Chalicotherium
Encyclopedia
Chalicotherium is a genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of extinct browsing odd-toed ungulate
Odd-toed ungulate
An odd-toed ungulate is a mammal with hooves that feature an odd number of toes. Odd-toed ungulates comprise the order Perissodactyla . The middle toe on each hoof is usually larger than its neighbours...

s of the order Perissodactyla and family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 Chalicotheriidae, found in Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

, Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

, and Asia
Asia
Asia is the world's largest and most populous continent, located primarily in the eastern and northern hemispheres. It covers 8.7% of the Earth's total surface area and with approximately 3.879 billion people, it hosts 60% of the world's current human population...

 during the Late Oligocene to Lower Pliocene, living from 16—7.75 mya, existing for approximately .

This animal would look much like other chalicotheriid species: an odd looking herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

 with long claw
Claw
A claw is a curved, pointed appendage, found at the end of a toe or finger in most mammals, birds, and some reptiles. However, the word "claw" is also often used in reference to an invertebrate. Somewhat similar fine hooked structures are found in arthropods such as beetles and spiders, at the end...

ed forelimb
Forelimb
A forelimb is an anterior limb on an animal's body. When referring to quadrupeds , the term foreleg is often instead used....

s and stouter weight bearing
Weight bearing
In orthopedics, weight-bearing is the amount of weight a patient puts on the leg on which surgery has been performed. In general, it is described as a percentage of the body weight, because each leg of a healthy person carries the full body weight when walking, in an alternating fashion.After...

 hindlimbs.

The type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

, Chalicotherium goldfussi, from Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...

 and Pliocene Europe, was described by J. J. Kaup in 1833 and since then 7 other species have been confidently assigned to this genus. According to current phylogenetic analyses Chalicotherium has two daughter genera nested within it, Anisodon
Anisodon
Anisodon is an extinct genus of chalicothere.-References:* Classification of Mammals by Malcolm C. McKenna and Susan K. Bell...

Lartet, 1851 and Nestoritherium
Nestoritherium
Nestoritherium is an extinct genus of chalicothere....

J. J. Kaup, 1859, thus rendering it paraphyletic.

Description

Chalicotherium, like many members of Perissodactyla, was adapted to browsing, though uniquely adapted to do so among its ungulate relatives. Its arms were long and heavily clawed, allowing them to walk on their knuckles only. The arms were used to reach for high branches and bring them close to its short-faced head to strip them clean of leaves. The horse
Horse
The horse is one of two extant subspecies of Equus ferus, or the wild horse. It is a single-hooved mammal belonging to the taxonomic family Equidae. The horse has evolved over the past 45 to 55 million years from a small multi-toed creature into the large, single-toed animal of today...

-like head itself shows adaptation to a diet of soft vegetation, since, as the animal reached sexual maturity
Sexual maturity
Sexual maturity is the age or stage when an organism can reproduce. It is sometimes considered synonymous with adulthood, though the two are distinct...

, the incisor
Incisor
Incisors are the first kind of tooth in heterodont mammals. They are located in the premaxilla above and mandible below.-Function:...

s and upper canines
Canine tooth
In mammalian oral anatomy, the canine teeth, also called cuspids, dogteeth, fangs, or eye teeth, are relatively long, pointed teeth...

 were shed, suggesting that its muscular lips and the resulting gum
Gingiva
The gingiva , or gums, consists of the mucosal tissue that lies over the mandible and maxilla inside the mouth.-General description:...

 pads were enough to crop fodder which was then processed by squarish, low-crowned molars.

Callosities
Callosity
A callosity is another name for callus, a piece of skin that has become thickened as a result of repeated contact and friction.- Monkeys :When occurring on an animal's buttocks, as with baboons, they are specifically called ischial 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
Phalanx bones
In anatomy, phalanx bones are those that form the fingers and toes. In primates such as humans and monkeys, the thumb and big toe have two phalanges, while the other fingers and toes consist of three. Phalanges are classified as long bones.The phalanges do not have individual names...

 is interpreted as evidence of knuckle-walking
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....

, which would probably be useful to avoid wearing down the claws, preserving them for use either as a forage-collecting rake
Rake (tool)
A rake is a broom for outside; an horticultural implement consisting of a toothed bar fixed transversely to a handle, and used to collect leaves, hay, grass, etc., and, in gardening, for loosening the soil, light weeding and levelling, removing dead grass from...

 or as a formidable defensive weapon.

All of these characteristics show some convergence with the unrelated ground sloth
Ground sloth
Ground sloths are a diverse group of extinct sloths, in the mammalian superorder Xenarthra. Their most recent survivors lived in the Antilles, where it has been proposed they may have survived until 1550 CE; however, the youngest AMS radiocarbon date reported is 4190 BP, calibrated to c. 4700 BP...

s, gorilla
Gorilla
Gorillas are the largest extant species of primates. They are ground-dwelling, predominantly herbivorous apes that inhabit the forests of central Africa. Gorillas are divided into two species and either four or five subspecies...

s and giant panda
Giant Panda
The giant panda, or panda is a bear native to central-western and south western China. It is easily recognized by its large, distinctive black patches around the eyes, over the ears, and across its round body. Though it belongs to the order Carnivora, the panda's diet is 99% bamboo...

s.

Taxonomic history

The type specimens for Chalicotherium goldfussi were found in the Upper Miocene strata of the Dinotherien-sande beds near Eppelsheim
Eppelsheim
Eppelsheim is an Ortsgemeinde – a municipality belonging to a Verbandsgemeinde, a kind of collective municipality – in the Alzey-Worms district in Rhineland-Palatinate, Germany.- Municipal council :...

, in the Grand Duchy of Hesse
Grand Duchy of Hesse
The Grand Duchy of Hesse and by Rhine , or, between 1806 and 1816, Grand Duchy of Hesse —as it was also known after 1816—was a member state of the German Confederation from 1806, when the Landgraviate of Hesse-Darmstadt was elevated to a Grand Duchy, until 1918, when all the German...

, Germany. 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
Sansan is a commune in the Gers department in southwestern France.The vicinity of Sansan is known for its Miocene fossil deposits where geologist Edouard Lartet unearthed the jaw of the primate Pliopithecus antiquus in 1837.-Population:...

 in the department of Gers
Gers
The Gers is a department in the Midi-Pyrénées region in the southwest of France named after the Gers River.Inhabitants are called les Gersois or Gersoises.-History:...

, Southwestern France, were first described as Macrotherium by Lartet in 1837. Further study of these fossil remains and subsequent finds by Filhol
Henri Filhol
Henri Filhol was a French medical doctor, malacologist and naturalist. He served as the expedition doctor and naturalist on the French 1874 Transit of Venus expedition to Campbell Island, New Zealand, with a peak on the island, Filhol Peak, being named after him...

 warranted a referral of the material described as Macrotherium to Chalicotherium. A more recently synonymized genus is Butleria, preoccupied by a butterfly
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...

 genus.

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 giganteum Pictet, 1844.
First described as Macrotherium giganteum Gervais, later sunk into the type species for lack of distinguishing skeletal morphology, it was found in the Upper Miocene strata located at Sansan, France.
  • Chalicotherium brevirostris
First described as Macrotherium brevirostris Colbert 1934, this species hails from the Upper Miocene Tung Gur Formation, Inner Mongolia
Inner Mongolia
Inner Mongolia is an autonomous region of the People's Republic of China, located in the northern region of the country. Inner Mongolia shares an international border with the countries of Mongolia and the Russian Federation...

, China.
  • Chalicotherium rusingense Butler, 1962.
This species hails from the Lower Miocene strata located in Kenya
Kenya
Kenya , officially known as the Republic of Kenya, is a country in East Africa that lies on the equator, with the Indian Ocean to its south-east...

 and Uganda
Uganda
Uganda , officially the Republic of Uganda, is a landlocked country in East Africa. Uganda is also known as the "Pearl of Africa". It is bordered on the east by Kenya, on the north by South Sudan, on the west by the Democratic Republic of the Congo, on the southwest by Rwanda, and on the south by...

.
  • Chalicotherium pilgrimi
Formerly known as Schizotherium pilgrimi Forster Cooper, then referred to Macrotherium, this species hails from the Lower Miocene Bugti beds of Pakistan
Pakistan
Pakistan , officially the Islamic Republic of Pakistan is a sovereign state in South Asia. It has a coastline along the Arabian Sea and the Gulf of Oman in the south and is bordered by Afghanistan and Iran in the west, India in the east and China in the far northeast. In the north, Tajikistan...

.
  • Chalicotherium wetzleri
Formerly named as Schizotherium wetzleri this species hails from Oligocene beds located in France and Germany, leading scientists to believe it was widespread in Western Europe.
  • Chalicotherium salinum
First described as Macrotherium salinum Forster Cooper, this species was first discovered at the Lower Pliocene Lower Siwaliks beds in India; its chronological and geographic range was later extended to the Middle and Upper Miocene, and to Pakistan and China, respectively.
  • Chalicotherium wuduensis
This species hails from Upper Miocene strata located in Gansu
Gansu
' is a province located in the northwest of the People's Republic of China.It lies between the Tibetan and Huangtu plateaus, and borders Mongolia, Inner Mongolia, and Ningxia to the north, Xinjiang and Qinghai to the west, Sichuan to the south, and Shaanxi to the east...

, 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.
  • Chalicotherium baltavarense
This name pertains to a specimen found in Slovakia
Slovakia
The Slovak Republic is a landlocked state in Central Europe. It has a population of over five million and an area of about . Slovakia is bordered by the Czech Republic and Austria to the west, Poland to the north, Ukraine to the east and Hungary to the south...

 that was later lumped into Chalicotherium pentelicum.
  • Chalicotherium minus
For sometime considered a species of Anisodon, it was later sunk into the type species.
  • Chalicotherium posterigenium
Junior synonym of Chalicotherium sivalense
  • Chalicotherium sindiense
Junior synonym of Chalicotherium sivalense
  • Chalicotherium sinense
This species hailing from Lower Pliocene beds found in China is now assigned to the genus Nestoritherium.
  • Chalicotherium sivalense
Erected from specimens found in Upper Miocene beds located in India it is now assigned to the genus Nestoritherium.
  • Chalicotherium pentelicum (pentelici in some older publications)
Hailing from Upper Miocene strata of Slovakia
Slovakia
The Slovak Republic is a landlocked state in Central Europe. It has a population of over five million and an area of about . Slovakia is bordered by the Czech Republic and Austria to the west, Poland to the north, Ukraine to the east and Hungary to the south...

 and Greece
Greece
Greece , officially the Hellenic Republic , and historically Hellas or the Republic of Greece in English, is a country in southeastern Europe....

, Samos Island
Samos Island
Samos is a Greek island in the eastern Aegean Sea, south of Chios, north of Patmos and the Dodecanese, and off the coast of Asia Minor, from which it is separated by the -wide Mycale Strait. It is also a separate regional unit of the North Aegean region, and the only municipality of the regional...

 being a notable locality of the latter, the species named Colodus pachygnathum was later subsumed with it. Further study referred this species to the genus Ancylotherium.
  • Chalicotherium grande Blainville, 1849.
Further study after this species' first description as Macrotherium grande Lartet, 1837, and subsequent referral to Chalicotherium by Blainville and later on of M. rhodanicum, M. magnum and M. sansaniense, warranted all the pertaining specimens to be clustered under the revived genus Anisodon Lartet, 1851, by Geraads et al. in 1995, as Anisodon grande.


Attributed specimens:
  • Chalicotherium cf. C. brevirostris Wang et al., 2001.
Hailing for the Tsaidam Basin, northern Qinghai
Qinghai
Qinghai ; Oirat Mongolian: ; ; Salar:) is a province of the People's Republic of China, named after Qinghai Lake...

-Tibetan Plateau
Tibetan Plateau
The Tibetan Plateau , also known as the Qinghai–Tibetan Plateau is a vast, elevated plateau in Central Asia covering most of the Tibet Autonomous Region and Qinghai, in addition to smaller portions of western Sichuan, southwestern Gansu, and northern Yunnan in Western China and Ladakh in...

, China.
  • "Chalicotherium modicum" Stehlin, 1905.
A nomen nudum
Nomen nudum
The phrase nomen nudum is a Latin term, meaning "naked name", used in taxonomy...

, actually a Schizotherium priscum tooth.
  • "Chalicotherium" bilobatum Cope.
Hailing from the Oligocene of Saskatchewan
Saskatchewan
Saskatchewan is a prairie province in Canada, which has an area of . Saskatchewan is bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, and on the south by the U.S. states of Montana and North Dakota....

, this very fragmentary specimen was the type on which Russel erected the genus Oreinotherium
Oreinotherium
Oreinotherium is a genus of brontothere.According to one source, the species of Oreinotherium are merged into the genus Megacerops.-References:...

.
  • Chalicotherium spp.
Specimens found in two Tajikistan
Tajikistan
Tajikistan , officially the Republic of Tajikistan , is a mountainous landlocked country in Central Asia. Afghanistan borders it to the south, Uzbekistan to the west, Kyrgyzstan to the north, and China to the east....

 localities, thought to pertain to at least two different species.

Phylogeny

unnamed

In popular culture

While the Chalicotherium is not well known in popular culture, the 2001 educational documentary Walking with Beasts
Walking with Beasts
Walking with Beasts is a 2001 six-part television documentary miniseries, produced by the BBC in the United Kingdom, narrated by Kenneth Branagh. In North America it has been retitled Walking with Prehistoric Beasts, and the original Discovery Channel broadcast was narrated by Stockard Channing...

included the animal, adding some publicity.

Chalicotherium does appear briefly in the animated sequel to Ice Age (film)
Ice Age (film)
Ice Age is a 2002 American computer-animated film created by Blue Sky Studios and released by 20th Century Fox. It was directed by Carlos Saldanha and Chris Wedge from a story by Michael J. Wilson. The story follows three Paleolithical mammals attempting to return a lost human baby to its parents...

, Ice Age: The Meltdown
Ice Age: The Meltdown
Ice Age: The Meltdown, also known as Ice Age 2: The Meltdown or simply as Ice Age 2, is the 2006 sequel to the 2002 computer-animated film Ice Age. It was produced by Blue Sky Studios for 20th Century Fox, and premiered in Belgium on March 1, 2006...

.

In addition, "chalikos" are used as steeds in Julian May's Saga of Pliocene Exile
Saga of Pliocene Exile
The Saga of Pliocene Exile is a series of science / speculative fiction books by Julian May, first published in the early 1980s...

, a time travel series in which humans go through a gateway back to the Pliocene Era.
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