Phenacodontidae
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Phenacodontidae is 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 large herbivorous
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 traditionally placed in the “
wastebasket taxon Wastebasket taxon (also called a wastebin taxon, dustbin taxon or catch-all taxon) is a term used by some taxonomists to refer to a taxon that has the sole purpose of classifying organisms that do not fit anywhere else. They are typically defined ...
Condylarthra Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a wast ...
, which may instead represent early-stage
perissodactyls 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 ...
. They lived in the Paleocene and Eocene
epochs In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided by ...
(about 60–50 million years ago) and their fossil remains have been found in North America and Europe.


Description

These animals had a variety of body sizes, and could be as small as domestic cats (''
Tetraclaenodon ''Tetraclaenodon'' was a genus of small and early ungulate mammals that was part of the Phenacodontidae Phenacodontidae is an extinct family (biology), family of large herbivorous mammals traditionally placed in the “wastebasket taxon” C ...
'' and ''
Ectocion ''Ectocion'' (sometimes ''Ectocyon'') is an extinct genus of placental mammals of the family Phenacodontidae. The genus was earlier classified as ''Gidleyina'' (Simpson 1935) and ''Prosthecion'' (Patterson and West 1973). Retrieved May 2013. Pa ...
'') and as large as sheep (''
Phenacodus ''Phenacodus'' (Greek: "deception" (phenax), "tooth' (odus)) is an extinct genus of mammals from the late Paleocene through middle Eocene, about 55 million years ago. It is one of the earliest and most primitive of the ungulates, typifying the fa ...
''). The skull of phenacodontids is long and narrow, and equipped with a small braincase. The skeleton of phenacodontids show several primitive characteristics (the long and heavy tail for example) but also a number of advanced, Perissodactyla-like adaptations: Their long legs, for example, had five fingers, but the first finger showed a clear reduction, and in some forms (like ''Phenacodus'') the fifth finger was reduced as well. Some species had tapir-like adaptations suggestive of the presence of a short proboscis or a strong prehensile lip. The teeth of phenacodontids, particularly in the latter forms, were quite specialized: The molars and premolars were equipped with low cusps that sometimes joined in ridges, similar to the condition found in some perissodactyls. Some forms, like ''
Meniscotherium ''Meniscotherium'' is an extinct genus of dog-sized mammal which lived 54–38 million years ago. It was a herbivore and had hooves. Fossils have been found in Utah, New Mexico. and Colorado. Many individuals have been found together, indicating ...
'', had enlarged ridges. This adaptation is unusual for mammals as old as phenacodontids. Only a few other archaic mammals possessed teeth with similar structures, such as ''
Pleuraspidotherium ''Pleuraspidotherium'' is an extinct genus of condylarth of the family Pleuraspidotheriidae, whose fossils have been found in the Late Paleocene Marnes de Montchenot of France and the Tremp Formation of modern Spain , image_flag ...
''.


Evolution

The phenacodontids evolved in the middle Paleocene in North America. Early forms were usually small; ''
Tetraclaenodon ''Tetraclaenodon'' was a genus of small and early ungulate mammals that was part of the Phenacodontidae Phenacodontidae is an extinct family (biology), family of large herbivorous mammals traditionally placed in the “wastebasket taxon” C ...
'', for example, was the size of a fox. Later forms were much larger and invaded Europe, although they never became as plentiful as in North America. Towards the beginning of the Eocene these animals slowly disappeared from the fossil record. Only a few forms survived into the middle Eocene: the ''
Phenacodus ''Phenacodus'' (Greek: "deception" (phenax), "tooth' (odus)) is an extinct genus of mammals from the late Paleocene through middle Eocene, about 55 million years ago. It is one of the earliest and most primitive of the ungulates, typifying the fa ...
'' in Europe and North America, ''
Almogaver ''Almogaver'' is an extinct possible odd-toed ungulate genus in the family Phenacodontidae. It was a ground-dwelling herbivore. It is known from the Tremp Basin in Spain, and some authorities consider it synonymous with ''Phenacodus ''Phenaco ...
'' in Europe and ''
Ectocion ''Ectocion'' (sometimes ''Ectocyon'') is an extinct genus of placental mammals of the family Phenacodontidae. The genus was earlier classified as ''Gidleyina'' (Simpson 1935) and ''Prosthecion'' (Patterson and West 1973). Retrieved May 2013. Pa ...
'' in North America. An exception to the scarcity of Eocene phenacodontids is the dog-sized genus ''
Meniscotherium ''Meniscotherium'' is an extinct genus of dog-sized mammal which lived 54–38 million years ago. It was a herbivore and had hooves. Fossils have been found in Utah, New Mexico. and Colorado. Many individuals have been found together, indicating ...
'', whose fossils are very abundant.


Classification

Phenacodontids have classically been included in the large group
Condylarthra Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a wast ...
, now considered
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
. In particular, the genus ''Phenacodus'' is often illustrated as a typical example of a "condylarth", due to the remarkable abundance of fossil remains. * Subfamily
Phenacodontinae Phenacodontinae was a subfamily of small herbivorous mammals that were part of the Phenacodontidae family. Genera The subfamily contains the following genera: * †''Almogaver'' * †''Copecion'' * †'' Lophocion'' * †'' Eodesmatodon'' * ...
Cope, 1881 ** Genus ''
Tetraclaenodon ''Tetraclaenodon'' was a genus of small and early ungulate mammals that was part of the Phenacodontidae Phenacodontidae is an extinct family (biology), family of large herbivorous mammals traditionally placed in the “wastebasket taxon” C ...
'' Scott, 1893 ***''Tetraclaenodon floverianus'' Cope, 1890 ***''Tetraclaenodon puercensis'' (Cope, 1881) ***''Tetraclaenodon septentrionalis'' Thewissen, 1990 ** Genus ''
Copecion ''Copecion'' was a genus of early herbivorous mammals that was part of the family Phenacodontidae Phenacodontidae is an extinct family (biology), family of large herbivorous mammals traditionally placed in the “wastebasket taxon” Condylart ...
'' Gingerich, 1989 ***''Copecion brachypternus'' (Cope, 1882) ***''Copecion davisi'' Gingerich, 1989 ** Genus ''
Ectocion ''Ectocion'' (sometimes ''Ectocyon'') is an extinct genus of placental mammals of the family Phenacodontidae. The genus was earlier classified as ''Gidleyina'' (Simpson 1935) and ''Prosthecion'' (Patterson and West 1973). Retrieved May 2013. Pa ...
'' Cope, 1882 ***''Ectocion cedrus'' Thewissen, 1990 ***''Ectocion collinus'' Russell, 1929 ***''Ectocion ignotum'' Novacek et al., 1991 ***''Ectocion major'' (Patteron & West, 1973) ***''Ectocion mediotuber'' Thewissen, 1990 ***''Ectocion osbornianus'' (Cope, 1882) ***''Ectocion parvus'' Granger, 1915 ***''Ectocion superstes'' Granger, 1915 ** Genus ''
Phenacodus ''Phenacodus'' (Greek: "deception" (phenax), "tooth' (odus)) is an extinct genus of mammals from the late Paleocene through middle Eocene, about 55 million years ago. It is one of the earliest and most primitive of the ungulates, typifying the fa ...
'' Cope, 1873 ***''Phenacodus bisonensis'' Gazin, 1956 ***''Phenacodus condali'' (Crusafont i Villalta, 1955) ***''Phenacodus grangeri'' Simpson, 1935 ***''Phenacodus intermedius'' Granger, 1915 ***''Phenacodus lemoinei'' Thewissen, 1990 ***''Phenacodus magnus'' Thewissen, 1990 ***''Phenacodus matthewi'' Simpson, 1835 ***''Phenacodus primaevus'' Cope, 1873 ***''Phenacodus teilhardi'' Simpson, 1929 ***''Phenacodus trilobatus'' Cope, 1882 ***''Phenacodus vortmani'' (Cope, 1880) ** Genus '' Lophocion'' Wang and Tong 1997 ***''Lophocion asiaticus'' Wang and Tong 1997 ***''Lophocion grangeri'' Bai et al. 2019 * Subfamiliy Meniscotheriinae Cope, 1882 ** Genus ''
Meniscotherium ''Meniscotherium'' is an extinct genus of dog-sized mammal which lived 54–38 million years ago. It was a herbivore and had hooves. Fossils have been found in Utah, New Mexico. and Colorado. Many individuals have been found together, indicating ...
'' Cope, 1874 ***''Meniscotherium chamense'' Cope, 1874 ***''Meniscotherium tapiacitum'' Cope, 1882 ** Genus ''
Orthaspidotherium ''Orthaspidotherium'' was a European Paleocene genus of early herbivorous mammals of the family Pleuraspidotheriidae. It was included in the family Meniscotheriidae by Teilhard de Chardin Pierre Teilhard de Chardin ( (); 1 May 1881 – 10 ...
'' Lemoine, 1878 ***''Orthaspidotherium edwardsi'' Lemoine, 1878 Some
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses have revealed effective relationships between the various groups of "condylarths". One phylogeny suggests there may be close correlations between a
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, ...
containing
proboscidea The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. From ...
ns,
hyracoids Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simila ...
, perissodactyls and phenacodontids and another clade with ''
Microhyus ''Microhyus'' is a genus of true weevils in the beetle family Curculionidae The Curculionidae are a family of weevils, commonly called snout beetles or true weevils. They are one of the largest animal families, with 6,800 genera and 83,000 spe ...
'' and the macroscelidsTaxeopoda, proposed in 1998.monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
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, ...
, the Condylarths are better understood as an
evolutionary grade A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit. Definition An evolutionary grade is a group of ...
that lead to the true
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, cam ...
s. Indeed, recent phylogenetic studies confirm that phenacodonts were most closely related to modern
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) ...
s.


Paleobiology

The specialized teeth found in at least some phenacodontids seem to indicate a primary
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 ...
lifestyle. The shape of the legs indicated that some phenacodontids (like ''Phenacodus'') were swift runners.


See also

* '' Radinskya'', a basal perissodactyl from the Paleocene of China


Notes


References

* * * * * * {{Taxonbar, from=Q3772948 Condylarths Paleocene mammals Eocene mammals Paleocene first appearances Eocene extinctions Prehistoric mammal families