also

* '' Anisodon'' * ''
Moropus ''Moropus'' (meaning "slow foot") is an extinct genus of large perissodactyl mammal in the chalicothere family. They were endemic to North America during the Miocene from ~20.4–13.6 Mya (unit), Mya, existing for approximately . ''Moropus'' belo ...
''


References


Further reading

* Grande Enciclopédia Portuguesa e Brasileira, vol. 5. (1936-1960). Editorial Enciclopédia, Lda, Lisbon. * * Butler, P. M., 1978: Chalicotheriidae. 368–370. in Maglio, V. J. & Cooke, H. B. S., (eds.) 1978: ''Evolution of African mammals.'' – Harvard University Press, Cambridge, Massachusetts & London, England, 1978, xiv-641 * Carroll, R. L., 1988: ''Vertebrate paleontology and evolution.'' – W. H. Freeman and company, New York, 1988, 698. * Carroll, R. L., 1988: Appendix. 594–648. in Carroll, R. L., 1988: ''Vertebrate paleontology and evolution''. – W. H. Freeman and company, New York, 1988, 698. * Coombs, M. C., 1989: Interrelationships and diversity in the Chalicotheriidae. 438–457. in Prothero, D. R. & Schoch, R. M., (eds.) 1989: ''The Evolution of Perissodactyls''. – Oxford University Press, New York, New York & Oxford, England, 1989, ix-537 * Coombs, M. C., Hunt, Jr, R. M., Stepleton, E., Albright III, L. B. & Fremd, T. J., 2001: Stratigraphy, chronology, biogeography, and taxonomy of Early Miocene small chalicotheres in North America. – ''Journal of Vertebrate Paleontology: Vol. 21, #3'', pp. 607–620 * Geraads, D., Spassov, N. & Kovachev, D., 2001: New Chalicotheriidae (Perissodactyla, Mammalia) from the Late Miocene of Bulgaria. – Journal of Vertebrate Paleontology: Vol. 21, #3, pp. 569–606 * Hooker, J. J. & Dashzeveg, D., 2004: The origin of chalicotheres (Perissodactyla, Mammalia) – ''Palaeontology: Vol. 47, #6'', pp. 1363–1386 * Lucas, S. G. & Schoch, R. M., 1989: Taxonomy and biochronology of Eomoropus and Grangeria, Eocene chalicotheres from the western United States and China. 422–437. in Prothero, D. R. & Schoch, R. M., (eds.) 1989: ''The Evolution of Perissodactyls''. – Oxford University Press, New York, New York & Oxford, England, 1989, ix-537 * McKenna, M. C. & Bell, S. K., (eds.) 1997: ''Classification of mammals – above the species level''. – Columbia University Press, New York, 1997, xii-631 * Prothero, D. R. & Schoch, R. M., 1989: Classification of the Perissodactyla. 530–537. in Prothero, D. R. & Schoch, R. M., (eds.) 1989: ''The Evolution of Perissodactyls''. – Oxford University Press, New York, New York & Oxford, England, 1989, ix-537 * Remy, J.-A., Jaeger, J.-J., Chaimanee, Y., Soe, U. A. N., Marivaux, L., Sudre, J., Tun, S. T., Marandat, B. & Dewaele, E., 2005: A new chalicothere from the Pondaung Formation (late Middle Eocene) of Myanmar. – ''Comptes Rendus de l'Académie des Sciencies, Paris: Palevol: Vol. 4'', pp. 341–349 * ''The America Heritage Dictionary of English Language''. 2004, 2000. Houghton Mifflin Company. * * Wang, Xiaoming; Wang, Banyue (2001): New material of Chalicotherium from the Tsaidam Basin in the northern Qinghai-Tibetan Plateau, China. ''Paläontologische Zeitschrift, Vol 75, Fascicle 2''. Pages 219–226. * Margery Chalifoux Coombs
Additional ''Schizotherium'' material from China, and a review of ''Schizotherium'' dentitions (Perissodactyla, Chalicotheriidae). April 24, 1978. ''American Museum Novitates nr 2647''. pages 1-18.
The American Museum Of Natural History. New York City, N. Y. * Edwin H. Colbert
Distributional and phylogenetic studies on Indian fossil mammals. III – A classification of the Chalicotherioidea. May 8, 1935 ''American Museum Novitates nr 798, 56.9 (54)''
The American Museum Of Natural History. New York City. * {{Taxonbar, from=Q133349 Chalicotheriinae Miocene Perissodactyla Zanclean extinctions Miocene mammals of Europe Miocene mammals of Africa Fossil taxa described in 1833