Chalicotheres (from
Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
''
chalix'', "gravel" and ''
therion'', "beast") are an extinct
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of
herbivorous
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
,
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) ...
(perissodactyl)
mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s that lived in
North America
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
,
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 a ...
, and
Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
from the
Middle Eocene
The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "dawn ...
until the
Early Pleistocene
The Early Pleistocene is an unofficial sub-epoch in the international geologic timescale in chronostratigraphy, being the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently estimated to span the time ...
, existing from 48.6 to 1.806
mya
Mya may refer to:
Brands and product names
* Mya (program), an intelligent personal assistant created by Motorola
* Mya (TV channel), an Italian Television channel
* Midwest Young Artists, a comprehensive youth music program
Codes
* Burmese ...
. They are one of the five major
radiations
An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rapid ...
of perissodactyls, with three groups living (
horses
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 ...
, plus the extinct
paleotheres;
rhinoceroses
A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species ...
;
tapirs
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 inhabit ...
), and two extinct (
brontotheres
Brontotheriidae is a family of extinct mammals belonging to the order Perissodactyla, the order that includes horses, rhinoceroses, and tapirs. Superficially, they looked rather like rhinos, although they were actually more closely related to ...
and chalicotheres).
Description
Unlike modern perissodactyls, chalicotheres had clawed feet. They had longer forelimbs and shorter hind limbs, lower incisors that cropped food against a toothless pad in the upper jaw, low-crowned molar teeth, and were browsers on trees and shrubs throughout their history. They evolved in two different directions, which became separate subfamilies, the
Schizotheriinae
Schizotheriines are one of the two subfamilies of the extinct family Chalicotheriidae, a group of herbivorous odd-toed ungulate (perissodactyl) mammals that lived from the Eocene to the Pleistoscene. The other clade is the Chalicotheriinae. Both ...
and the
Chalicotheriinae
Chalicotheriines are one of the two subfamilies of the extinct family Chalicotheriidae, a group of herbivorous, odd-toed ungulate ''(perissodactyl)'' mammals that lived from the Eocene to the Pleistocene. The other subfamily is the Schizotherii ...
.
Schizotherine chalicotheres such as ''
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 . ...
'' lived in a variety of forest, woodland, and savannah habitats in Asia, Africa, and North and Central America. They developed long necks and skull adaptations that suggest they had long, extensible tongues to reach browse, like those of
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 ...
. Strong hindlimbs and an elongated pelvis suggest they could have reared upright as modern
goats
The goat or domestic goat (''Capra hircus'') is a domesticated species of Caprinae, goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a membe ...
do, and used their front claws to pull branches within reach of the tongue. The claws were retractable, and they walked normally on the bottom of the foot. Studies of tooth wear suggest they ate leaves, twigs, fruit, and bark.
Chalicotheriines, such as ''
Anisodon
''Anisodon'' (Greek: "unequal" (anisos), "teeth" (odontes)) is an extinct genus of chalicothere that lived in Europe during the late Miocene. It stood at about 150 cm and weighed around 600 kg. It is thought that the animal's clawed forelimbs wo ...
,'' lived only in moist, closed-canopy forests, never reached the Americas, and developed very unusual anatomy for an ungulate. Their shorter necks and horse-like heads did not show adaptations to reach high. Instead, they developed very long forelimbs with mobile shoulder joints and hooklike claws. The pelvis and hindlimbs were specialized to stand upright, and to sit for hours while feeding, like the living
gelada
The gelada (''Theropithecus gelada'', am, ጭላዳ, translit=č̣əlada), sometimes called the bleeding-heart monkey or the gelada baboon, is a species of Old World monkey found only in the Ethiopian Highlands, living at elevations of above se ...
monkey. Some early paleontologists thought the claws were used to dig up roots and tubers, but their teeth were designed for soft foods, and studies of tooth wear show they ate fruit and seeds. Their forelimbs were specialized to reach, grasp, and strip or sweep plants to the mouth. They could not retract the huge front claws, and knuckle-walked on their forelimbs. The chalicotheriines' anatomical design, posture, and locomotion show
convergence
Convergence may refer to:
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* "Convergence" (comics), two separate story lines published by DC Comics:
**A four-part crossover storyline that united the four Wei ...
with other large browsers that feed selectively in a bipedal position, such as the
ground sloth
Ground sloths are a diverse group of extinct sloths in the mammalian superorder Xenarthra. The term is used to refer to all extinct sloths because of the large size of the earliest forms discovered, compared to existing tree sloths. The Caribbe ...
s,
gorillas
Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
, and
giant panda
The giant panda (''Ailuropoda melanoleuca''), also known as the panda bear (or simply the panda), is a bear species endemic to China. It is characterised by its bold black-and-white coat and rotund body. The name "giant panda" is sometimes us ...
s.
Chalicothere fossils are uncommon even in areas where other taxa of similar size are well-preserved, which suggests they were mostly solitary animals, and unlike horses, rhinos, and brontotheres, never evolved species that lived in herds. Only two species of chalicothere are known from complete skeletons, the schizotheriine ''Moropus'' from the early Miocene of North America, and the chalicotheriine ''Anisodon'' from the middle Miocene of Europe. Fossils of other species range from very fragmentary to moderately complete. Chalicotheres ranged in size from an antelope to a large draft horse.
Evolution
Chalicotheres can be first identified with certainty around 46 million years ago, in the Eocene of Asia. The family is thought to have evolved there, but appeared in North America by the Eocene. By the late
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
, they had divided into schizotheriines and chalicotheriines. (Earlier chalicotheres are often referred to the family Eomoropidae; it is not yet clear whether they had claws or how the two subfamilies diverged.)
Both subfamilies were successful over many millions of years, and reached their greatest diversity in the Miocene. Advanced schizotheriines (''Moropus'') entered North America via the Bering land bridge at the Oligicene-Miocene boundary, and expanded southward into Central America. There were multiple radiations into Africa, where chalcotheriines were later replaced by schizotheriines. Both groups spread early into Europe. In the Pliocene, they would have faced new competition in North America from megalonychid ground sloths which emigrated from South America in the
Great American Interchange
The Great American Biotic Interchange (commonly abbreviated as GABI), also known as the Great American Interchange and the Great American Faunal Interchange, was an important late Cenozoic paleozoogeographic biotic interchange event in which lan ...
, and from evolving great apes in African and European forests. The family became less successful after the Miocene but persisted in Africa until the end of the Pleistocene; the latest surviving species was the schizotheriine ''
Ancylotherium
''Ancylotherium'' (from Greek, meaning "hooked beast") is an extinct genus of the family Chalicotheriidae, subfamily Schizotheriinae, endemic to Europe, Asia, and Africa during the Late Miocene-Early Pleistocene (9.0—1.8 mya), existing for app ...
hennigi.''
Chalicotheres are related to the extinct
brontothere
Brontotheriidae is a family of extinct mammals belonging to the order Perissodactyla, the order that includes horses, rhinoceroses, and tapirs. Superficially, they looked rather like rhinos, although they were actually more closely related to ...
s, as well as to modern day
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 y ...
s,
rhinoceros
A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species o ...
es, and
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 inhabit ...
s.
As the early evolution of perissodactyls is still unresolved, their closest relatives among other perissodactyl groups is obscure. They have been traditionally ranked as closer to Ceratomorpha (tapirs+rhinos) than Equoidea (horses, etc.). However, a 2004 cladistic study recovers chalicotheres as the sister group to
Lophiodontidae
Lophiodontidae is a family of browsing, herbivorous, mammals in the Perissodactyla suborder Ancylopoda that show long, curved and cleft claws. They lived in southern Europe during the Eocene epoch. Previously thought to be related to tapir
T ...
, and the combined group (
Ancylopoda
Ancylopoda is a group of browsing, herbivorous, mammals in the Perissodactyla that show long, curved and cleft claws. Morphological evidence indicates the Ancylopoda diverged from the tapirs, rhinoceroses and horses ( Euperissodactyla) after t ...
) as sister to all modern perissodactyls (Equoidea+Ceratomorpha), with the
brontotheres
Brontotheriidae is a family of extinct mammals belonging to the order Perissodactyla, the order that includes horses, rhinoceroses, and tapirs. Superficially, they looked rather like rhinos, although they were actually more closely related to ...
as the most distantly related within the order
Perissodactyla.
References
{{Spoken Wikipedia, Chalicothere.ogg, date=2017-12-30
Piacenzian extinctions
Clawed herbivores
Eocene first appearances
Taxa named by Theodore Gill