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| Pierolapithecus | |
|---|---|
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| Pierolapithecus catalaunicus | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Primates |
| Suborder: | Haplorhini |
| 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 ofprimate which lived around 12.5-13 million years ago during theMiocene in what is nowHostalets de Pierola,Catalonia,Spain. Some researchers believe that it is a candidate forcommon ancestor to thegreat ape clade, or is at least closer than any previous fossil discovery.[1] Others suggest it being apongine,[2] or adryopith.[3] On 16 October 2023, scientists reported the facial reconstruction of the great ape.[4][5]
Thesplanchnocranium was discovered in 2002 and systematic excavations took place during May and June 2003.[6] The species was described by a team of Spanishpaleoanthropologists led bySalvador Moyà-Solà on the basis of afossil skeleton,IPS21350 (nicknamedPau ("peace" in Catalan as it was announced alongside Spanish demonstrations against theIraq War)[7]), discovered in December 2002. The finding was first reported in thejournalScience on November 19, 2004. The skeleton is of an adult male individual, composed of 83 bones that make up the splanchnocranium, bothmaxillae, a complete set of cheek teeth, both canines, a right central incisor,zygomatics,lacrimals, a partialfrontal,carpals,metacarpals,manual phalanges from two hands,tarsals,metatarsals, pedal phalanges, rightpatellar distalepiphysis, a leftradius, some long bonediaphyses, 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-Solàet 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, thenasals are flat and sit beneath the lower rims of theorbit, theglabella is posteriorly oriented, the face is low, the brows are thin, the zygomatic root is high, and thenasoalveolar clivus is high. The rear border of theincisive foramen is in line with the P3, thepalate is deep and stout, thenasal aperture is widest at the base, theinterorbital 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 athorax that is anteroposteriorly compressed, theclavicle is robust, lacking a ventral keel on the mid-lumbars, thepedicles of the neural arch are robust and stout, thespinous processes are slightly caudally inclined, the pedicle-body inserts thetransverse processes, dorsally oriented and pedicle-born transverse process, the metacarpals and phalanges are short, theos centrale are unfused, thetriquetrum is small and non-articulating with theulnar styloid, and the crevice insertingmeniscus attachment andpisiform facet is distally shifted.[8]
The holotypic individual is estimated to have weighed 30 kg (66.13 lbs).[7]
Overall, theadduction andsupination 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 andorthograde locomotion were emphasized. This type of movement is diagnostic for all extantapes includinghumans, but it is rarely documented in the fossil record. Other hominids that have this suite areOreopithecus and, although less skeletally complete,Dryopithecus. Earlier taxa—Proconsul,Afropithecus,Equatorius,Nacholapithecus—retain basal characters and the similarly-agedMorotopithecus practiced orthograde locomotion but was probably sister to apes (based on facial structure). The shortened phalanges suggest ancestralpalmigrade adaptations, but it is unlikelyPierolapithecus practiced much or anysuspensory behavior. However, vertical climbing and suspension are independent abilities that are integral to ape evolution. Below-branch suspension may have evolved repeatedly or inconvergence 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 thisgenus is related to a retainedpronograde 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 basalhominoids through thicker patellae. As such, a derived knee might be related to enhanced climbing, notably vertical climbing.[11] The pelvis shares an ancestral template withProconsul nyanzae, which was modified for orthograde behavior (assuming that hypothesis is accepted), and suggestshomoplasy in ape pelves.[12]
Pierolapithecus demonstrates derived facial features and apelike skeletal adaptations that suggest that it is an early member of the apeclade. 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 considerabletaxonomic confusion. The lateMiddle Miocene is the farthest trace of aPierolapithecus-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 andAnoiapithecus may be synonymous withDryopithecus,[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 theafropithecids, 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 andgorillas but notorangutans, given certain facial characteristics.[13] This genus is distinguished from pongines and share traits with extanthominines, suggesting a sister relationship withDryopithecus (possibly in a tribe called Dryopithecini, having thick and thin enamel much likeArdipithecus/australopiths andPan/hominins).[3] A reconstruction of the face ofPierolapithecus catalaunicus indicates its morphology as most consistent with a phylogenetic placement as astemhominid.[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 northwesternMediterranean rifted to form a stretch between two mountain ranges. The proximal to distal-marginalalluvial-fan sediments cover the Miocene. It was discovered as a fossiliferous area by Guerín in the 1920s with an ape M2 mistaken as asuid, followed by the discovery ofDryopithecus fontani,Hispanopithecus laietanus, andSivapithecus occidentalis in the area. From the area hailingPierolapithecus specifically was explored from the 1950s–1970s from a garbage dump. 19 large and small mammals were discovered at the site, almost 300macroinvertebrate fossils, and 83 hominid fossils (composing the holotype skeleton).[6]
Thefauna comprises thePierolapithecus,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. Themicromammals show signs ofdigestion by predators, probably by barn owls and others. The environment was quite humid, warm, and forested. The fauna is most likeFrance andcentral Europe in composition. Moreinsectivores, arborealdormice, andflying squirrels support a humid environment, and the open woodlands of other sites would have made hominid occupation impossible.[6]
ThatPierolapithecus would be ancestral to modern great apes is debated largely because this great ape was found in theIberian Peninsula, while most of the fossil evidence of the evolution of hominids and hominins has been located inEast Africa andSoutheast Asia. Because the Mediterranean Sea contracted several times in the past, migration of terrestrial fauna betweenAfrica andEurope was permitted.[17]
Pierolapithecus, like many other Miocene apes, is predicted to have beenpolygynous by its low second-to-fourth digit ratios, which are reflective of high prenatal androgen effects and correlated with polygyny in apes.[18]
