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The Palaeomerycidae are an extinct
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
ruminants Ruminants (suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microbial actions. The ...
in the order
Artiodactyla The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
. Palaeomerycids 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 ...
,
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 Continent#Subcontinents, subcontinent of Eurasia ...
,
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 ...
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 area ...
from 33 to 4.9 million years ago (from the Late
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
to
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
, but its identity as a palaeomerycid was subsequently disputed. Specimens of '' Amphitragulus'', the earliest known genus, have been found in
Aragon Aragon ( , ; Spanish and an, Aragón ; ca, Aragó ) is an autonomous community in Spain, coextensive with the medieval Kingdom of Aragon. In northeastern Spain, the Aragonese autonomous community comprises three provinces (from north to sou ...
(
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") , i ...
), Ronheim (Germany),
Sardinia Sardinia ( ; it, Sardegna, label=Italian, Corsican and Tabarchino ; sc, Sardigna , sdc, Sardhigna; french: Sardaigne; sdn, Saldigna; ca, Sardenya, label=Algherese and Catalan) is the second-largest island in the Mediterranean Sea, after ...
(
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...
),
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
, and
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a transcontinental country located mainly in Central Asia and partly in Eastern Europe. It borders Russia to the north and west, China to the east, Kyrgyzstan to the southeast, Uzbeki ...
and range from 33 to 15.97 million years old. Fossils of '' Lagomeryx'' and ''
Palaeomeryx ''Palaeomeryx'' is an extinct genus of Artiodactyla, of the family Palaeomerycidae, endemic to Europe and Asia from the Miocene epoch, 16.9 – 7.25 Ma, existing for approximately . Taxonomy ''Palaeomeryx'' was named by von Meyer (1834). ...
feignouxi'' were recovered in
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
and
Slovakia Slovakia (; sk, Slovensko ), officially the Slovak Republic ( sk, Slovenská republika, links=no ), is a landlocked country in Central Europe. It is bordered by Poland to the north, Ukraine to the east, Hungary to the south, Austria to the s ...
from the Middle Eocene. The oldest known North American specimen, '' Barbouromeryx'', was discovered in
Scotts Bluff, Nebraska Scottsbluff is a city in Scotts Bluff County, in the western part of the state of Nebraska, in the Great Plains region of the United States. The population was 14,436 at the 2020 census. Scottsbluff is the largest city in the Nebraska Panhand ...
, and estimated at 23.03 million years old. Palaeomerycids such as ''
Cranioceras ''Cranioceras'' is an extinct genus of artiodactyl from the Miocene to the Pliocene in the United States. Sources * After the Dinosaurs: The Age of Mammals (Life of the Past) by Donald R. Prothero * The Book of Life: An Illustrated History ...
'' and other Dromomerycinae continued to live in North America until the early Pliocene.


Description

Palaeomerycids were a group of horned, long-legged and massive ruminants that could attain a weight of . One of the first known members of this group, ''
Palaeomeryx ''Palaeomeryx'' is an extinct genus of Artiodactyla, of the family Palaeomerycidae, endemic to Europe and Asia from the Miocene epoch, 16.9 – 7.25 Ma, existing for approximately . Taxonomy ''Palaeomeryx'' was named by von Meyer (1834). ...
'', was thought to be a hornless form distantly related to the
Giraffidae The Giraffidae are a family of ruminant artiodactyl mammals that share a common ancestor with deer and bovids. This family, once a diverse group spread throughout Eurasia and Africa, presently comprises only two extant genera, the giraffe (one or ...
before paleontologist Miguel Crusafont found remains of '' Triceromeryx'' in middle Miocene Spain. This ''Palaeomeryx''-like form carried two
ossicone Ossicones are columnar or conical skin-covered bone structures on the heads of giraffes, male okapi, and some of their extinct relatives. Ossicones are distinguished from the superficially similar structures of horns and antlers by their uniqu ...
s over its orbits that were straight and short, similar to those of true giraffids. However, the most striking feature of ''Triceromeryx'' was the third, Y-shaped appendage that prolonged the
occipital bone The occipital bone () is a neurocranium, cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobe ...
at the back of the skull. Discoveries during the 1980s and 1990s showed a surprising variety in these occipital appendages. '' Ampelomeryx'', a genus of palaeomerycids found at the early Miocene site of Els Casots, Valles-Penedes Basin, Spain, had a three-horned system of appendages similar to those of ''Triceromeryx''. These appendages were, however, quite different, with the paired appendages extending laterally over the orbits, flat and wide, forming an eye-shade, while the third spectacular posterior appendage was about long. Another species of ''Triceromeryx'', '' T. conquensis'' found in La Retama in Spain, showed an even more spectacular appendage — instead of a Y-shaped structure, its posterior appendage was T-shaped with the lateral branches expanding toward the front. In primitive members of the group (e.g. ''Ampelomeryx''), this appendage was a posterior expansion of the occipital bone lying close to the powerful muscles supporting the skull in a normal position, thus suggesting that this appendage was actually used for fighting between males during the breeding season. The reduced shapes of the flat and laterally oriented appendages of later species suggests that these were not used in active fighting, instead forming a function of passive display. The limbs were more similar to those of modern large
bovid The Bovidae comprise the biological family of cloven-hoofed, ruminant mammals that includes cattle, bison, buffalo, antelopes, and caprines. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, ...
s (e.g. buffaloes) than to those of
okapi The okapi (; ''Okapia johnstoni''), also known as the forest giraffe, Congolese giraffe, or zebra giraffe, is an artiodactyl mammal that is endemic to the northeast Democratic Republic of the Congo in central Africa. It is the only species ...
s. They probably lived in boggy forests, living on soft leaves and aquatic plants (as hinted to by the
brachydont The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
teeth, which are similar to those of primitive giraffids). Their diets consisted of coarse plant material. As a group, the palaeomerycids appear to have formed a successful part of an independent radiation of horned ruminants that diversified into a variety of forms during the early to middle Miocene, with a geographic range reaching from Spain to China. The giraffids seem to have originated in Asia south of the Alpine belt, while the cervoids seem to have originated north of the Alpine belt.


Taxonomy

The Palaeomerycidae were named by Lydekker (1883). The type genus is ''Palaeomeryx''. The family was assigned to the Artiodactyla by Hulbert and Whitmore (2006), and to Cervoidea by Carroll (1988), Sach and Heizmann (2001) and Prothero and Liter (2007).


Classification

*'' Amphitragulus'' * Dromomerycinae **'' Asiagenes'' ** Aletomerycini ***'' Sinclairomeryx'' ***'' Aletomeryx'' ***'' Surameryx''? – interpretation as a palaeomerycid disputed ***'' Diabolocornis'' ** Dromomerycini ***'' Drepanomeryx'' ***'' Rakomeryx'' ***''
Dromomeryx ''Dromomeryx'' is an extinct genus of Artiodactyla The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes a ...
'' ***'' Subdromomeryx'' ** Cranioceratini ***'' Barbouromeryx'' ***'' Bouromeryx'' ***'' Procranioceras'' ***''
Cranioceras ''Cranioceras'' is an extinct genus of artiodactyl from the Miocene to the Pliocene in the United States. Sources * After the Dinosaurs: The Age of Mammals (Life of the Past) by Donald R. Prothero * The Book of Life: An Illustrated History ...
'' ***'' Pediomeryx'' ***'' Yumaceras'' *
Palaeomerycinae The Palaeomerycidae are an extinct family of ruminants in the order Artiodactyla. Palaeomerycids lived in North America, Europe, Africa and Asia from 33 to 4.9 million years ago (from the Late Eocene to Pliocene epochs), existing for about 28 m ...
**'' Ampelomeryx'' **''
Palaeomeryx ''Palaeomeryx'' is an extinct genus of Artiodactyla, of the family Palaeomerycidae, endemic to Europe and Asia from the Miocene epoch, 16.9 – 7.25 Ma, existing for approximately . Taxonomy ''Palaeomeryx'' was named by von Meyer (1834). ...
'' **'' Triceromeryx'' **'' Xenokeryx'' Genus '' Lagomeryx'' (together with its proposed subfamily Lagomerycinae) may be a further member of this family, as well as '' Ligeromeryx'' and '' Stephanocemas'' (formerly classified as members of
Cervidae Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...
).


References


External links

* {{Taxonbar, from=Q5493096 Eocene first appearances Miocene extinctions Prehistoric mammal families Pecora