| Pliopithecoidea | |
|---|---|
| Epipliopithecus skull | |
| Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Primates |
| Suborder: | Haplorhini |
| Parvorder: | Catarrhini |
| Superfamily: | †Pliopithecoidea |
| Families, subfamilies & unplaced genera | |
Dionysopithecidae | |
Pliopithecoidea is an extinct superfamily ofcatarrhineprimates that inhabited Asia and Europe during theMiocene.[1][2] Although they were once a widespread and diverse group of primates, the pliopithecoids have no living descendants.
The first fossil specimens attributed to Pliopithecoidea were discovered byÉdouard Lartet in Sansan, France in 1837. These fossils were later referenced byHenri Marie Ducrotay de Blainville in 1839, who named the type speciesPliopithecus 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.[1]
In the mid-twentieth century, paleontologistsJohannes Hürzeler andHelmuth Zapfe reinvigorated interest in the pliopithecoids with a series of publications in which they named a number of new species, includingPliopithecus 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 theHylobatidae lineage.[3]
With the discovery of more European pliopithecoid fossils in the mid to late 1970s,[4][5] and subsequent discovery of pliopithecoid fossils in China,[6] the idea that pliopithecoids were ancestral to gibbons fell out of favor. Today, most paleontologists agree that pliopithecoids hold a basal position in thecatarrhine family tree.[1][2][7] As such, pliopithecoids represent something similar to the common ancestor ofOld World monkeys andapes.
Afemur discovered inEppelsheim and given the genus namePaidopithex was for many years controversial, as its large size compared to Pliopithecoids led to suggestions that it was instead related to theDryopithecini. 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 discoveredDryopithecus laietanus 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.[8]
A worn tooth found nearHaritalyangar 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.[9] However, two recently discovered molars in the same area appear to support this, with placement within the superfamily uncertain (but clearly not Crouzeliinae).[10]
The pliopithecoid fossil record mostly consists of teeth with a few mandibular and maxillary fragments.[1][2] The dental formula (2.1.2.3) and shape of the teeth are the primary factors which include pliopithecoids among thecatarrhini. Although some authors have argued that the narrow upper molars and broad upper molars of pliopithecoids demonstrate their affinity with modern catarrhines,[11] others have demonstrated that these traits are variable between species.[1] In fact, pliopithecoids are more similar toNew World monkeys in some aspects of their dentition, including narrow lower incisors (mesiodistally waisted towards the base of the crown).[1][12] 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.[1][13] 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.[1]
The crania ofP. vindobonesis,Laccopithecus robustus,Pliopithecus zhanxiangi, andAnapithecus hernyaki demonstrate that pliopithecoids had relatively large and globular braincases with a projecting snout.[1][14] 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.[1] Most importantly, however, pliopithecoids had an incompletely ossified ectotympanic tube. This anatomical feature represents an intermediate stage between what is found inplatyrrhines, which do not have an ossified ectotympanic tube, andcatarrhines, which have a completely ossified ectotympanic tube.[1][2][14]
Nearly all of what is known about the body proportions and post-cranial morphology of this family are derived fromPliopithecus vindobonensis, as it is the only species for which a complete skeleton has been found.[3] 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).[15] Köhler estimates a slightly higher average weight of 10 kg.[8] Post-cranially, pliopithecoids had an interesting mix of platyrrhine and catarrhine traits. The brachial index ofP. 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.[1] Proportionally, however, the forelimbs ofP. vindobonesis were shorter than their hindlimbs, making them comparable to a baboon. The hands and feet ofP. vindobonesis were long and curved, suggesting that pliopithecoids were adept and agile climbers.[1][3] The post-crania ofP. vindobonesis also shows that Pliopithecoids had an entepicondylar foramen, which is a primitive trait not found in any other catarrhine primates (extant or extinct).[1][2][14] 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.[13] Pliopithecoids also had a tail.[1][14]
The following classification scheme represents multiple sources.[citation needed]
Begun divides Pliopithecoidea into two - FamilyDionysopithecidae and FamilyPliopithecidae, with thePliopithecidae sub-divided intoSubfamiliesPliopithecinae andCrouzeliinae.[9]