Macrotherium
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''Chalicotherium'' (
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic p ...
/, -: pebble/ gravel + /, diminutive of / : beast) is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of extinct
odd-toed ungulate Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) o ...
s of the order Perissodactyla and
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Chalicotheriidae. The genus is known from
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
and
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
, from the Middle Miocene to Late Miocene. This animal would look much like other chalicotheriid species: an odd-looking herbivore 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 tarsus ...
ed forelimbs and stouter weight-bearing hindlimbs. The
type species In 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 species that contains the biological type specime ...
, ''Chalicotherium goldfussi'', from Late Miocene Europe, was described by Johann Jakob Kaup in 1833. When the French naturalist George Cuvier first received a cleft claw from Eppelheim, Germany, he identified it as the toe bone of a gigantic pangolin.


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 million yea ...
-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 might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definit ...
, 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 Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth, in mammalian oral anatomy People with the surn ...
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 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 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, bonobos, and chimpanzees use this style o ...
, which would probably be useful to avoid wearing down the claws, preserving them for use either as a forage-collecting rake or as a formidable defensive weapon, 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 sloths,
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, bears (especially giant pandas), and a group of theropod dinosaurs known as
therizinosaur Therizinosaurs (once called segnosaurs) were large herbivorous theropod dinosaurs whose fossils have been found across the Early to Late Cretaceous deposits in Asia and North America. Various features of the forelimbs, skull and pelvis unite thes ...
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, Germany. Johann Jakob 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 in the department of Gers, 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. Biography Lartet was born near Castelnau-Barbarens, ' of Gers, France, where his family had lived for more than ...
in 1837. Further study of these fossil remains and subsequent finds by Filhol warranted a referral of the material described as ''Macrotherium'' to ''Chalicotherium''.Anquetin,J.,Antoine,P.-O.,Tassy,P.,2007.Middle Miocene Chalicotheriinae(Mammalia, Perissodactyla)from France, with a discussion on chalicotheriine phylogeny. Zoological Journal of the Linnean Society 151, 577–608. 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'' :First described as ''Macrotherium brevirostris'' Colbert 1934, this species hails from the Upper Miocene Tung Gur Formation,
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an autonomous region of the People's Republic of China. Its border includes most of the length of China's border with the country of Mongolia. Inner Mongolia also accounts for a ...
, China. 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 (; alternately romanized as Tsinghai, Ch'inghai), also known as Kokonor, is a landlocked province in the northwest of the People's Republic of China. It is the fourth largest province of China by area and has the third smallest po ...
-
Tibetan Plateau The Tibetan Plateau (, also known as the Qinghai–Tibet Plateau or the Qing–Zang Plateau () or as the Himalayan Plateau in India, is a vast elevated plateau located at the intersection of Central, South and East Asia covering most of the ...
, China. * "Chalicotherium modicum" Stehlin, 1905. :A '' nomen nudum'', actually a ''Schizotherium priscum'' tooth. * "''Chalicotherium''" ''bilobatum'' Cope. :Hailing from the Oligocene of
Saskatchewan Saskatchewan ( ; ) is a province in western Canada, bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and on the south by the U.S. states of Montana and North Dak ...
, this very fragmentary specimen was the type on which Russel erected the genus '' Oreinotherium''. * ''Chalicotherium'' spp. :Specimens found in two
Tajikistan Tajikistan (, ; tg, Тоҷикистон, Tojikiston; russian: Таджикистан, Tadzhikistan), officially the Republic of Tajikistan ( tg, Ҷумҳурии Тоҷикистон, Jumhurii Tojikiston), is a landlocked country in Centr ...
localities, thought to pertain to at least two different species.


See also

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


References

* Grande Enciclopédia Portuguesa e Brasileira, vol. 5. (1936-1960). Editorial Enciclopédia, Lda, Lisbon.
''Chalicotherium'' at Atlas Virtual da Pré-História
Retrieved 22 March 2007. * 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.
www.angellis.net/Web/PDfiles/ungperis.pdf
{{Taxonbar, from=Q133349 Chalicotheres Miocene odd-toed ungulates Zanclean extinctions Neogene mammals of Europe Cenozoic mammals of Africa Fossil taxa described in 1833