Pierolapithecus Temporal range: Miocene, | |
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Pierolapithecus catalaunicus | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Suborder: | Haplorhini |
Infraorder: | Simiiformes |
Family: | Hominidae |
Subfamily: | incertae sedis |
Genus: | † Pierolapithecus Moyà-Solà et al., 2004 |
Species: | †P. catalaunicus |
Binomial name | |
†Pierolapithecus catalaunicus | |
Pierolapithecus catalaunicus is an extinct species of primate which lived around 12.5-13 million years ago during the Miocene in what is now Hostalets de Pierola, Catalonia, Spain. Some researchers believe that it is a candidate for common ancestor to the great ape clade, or is at least closer than any previous fossil discovery. [1] Others suggest it being a pongine, [2] or a dryopith. [3] On 16 October 2023, scientists reported the facial reconstruction of the great ape. [4] [5]
The splanchnocranium was discovered in 2002 and systematic excavations took place during May and June 2003. [6] The species was described by a team of Spanish paleoanthropologists led by Salvador Moyà-Solà on the basis of a fossil skeleton, IPS21350 (nicknamed Pau ("peace" in Catalan as it was announced alongside Spanish demonstrations against the Iraq War) [7] ), discovered in December 2002. The finding was first reported in the journal Science on November 19, 2004. The skeleton is of an adult male individual, composed of 83 bones that make up the splanchnocranium, both maxillae, a complete set of cheek teeth, both canines, a right central incisor, zygomatics, lacrimals, a partial frontal, carpals, metacarpals, manual phalanges from two hands, tarsals, metatarsals, pedal phalanges, right patellar distal epiphysis, a left radius, some long bone diaphyses, two pelvic pieces, three vertebrae, two intact ribs, and twelve rib fragments of large size. They named their new genus after the nearby village Els Hostalets de Pierola, and Catalonia respectively. [8]
Moyà-Solà et al. initially founded the species on a set of unique characteristics, of which are the following. The frontal processes of the face remain on the same plane, the nasals are flat and sit beneath the lower rims of the orbit, the glabella is posteriorly oriented, the face is low, the brows are thin, the zygomatic root is high, and the nasoalveolar clivus is high. The rear border of the incisive foramen is in line with the P3, the palate is deep and stout, the nasal aperture is widest at the base, the interorbital distance is wide, the zygomatics expand to the side, the P3 is similar in size to the P4, there is reduced cusp heteromorphy, all molars save from the M3 are elongated, the upper molars and premolars lack cingula, the lingual cusps of the upper molars are positioned peripherally, the M2 is large and has cusp heteromorphy, and the upper canine is large and compressed in the crown. The ribs are very curved to form a thorax that is anteroposteriorly compressed, the clavicle is robust, lacking a ventral keel on the mid-lumbars, the pedicles of the neural arch are robust and stout, the spinous processes are slightly caudally inclined, the pedicle-body inserts the transverse processes, dorsally oriented and pedicle-born transverse process, the metacarpals and phalanges are short, the os centrale are unfused, the triquetrum is small and non-articulating with the ulnar styloid, and the crevice inserting meniscus attachment and pisiform facet is distally shifted. [8]
The holotypic individual is estimated to have weighed 30 kg (66.13 lbs). [7]
Overall, the adduction and supination capacity of the wrist, specially built thorax, scapular shift to the back (which was inferred through the long, chimpanzee-like clavicles), and stiff lumbar vertebrae suggest that positional behavior and orthograde locomotion were emphasized. This type of movement is diagnostic for all extant apes including humans, but it is rarely documented in the fossil record. Other hominids that have this suite are Oreopithecus and, although less skeletally complete, Dryopithecus . Earlier taxa— Proconsul , Afropithecus , Equatorius , Nacholapithecus —retain basal characters and the similarly-aged Morotopithecus practiced orthograde locomotion but was probably sister to apes (based on facial structure). The shortened phalanges suggest ancestral palmigrade adaptations, but it is unlikely Pierolapithecus practiced much or any suspensory behavior. However, vertical climbing and suspension are independent abilities that are integral to ape evolution. Below-branch suspension may have evolved repeatedly or in convergence later and independently in the ape lineage. [8]
Further analysis suggests that very long and curved phalanges is decoupled with orthograde features related to vertical climbing being acquired. The condition in this genus is related to a retained pronograde plan. Although the lumbars, ribs, and carpals are orthograde, the degree of this in the phalanges is only slight. Many traits were independently acquired, leading to new advances being superimposed and basal characters retained for an extended time. Pierolapithecus lacked adaptations for suspensory hanging, but it may have still been capable of doing so, only that it was not adaptively relevant. [9] Although, the latter remains disputed. [10]
The patella was like extant apes in dimensions, which is typically regarded as having a mobile knee. Pierolapithecus differs from monkeys, hylobatids, and basal hominoids through thicker patellae. As such, a derived knee might be related to enhanced climbing, notably vertical climbing. [11] The pelvis shares an ancestral template with Proconsul nyanzae, which was modified for orthograde behavior (assuming that hypothesis is accepted), and suggests homoplasy in ape pelves. [12]
Pierolapithecus demonstrates derived facial features and apelike skeletal adaptations that suggest that it is an early member of the ape clade. This genus, 12.5-13 mya in age, postdates the hylobatid-hominid split, which occurred anywhere from 14.9±2 or 14.6±2.6 mya. Much of the skeleton is derived, but the shortened phalanges are indicative of palmigrade adaptations that are primitive. This mosaic is important in ape evolution. [8] The large amount of homoplasy in ape locomotion creates considerable taxonomic confusion. The late Middle Miocene is the farthest trace of a Pierolapithecus-like character group, and assuming that this identifies the earliest apes, is the farthest trace of hominids. As well, early hominids are substantially more primitive than estimated, which may explain why no early great apes were previously reported. [8] These early traits would have been maintained, overlaid, and modified to suite new adaptations that occurred independently. [9]
Pierolapithecus and Anoiapithecus may be synonymous with Dryopithecus, [13] but both have enough craniofacial differences to justify separation. [14] Placement as a basal hominid is supported by the comparison of the thick enamel from the afropithecids, which may be ancestral to apes. [14] Other hypotheses are that the taxon represents a stem pongine. [2] Rather than a full common ancestor, it has been suggested that the species may be ancestral to humans, chimpanzees and gorillas but not orangutans, given certain facial characteristics. [13] This genus is distinguished from pongines and share traits with extant hominines, suggesting a sister relationship with Dryopithecus (possibly in a tribe called Dryopithecini, having thick and thin enamel much like Ardipithecus /australopiths and Pan /hominins). [3] A reconstruction of the face of Pierolapithecus catalaunicus indicates its morphology as most consistent with a phylogenetic placement as a stem hominid. [15]
Pierolapithecus bore thick enamel found in hard-object feeders, but its diet is not yet known aside from possibly having fed in trees like orangutans. [7] [14] It was discovered at the locality of BCV1, [16] which formed when the northwestern Mediterranean rifted to form a stretch between two mountain ranges. The proximal to distal-marginal alluvial-fan sediments cover the Miocene. It was discovered as a fossiliferous area by Guerín in the 1920s with an ape M2 mistaken as a suid, followed by the discovery of Dryopithecus fontani, Hispanopithecus laietanus , and Sivapithecus occidentalis in the area. From the area hailing Pierolapithecus specifically was explored from the 1950s–1970s from a garbage dump. 19 large and small mammals were discovered at the site, almost 300 macroinvertebrate fossils, and 83 hominid fossils (composing the holotype skeleton). [6]
The fauna comprises the Pierolapithecus, megaherbivores like elephants, and various others like carnivores, artiodactyls, turtles, and small-medium fragments. Pierolapithecus bears evidence of scavenging whereas the other fossils show signs of being scattered across an alluvial plain. The micromammals show signs of digestion by predators, probably by barn owls and others. The environment was quite humid, warm, and forested. The fauna is most like France and central Europe in composition. More insectivores, arboreal dormice, and flying squirrels support a humid environment, and the open woodlands of other sites would have made hominid occupation impossible. [6]
That Pierolapithecus would be ancestral to modern great apes is debated largely because this great ape was found in the Iberian Peninsula, while most of the fossil evidence of the evolution of hominids and hominins has been located in East Africa and Southeast Asia. Because the Mediterranean Sea contracted several times in the past, migration of terrestrial fauna between Africa and Europe was permitted. [17]
Homininae, also called "African hominids" or "African apes", is a subfamily of Hominidae. It includes two tribes, with their extant as well as extinct species: 1) the tribe Hominini ―and 2) the tribe Gorillini (gorillas). Alternatively, the genus Pan is sometimes considered to belong to its own third tribe, Panini. Homininae comprises all hominids that arose after orangutans split from the line of great apes. The Homininae cladogram has three main branches, which lead to gorillas and to humans and chimpanzees. There are two living species of Panina and two living species of gorillas, but only one extant human species. Traces of extinct Homo species, including Homo floresiensis have been found with dates as recent as 40,000 years ago. Organisms in this subfamily are described as hominine or hominines.
Orrorin is an extinct genus of primate within Homininae from the Miocene Lukeino Formation and Pliocene Mabaget Formation, both of Kenya.
Pongidae, or the pongids is an obsolete primate taxon containing chimpanzees, gorillas and orangutans. By this definition pongids were also called "great apes". This taxon is not used today but is of historical significance. The great apes are currently classified as Hominidae. This entry addresses the old usage of pongid.
Oreopithecus is an extinct genus of hominoid primate from the Miocene epoch whose fossils have been found in today's Tuscany and Sardinia in Italy. It existed nine to seven million years ago in the Tusco-Sardinian area when this region was an isolated island in a chain of islands stretching from central Europe to northern Africa in what was becoming the Mediterranean Sea.
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 being described from single remains based on minute differences amongst each other, and the fragmentary nature of the holotype specimen makes differentiating remains difficult. There is currently only one uncontested species, the type species D. fontani, though there may be more. The genus is placed into the tribe Dryopithecini, which is either an offshoot of orangutans, African apes, or is its own separate branch.
Ekembo nyanzae, originally classed as a species of Proconsul, is a species of fossil primate first discovered by Louis Leakey on Rusinga Island in 1942, which he published in Nature in 1943. It is also known by the name Dryopithecus africanus. A joint publication of Wilfrid Le Gros Clark and Louis Leakey in 1951, "The Miocene Hominoidea of East Africa", first defines Proconsul nyanzae. In 1965 Simons and Pilbeam replaced Proconsul with Dryopithecus, using the same species names.
Ouranopithecus is a genus of extinct Eurasian great ape represented by two species, Ouranopithecus macedoniensis, a late Miocene hominoid from Greece and Ouranopithecus turkae, also from the late Miocene of Turkey.
Indopithecus giganteus is an extinct species of large ape that lived in the late Miocene of the Siwalik Hills in northern India. Although frequently assigned to the more well-known genus Gigantopithecus, recent authors consider it to be a distinct genus in its own right.
Chororapithecus is an extinct great ape from the Afar region of Ethiopia roughly 8 million years ago during the Late Miocene, comprising one species, C. abyssinicus. It is known from 9 isolated teeth discovered in a 2005–2007 survey of the Chorora Formation. The teeth are indistinguishable from those of gorillas in terms of absolute size and relative proportions, and it has been proposed to be an early member of Gorillini. However, this is controversial given the paucity of remains, and notable anatomical differences between Chororapithecus and gorilla teeth. The Kenyan ape Nakalipithecus has been proposed to be an ancestor of Chororapithecus or at least closely related. If correct, they would be the only identified fossil members of any modern non-human great ape lineage, and would push the gorilla–human last common ancestor from 8 million years ago to 10 million years ago. The teeth are adapted for processing tough plant fibres as well as hard, brittle food, and the formation is thought to represent a forested lakeside habitat.
The Hominidae, whose members are known as the great apes or hominids, are a taxonomic family of primates that includes eight extant species in four genera: Pongo ; Gorilla ; Pan ; and Homo, of which only modern humans remain.
Anoiapithecus is an extinct ape genus thought to be closely related to Dryopithecus. Both genera lived during the Miocene, approximately 12 million years ago. Fossil specimens named by Salvador Moyà-Solà are known from the deposits from Spain.
Salvador Moyà-Solà is a Spanish paleontologist. He works in the Institut Català de Paleontología Miquel Crusafont in Sabadell, Catalonia. From 1983 to 2006, he was affiliated with the Diputación Provincial de Barcelona. In 2006 he became investigator of the Physical Anthropology unit of the Universidad Autónoma de Barcelona.
Dryopithecini is an extinct tribe of Eurasian and African great apes that are believed to be close to the ancestry of gorillas, chimpanzees and humans. Members of this tribe are known as dryopithecines.
Hominid dispersals in Europe refers to the colonisation of the European continent by various species of hominid, including hominins and archaic and modern humans.
Nacholapithecus kerioi was an ape that lived 14-15 million years ago during the Middle Miocene. Fossils have been found in the Nachola formation in northern Kenya. The only member of the genus Nacholapithecus, it is thought to be a key genus in early hominid evolution. Similar in body plan to Proconsul, it had a long vertebral column with six lumbar vertebrae, no tail, a narrow torso, large upper limbs with mobile shoulder joints, and long feet.
Graecopithecus is an extinct genus of hominid that lived in southeast Europe during the late Miocene around 7.2 million years ago. Originally identified by a single lower jaw bone bearing teeth found in Pyrgos Vasilissis, Athens, Greece, in 1944, other teeth were discovered from Azmaka quarry in Bulgaria in 2012. With only little and badly preserved materials to reveal its nature, it is considered as "the most poorly known European Miocene hominoids." The creature was popularly nicknamed 'El Graeco' by scientists.
Hispanopithecus is a genus of apes that inhabited Europe during the Miocene epoch. It was first identified in a 1944 paper by J. F. Villalta and M. Crusafont in Notas y Comunicaciones del Instituto Geologico y Minero de España. Anthropologists disagree as to whether Hispanopithecus belongs to the subfamily Ponginae or Homininae.
Pliobates cataloniae is a primate from 11.6 million years ago, during the Iberian Miocene. Originally described as a species of stem-ape that was found to be the sister taxon to gibbons and great apes like humans, it was subsequently reinterpreted as a non-ape catarrhine belonging to the group Crouzeliidae within the superfamily Pliopithecoidea on the basis of discovery of new dental remains with crouzeliid synapomorphies. Its anatomy is gibbon-like; prior to this discovery, it was assumed that the ancestral ape bauplan was robust like Proconsul. This species has mosaic characteristics of primitive, monkey-like features and the more derived ape characteristics; however, even when originally described it wasn't interpreted as a direct ancestor of modern apes but rather a side-branch that retained the ancestral morphotype and was thus placed in its own family Pliobatidae. Its subsequent placement within Pliopithecoidea indicates that it was convergent with apes in elbow and wrist morphology.
Pliopithecoidea is an extinct superfamily of catarrhine primates that inhabited Asia and Europe during the Miocene. Although they were once a widespread and diverse group of primates, the pliopithecoids have no living descendants.