Plesiadapis
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''Plesiadapis'' is one of the oldest known
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians ( monkeys and apes, the latter includin ...
-like
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 ...
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 Adapiformes is a group of early primates. Adapiforms radiated throughout much of the northern continental mass (now Europe, Asia and North America), reaching as far south as northern Africa and tropical Asia. They existed from the Eocene to the ...
primate of the Eocene period, ''
Adapis ''Adapis'' is an extinct adapiform primate from the Eocene of Europe. While this genus has traditionally contained five species (''A. magnus, A. bruni, A. collinsonae, A. parisiensis,'' and ''A. sudrei''), recent research has recognized at least ...
''. ''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 ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
probably arose in North America and colonized Europe on a landbridge via
Greenland Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an island country in North America that is part of the Kingdom of Denmark. It is located between the Arctic and Atlantic oceans, east of the Canadian Arctic Archipelago. Greenland ...
. 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 continental sediments and in the correlation of faunas on both sides of the Atlantic. Two remarkable
skeletons A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure insid ...
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 that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
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 itself the lone extant family within the infraorder Tarsiiformes. Although the group was once more widespread, all of its species living today are found in Maritime Southeas ...
-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 ''Archicebus'' is a genus of fossil primates that lived in the early Eocene forests (~55.8–54.8 million years ago) of what is now Jingzhou in the Hubei Province in central China, discovered in 2003. The only known species, ''A. achil ...
'' 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 ''lemures'' – ghosts or spirits) are wet-nosed primates of the superfamily Lemuroidea (), divided into 8 families and consisting of 15 genera and around 100 existing species. They are endemic to the island of Madagas ...
, 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 Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth, in mammalian oral anatomy People with the surn ...
, three
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s, and three
molars 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 ...
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, but others are exclusively arboreal. The habitats pose nu ...
lifestyle in the trees. The
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, 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 o ...
of ''P. gidleyi'' is similar to that of the
eastern gray squirrel The eastern gray squirrel (''Sciurus carolinensis''), also known, particularly outside of North America, as simply the grey squirrel, is a tree squirrel in the genus ''Sciurus''. It is native to eastern North America, where it is the most prodi ...
, though lacking the strong spines. The
humerus The humerus (; ) 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 and ulna, and consists of three sections. The humeral upper extremity consists of a r ...
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 omnivority and herbivority. 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 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 ...
. 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, but others are exclusively arboreal. The habitats pose nu ...
quadruped Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
. Remains found showed that it had a body mass of around .


References


External links


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