Pliopithecoidea
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Pliopithecoidea is an extinct superfamily of catarrhine
primate Primates are a diverse order (biology), order of mammals. They are divided into the Strepsirrhini, strepsirrhines, which include the lemurs, galagos, and lorisids, and the Haplorhini, haplorhines, which include the Tarsiiformes, tarsiers and ...
s that inhabited Asia and Europe during the
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...
. Although they were once a widespread and diverse group of primates, the pliopithecoids have no living descendants.


History of discovery

The first fossil specimens attributed to Pliopithecoidea were discovered by Édouard Lartet in Sansan, France in 1837. These fossils were later referenced by
Henri Marie Ducrotay de Blainville Henri Marie Ducrotay de Blainville (; 12 September 1777 – 1 May 1850) was a French zoologist and anatomist. Life Blainville was born at Arques, near Dieppe. As a young man he went to Paris to study art, but ultimately devoted himself to nat ...
in 1839, who named the type species '' Pliopithecus antiquus''. A second species, ''Pliopithecus platyodon'', was discovered in Switzerland by Biedermann in 1863. Following this, a small number of other pliopithecoid species were described from fossil collections found in France, Germany, and Poland. In the mid-twentieth century, paleontologists Johannes HĂĽrzeler and
Helmuth Zapfe Helmuth is both a masculine German given name and a surname. Notable people with the name include: Given name; * Helmuth Theodor Bossert (1889–1961), German art historian, philologist and archaeologist *Helmuth Duckadam (born 1959), Romanian form ...
reinvigorated interest in the pliopithecoids with a series of publications in which they named a number of new species, including ''Pliopithecus vindobonensis'', which consists of the most complete cranial and post-cranial pliopithecoid specimens ever discovered. Based on their size, and some superficial similarities to modern day gibbons, Zapfe suggested that pliopithecoids were ancestral to the
Hylobatidae Gibbons () are apes in the Family (biology), family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical rainforest from eastern Bangladesh ...
lineage. With the discovery of more European pliopithecoid fossils in the mid to late 1970s, and subsequent discovery of pliopithecoid fossils in China, the idea that pliopithecoids were ancestral to gibbons fell out of favor. Today, most paleontologists agree that pliopithecoids hold a basal position in the catarrhine family tree. As such, pliopithecoids represent something similar to the common ancestor of
Old World monkey Old World monkey is the common English name for a family of primates known taxonomically as the Cercopithecidae (). Twenty-four genera and 138 species are recognized, making it the largest primate family. Old World monkey genera include baboons ...
s and apes. A
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
discovered in
Eppelsheim Eppelsheim is an ''Ortsgemeinde'' – a Municipalities of Germany, municipality belonging to a ''Verbandsgemeinde'', a kind of collective municipality – in the Alzey-Worms district in Rhineland-Palatinate, Germany. Geography The municipality ...
and given the genus name '' Paidopithex'' was for many years controversial, as its large size compared to Pliopithecoids led to suggestions that it was instead related to the Dryopithecini. A lack of femurs for Dryopithecini meant that the suggestion was not ruled out for many years, but in 2002 work by Köhler et al comparing it to a recently discovered ''
Dryopithecus laietanus ''Dryopithecus'' is a genus of extinct great apes from the middle–late Miocene boundary of Europe 12.5 to 11.1 million years ago (mya). Since its discovery in 1856, the genus has been subject to taxonomic turmoil, with numerous new species b ...
'' skeleton showed that it was very different from the Dryopithecini. However, Köhler felt unable to definitely place Paidopithex in the Pliopithecoid superfamily, stating it was either an unusually large Pliopithecoid (estimated bodyweight 22 kg) or could be the sole known species of a separate superfamily. A worn tooth found near Haritalyangar in India and dated from around 9 to 8 million years ago has been suggested as possibly a Pliopithecoid species, Krishnapithecus krishnai, but the wear has made this difficult to determine. However, two recently discovered molars in the same area appear to support this, with placement within the superfamily uncertain (but clearly not Crouzeliinae).


Physical characteristics

The pliopithecoid fossil record mostly consists of teeth with a few mandibular and maxillary fragments. The dental formula (2.1.2.3) and shape of the teeth are the primary factors which include pliopithecoids among the catarrhini. Although some authors have argued that the narrow upper molars and broad upper molars of pliopithecoids demonstrate their affinity with modern catarrhines, others have demonstrated that these traits are variable between species. In fact, pliopithecoids are more similar to
New World monkey New World monkeys are the five families of primates that are found in the tropical regions of Mexico, Central and South America: Callitrichidae, Cebidae, Aotidae, Pitheciidae, and Atelidae. The five families are ranked together as the Ceboid ...
s in some aspects of their dentition, including narrow lower incisors (mesiodistally waisted towards the base of the crown). Many species have what is often referred to as a 'pliopithecine triangle', a subtle set of ridges defining a small triangular shaped pit between the protocone and hypocone of the lower molars, but even this trait is variable. Instead, the most defining dental trait present in all pliopithecoids is a tall crowned lower third premolar, which is relatively triangular in outline with a comparatively short, vertically oriented mesiobucal face. The crania of ''P. vindobonesis'', ''Laccopithecus robustus'', ''Pliopithecus zhanxiangi'', and ''Anapithecus hernyaki'' demonstrate that pliopithecoids had relatively large and globular braincases with a projecting snout. The snout projects less than the propliopithecoids of North Africa (''i.e.'' '' Aegyptopithecus''), suggesting some prognathic reduction from the inferred common ancestor of these two primate families. The orbits are widely spaced and the mandible is long and robust, with a relatively broad ramus. Most importantly, however, pliopithecoids had an incompletely ossified ectotympanic tube. This anatomical feature represents an intermediate stage between what is found in platyrrhines, which do not have an ossified ectotympanic tube, and catarrhines, which have a completely ossified ectotympanic tube. Nearly all of what is known about the body proportions and post-cranial morphology of this family are derived from ''Pliopithecus vindobonensis'', as it is the only species for which a complete skeleton has been found. Still, the majority of fossil material indicates that pliopithecoids were medium sized primates, approximately the size of a howler monkey or a gibbon (8 kg). Köhler estimates a slightly higher average weight of 10 kg. Post-cranially, pliopithecoids had an interesting mix of platyrrhine and catarrhine traits. The brachial index of ''P. vindobonesis'' (the length of the radius divided by the length of the humerus) is similar to that of a howler monkey, but the crural index (the length of the tibia divided by the length of the femur) is similar to that of a gibbon. Proportionally, however, the forelimbs of ''P. vindobonesis'' were shorter than their hindlimbs, making them comparable to a baboon. The hands and feet of ''P. vindobonesis'' were long and curved, suggesting that pliopithecoids were adept and agile climbers. The post-crania of ''P. vindobonesis'' also shows that Pliopithecoids had an entepicondylar foramen, which is a primitive trait not found in any other catarrhine primates (extant or extinct). The wrist and hands of pliopithecoids were seemingly much more similar to platyrrhines than to catarrhines, as the carpo-metacarpal joint of the thumb is a modified “hinge joint” compared to the "saddle-like" thumb joint found in Old World monkeys and apes. Pliopithecoids also had a tail.


Classification

The following classification scheme represents multiple sources. *Order
Primate Primates are a diverse order (biology), order of mammals. They are divided into the Strepsirrhini, strepsirrhines, which include the lemurs, galagos, and lorisids, and the Haplorhini, haplorhines, which include the Tarsiiformes, tarsiers and ...
s (Linnaeus, 1758) **Infraorder Catarrhini (Geoffroy Saint-Hilaire, 1812) ***Superfamily Pliopithecoidea (Zapfe, 1960) ****Family Pliopithecidae (Zapfe, 1960) *****Subfamily Dionysopithecinae (Li, 1978) ******Genus '' Dionysopithecus'' (Li, 1978) *******''Dionysopithecus shuangoeuensis'' (Li, 1978) *******''Dionysopithecus orientalis'' (Suteethorn, 1990) ******Genus '' Platodontopithecus'' (Li, 1978) *******''Platodontopithecus jianghuaiensis'' (Gu and Lin, 1983) *****Subfamily Pliopithecinae (Zapfe, 1960) ******Genus '' Pliopithecus'' (Gervais, 1849) *******''Pliopithecus antiquus'' (Blainville, 1839) *******''Pliopithecus piveteaui'' (Hürzeler, 1954) *******''Pliopithecus platydon'' (Biederman, 1863) *******''Pliopithecus zhanxiangi'' (Harrison, Delson, and Guan, 1991) *******''Pliopithecus bii'' (Yu, Jin, Jie 2003) *******''Pliopithecus canmatensis'' (Alba, Moyá-Solá, Robles, Galindo, 2012) ******Genus ''
Epipliopithecus ''Epipliopithecus vindobonensis'' is an extinct species of pliopithecoid primate recovered from the Middle Miocene deposits of Devínska Nová Ves fissure in western Slovakia. ''Epipliopithecus'' is one of the few pliopithecoids for which both cr ...
'' (Zapfe and HĂĽrzeler, 1957) *******''
Epipliopithecus vindobonensis ''Epipliopithecus vindobonensis'' is an extinct species of pliopithecoid primate recovered from the Middle Miocene deposits of Devínska Nová Ves fissure in western Slovakia. ''Epipliopithecus'' is one of the few pliopithecoids for which both cr ...
'' (Zapfe and Hürzeler, 1957) *****Subfamily Crouzeliinae (Ginsburg, 1975) ******Tribe Crouzeliini (Ginsburg, 1975) *******Genus '' Plesiopliopithecus'' (Zapfe, 1961) ********''Pleisopliopithecus auscitanensis'' (Ginsburg, 1975) ********''Pleisopliopithecus rhondanica'' (Ginsburg and Mein, 1980) ********''Pleisopliopithecus lockeri'' (Zapfe, 1961) ********''Pleisopliopithecus priensis'' (Welcomme, Aguilar, and Ginsburg, 1991) ******Tribe Anapithecini (Kretzoi, 1975) *******Genus '' Anapithecus'' (Kretzoi, 1975) ********''Anapithecus hernyaki'' (Kretzoi, 1975) *******Genus '' Laccopithecus'' (Wu & Pan, 1984) ********''Laccopithecus robustus'' (Wu and Pan, 1984) *******Genus '' Barberapithecus'' (Alba and Moyá-Solá, 2012) ********''Barberapithecus huerzeleri'' (Alba and Moyá-Solá, 2012) *******''
Genus Egarapithecus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
'' (Moyá-Solá, Köhler, and Alba, 2001) ********''Egarapithecus narcisoi'' (Moyá-Solá, Köhler, and Alba, 2001) *****''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'' ******Genus '' Paidopithex'' (Pohlig, 1895) ******Genus '' Krishnapithecus'' ******Genus '' Kapi'' Begun divides Pliopithecoidea into two - Family
Dionysopithecidae Dionysopithecidae is an extinct family of fossil catarrhines and the earliest-known and most primitive members of the Pliopithecoidea Pliopithecoidea is an extinct superfamily of catarrhine primates that inhabited Asia and Europe during the M ...
and Family Pliopithecidae, with the Pliopithecidae sub-divided into Subfamilies Pliopithecinae and Crouzeliinae.


Notes


References

{{Taxonbar, from=Q10350553 Prehistoric primates Catarrhini Mammal superfamilies Miocene first appearances Miocene extinctions