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''Plesiadapis'' is one of the oldest known
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
-like
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 ...
genera which existed about 58–55 million years ago in North America and Europe. ''Plesiadapis'' means "near-Adapis", which is a reference to the adapiform primate of the Eocene period, '' Adapis''. ''Plesiadapis tricuspidens'', the type specimen, is named after the three cusps present on its upper incisors.


Taxonomy

The first discovery of ''Plesiadapis'' was made by François Louis Paul Gervaise in 1877, who first discovered ''Plesiadapis tricuspidens'' in France. The type specimen is MNHN Crl-16, and is a left mandibular fragment dated to the early Eocene epoch. This
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
probably arose in North America and colonized Europe on a land bridge via
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
. Thanks to the abundance of the genus and to its rapid evolution, species of ''Plesiadapis'' play an important role in the zonation of
Late Paleocene The Thanetian is, in the International Commission on Stratigraphy, ICS Geologic timescale, the latest age (geology), age or uppermost stage (stratigraphy), stratigraphic stage of the Paleocene epoch (geology), Epoch or series (stratigraphy), Serie ...
continental sediments and in the correlation of faunas on both sides of the Atlantic. Two remarkable skeletons of ''Plesiadapis'', one of them nearly complete, have been found in lake deposits at Menat, France. Although the preservation of the hard parts is poor, these skeletons still show remains of skin and hair as a carbonaceous film—something unique among
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), ...
mammals. Details of the bones are better preserved in fossils from Cernay, also in France, where ''Plesiadapis'' is one of the most common mammals.


Classification

The following are possible shared derived features of Plesiadapiformes: maxillary-frontal contact in orbit, the presence of a suboptic foramen, an ossified external auditory meatus, the absence of a promontory artery, the absence of a stapedial artery, and a strong mastoid tubercle. Although the placement of the ''Plesiadapis'' lineage is still up for debate, the consensus in the 1970s was that they were closest to early
tarsier Tarsiers ( ) are haplorhine primates of the family Tarsiidae, which is the lone extant family within the infraorder Tarsiiformes. Although the group was prehistorically more globally widespread, all of the existing species are restricted to M ...
-like primates. Plesiadapiformes have also been proposed as a nonprimate sister group to Eocene-Recent primates. A study done in 1987 linked Plesiadapiformes with adapids and omomyids through nine shared-derived features, six of which are cranial or dental: (1) auditory bulla inflated and formed by the petrosal bone, (2) ectotympanic expanded laterally and fused medially to the wall of the bulla, (3) promontorium centrally positioned in the bulla, and large hypotympanic sinus widely separating promontorium from the basisphenoid, (4) internal carotid entering the bulla posteriolaterally and enclosed in a bony tube, (5) nannopithex fold on the upper molars, and (6) loss of one pair of incisors. In 2013, a phylogenetic analysis that includes also the basal primate '' Archicebus'' positions ''Plesiadapis'' firmly outside of the Primates, as a sister group to both Primates and Dermoptera.


Anatomy and remains

''Plesiadapis'' is one of the most completely known early primatomorphs, with a significant amount of the skeleton known. Though, the skeleton is mostly known from ''P. gidleyi'' and the relatively late (derived) ''P. tricuspidens''. The skull is overall reminiscent of a
lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
, though it lacks the postorbital bars (a vertical bar bordering the posterior margin of the eye socket). The brain was probably relatively large compared to similarly sized contemporary mammals, namely the arctocyonids. The dental formula is usually , with two
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
s, one canine, three
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
s, and three
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
in either half of the upper jaw; and one incisor, one canine, three premolars, and one molar in either half of the lower jaw. The incisors are quite long. Already, ''Plesiadapis'' had lost the first premolar from the mammalian common ancestor, but later primatomorphs would lose the second premolar as well. ''P. dubius'' consistently lacks the lower second premolar, and about half of ''P. rex'' specimens lack it too. ''P. gidleyi'' and European ''Plesiadapis'' lack the lower canines. The skeletal adaptations are consistent with a largely
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
lifestyle in the trees. The
sacrum The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part of the pelvic cavity, ...
of ''P. gidleyi'' is similar to that of the eastern gray squirrel, though lacking the strong spines. The
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
is robust and features a strong S-curve. Nearly all of what is known about the anatomy of plesiadapiforms comes from fragmentary jaws and teeth, so most definitions of plesiadapiform genera and species are based on dentition. ''Plesiadapis dentition shows a functional shift toward grinding and crushing in the cheek teeth as an adaptation towards increasing omnivory and herbivory. The skull of ''Plesiadapis'' is relatively broad and flat, with a long snout with rodent-like jaws and teeth and long, gnawing
incisors Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
separated by a gap from its
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
. Orbits are still directed to the side, unlike the forward-facing eyeballs of modern primates that enable three-dimensional vision. ''Plesiadapis'' had mobile limbs that terminated in strongly curved claws, and it sported a long bushy tail which is preserved in the Menat skeletons. The way of life of ''Plesiadapis'' has been much debated in the past. Climbing habits could be expected in a relative of the primates, but tree-dwelling animals are rarely found in such high numbers. Based on this and other evidence, some paleontologists have concluded that these animals were mainly living on the ground, like today's marmots and ground squirrels. However, more recent investigations have confirmed that the skeleton of ''Plesiadapis'' is that of an adept climber, which can be best compared to tree squirrels or to tree-dwelling marsupials such as possums. The short, robust limbs, the long, laterally compressed claws, and the long, bushy tail indicate that it was an
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
quadruped Quadrupedalism is a form of locomotion in which animals have four legs that are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four legs is said to be a quadruped (fr ...
. Remains found showed that it had a body mass of around .


References


External links


Mikko's Phylogeny Archive
{{Taxonbar, from=Q1191111 Paleocene mammals Plesiadapiformes Prehistoric placental genera Paleogene mammals of North America Paleogene mammals of Europe Ypresian extinctions Fossil taxa described in 1877 Eocene mammals