Embrithopoda ("heavy-footed") is an
order
Order, ORDER or Orders may refer to:
* A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
...
of extinct
mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s known from Asia, Africa and
Eastern Europe
Eastern Europe is a subregion of the Europe, European continent. As a largely ambiguous term, it has a wide range of geopolitical, geographical, ethnic, cultural and socio-economic connotations. Its eastern boundary is marked by the Ural Mountain ...
. Most of the embrithopod genera are known exclusively from jaws and teeth dated from the late
Paleocene
The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
to the late
Eocene
The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
; however, the order is best known from its terminal member, the elephantine ''
Arsinoitherium
''Arsinoitherium'' is an extinct genus of paenungulate mammals belonging to the extinct Order (biology), order Embrithopoda. It is related to elephants, sirenians, and hyraxes. Arsinoitheres were superficially rhinoceros-like herbivores that live ...
''.
Description
While embrithopods bore a superficial resemblance to
rhinoceros
A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...
es, their horns had bony cores covered in
keratin
Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
ized skin. Not all embrithopods possessed horns, either. Despite their appearance, they have been regarded as related to
elephant
Elephants are the largest living land animals. Three living species are currently recognised: the African bush elephant ('' Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian elephant ('' Elephas maximus ...
s, not
perissodactyl
Perissodactyla (, ), or odd-toed ungulates, is an order of Ungulate, ungulates. The order includes about 17 living species divided into three Family (biology), families: Equidae (wild horse, horses, Asinus, asses, and zebras), Rhinocerotidae ( ...
s.
As
tethytheres, the Embrithopoda have been believed to be part of the clade
Afrotheria
Afrotheria ( from Latin ''Afro-'' "of Africa" + ''theria'' "wild beast") is a superorder of placental mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephan ...
. However, a study of the basal arsinoitheriid, ''Palaeoamasia'', suggests that embrithopods are not
tethytheres or even
paenungulates, and that they need to be better sampled in an analysis of
eutheria
Eutheria (from Greek , 'good, right' and , 'beast'; ), also called Pan-Placentalia, is the clade consisting of Placentalia, placental mammals and all therian mammals that are more closely related to placentals than to marsupials.
Eutherians ...
n relationships to clarify if they are even
afrotheria
Afrotheria ( from Latin ''Afro-'' "of Africa" + ''theria'' "wild beast") is a superorder of placental mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephan ...
ns.
[[Erdal et al.'s inclusion of Embrithopoda in Tabuce et al. (2007) finds them outside of (Ungulata + Afrotheria). Since this clade is not supported by molecular data, it suggests the need to explore the relationships of embrithopods, as they could potentially have affinities with laurasiatheres and "true" ungulates.]] It is also not clear if embrithopods originated in Africa or Eurasia.
However, recent findings demonstrate an African origin for embrithopods and furthermore a relationship with other paenungulates, albeit having diverged earlier than previously thought.
Fossils of embrithopods, such as ''
Arsinoitherium
''Arsinoitherium'' is an extinct genus of paenungulate mammals belonging to the extinct Order (biology), order Embrithopoda. It is related to elephants, sirenians, and hyraxes. Arsinoitheres were superficially rhinoceros-like herbivores that live ...
'', have been found in
Egypt
Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
,
Ethiopia
Ethiopia, officially the Federal Democratic Republic of Ethiopia, is a landlocked country located in the Horn of Africa region of East Africa. It shares borders with Eritrea to the north, Djibouti to the northeast, Somalia to the east, Ken ...
,
Kenya
Kenya, officially the Republic of Kenya, is a country located in East Africa. With an estimated population of more than 52.4 million as of mid-2024, Kenya is the 27th-most-populous country in the world and the 7th most populous in Africa. ...
,
Morocco
Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
,
Mongolia
Mongolia is a landlocked country in East Asia, bordered by Russia to the north and China to the south and southeast. It covers an area of , with a population of 3.5 million, making it the world's List of countries and dependencies by po ...
,
Turkey
Turkey, officially the Republic of Türkiye, is a country mainly located in Anatolia in West Asia, with a relatively small part called East Thrace in Southeast Europe. It borders the Black Sea to the north; Georgia (country), Georgia, Armen ...
,
Romania
Romania is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern and Southeast Europe. It borders Ukraine to the north and east, Hungary to the west, Serbia to the southwest, Bulgaria to the south, Moldova to ...
,
Namibia
Namibia, officially the Republic of Namibia, is a country on the west coast of Southern Africa. Its borders include the Atlantic Ocean to the west, Angola and Zambia to the north, Botswana to the east and South Africa to the south; in the no ...
,
Tunisia
Tunisia, officially the Republic of Tunisia, is a country in the Maghreb region of North Africa. It is bordered by Algeria to the west and southwest, Libya to the southeast, and the Mediterranean Sea to the north and east. Tunisia also shares m ...
and
Croatia
Croatia, officially the Republic of Croatia, is a country in Central Europe, Central and Southeast Europe, on the coast of the Adriatic Sea. It borders Slovenia to the northwest, Hungary to the northeast, Serbia to the east, Bosnia and Herze ...
. Until the 1970s, only ''Arsinoitherium'' itself was known, appearing isolated in the fossil record.
Classification
and considered ''Phenacolophus'' from Mongolia a primitive embrithopod, although this attribution was challenged by several other authors. A 2016 cladistic study found ''Phenacolophus'' as a stem-perissodactyl and the embrithopods at the base of
Altungulata
Altungulata or Pantomesaxonia ('' sensu'' and later authors) is an invalid clade ( mirorder) of ungulate mammals comprising the perissodactyls, hyracoids, and tethytheres ( sirenians, proboscideans, and related extinct taxa).
The name "Pant ...
.
[; ; ] More recently, an afrothere identity has been vindicated, albeit more basal than previously assumed.
Order Embrithopoda
sensu Prothero & Schoch 1989 (=Barypoda
Andrews 1904)
[ Mikko's Phylogeny Archivebr>]
* Genus †'
Gheerbrant ''et al'', 2018
* Family †
Arsinoitheriidae
** Genus †'
Pickford ''et al.'', 2008
** Genus †''
Arsinoitherium
''Arsinoitherium'' is an extinct genus of paenungulate mammals belonging to the extinct Order (biology), order Embrithopoda. It is related to elephants, sirenians, and hyraxes. Arsinoitheres were superficially rhinoceros-like herbivores that live ...
''
* Family †
Palaeoamasiidae
** Genus †''
Hypsamasia''
** Genus †''
Palaeoamasia''
** Genus †''
Crivadiatherium
''Crivadiatherium'' is an extinct genus of Palaeoamasiidae, which fossil remains—teeth and mandible fragments—have been discovered in the Crivadia site in the Hațeg depression, Romania. The age of the Crivadia site is not clear, but seem ...
''
Notes
References
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{{Taxonbar, from=Q131666
Eocene mammals
Oligocene mammals
Rupelian extinctions