Editing Pierolapithecus

{{short description|Extinct species of ape from Miocene Europe}}
{{Speciesbox
| fossil_range = [[Miocene]], {{fossilrange|12.5|13}}
| image = Pierolapithecus catalaunicus (Kopie).jpg
| image_alt = Reconstruction of "Pierolapithecus catalaunicus"
| image_caption = ''Pierolapithecus catalaunicus''
| status = 
| status_system = 
| status_ref = 
| genus = Pierolapithecus
| species = catalaunicus
| parent_authority = [[Salvador Moyà-Solà|Moyà-Solà]] et al., [[2004 in paleontology|2004]]
| authority = [[Salvador Moyà-Solà|Moyà-Solà]] et al., [[2004 in paleontology|2004]]
| synonyms = 
| synonyms_ref = 
| subdivision_ranks = 
| subdivision = 
}}

'''''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.<ref>{{Cite web | title = 'Original' great ape discovered | url = http://news.bbc.co.uk/1/hi/sci/tech/4014351.stm | first = Paul | last = Rincon | publisher = BBC | date = 18 November 2004 | access-date = 23 January 2013}}</ref> Others suggest it being a [[Ponginae|pongine]],<ref name="Pérez de los Ríos-2012" /> or a [[Dryopithecini|dryopith]].<ref name="Sevim-Erol-2023" /> On 16 October 2023, scientists reported the facial reconstruction of the great ape.<ref name="Johnson">{{cite news |last=Johnson |first=Mark |title=Scientists reconstructed the face of a 12 million-year-old great ape - Fossils of the extinct species Pierolapithecus catalaunicus may reveal clues about our origins |url=https://www.washingtonpost.com/science/2023/10/24/ancient-ape-face-reconstructed-fossils-landfill/ |date=24 October 2023 |newspaper=[[The Washington Post]] |url-status=live |archiveurl=https://archive.today/20231024143144/https://www.washingtonpost.com/science/2023/10/24/ancient-ape-face-reconstructed-fossils-landfill/ |archivedate=24 October 2023 |accessdate=25 October 2023 }}</ref><ref name="Pugh">{{cite journal |author=Pugh, Kelsey D. |display-authors=et al. |title=The reconstructed cranium of Pierolapithecus and the evolution of the great ape face |date=16 October 2023 |journal=[[Proceedings of the National Academy of Sciences of the United States of America|PNAS]] |volume=120 |issue=44 |pages=e2218778120 |doi=10.1073/pnas.2218778120 |pmid=37844214 |doi-access=free |pmc=10622906 |bibcode=2023PNAS..12018778P }}</ref>

== History ==
The [[splanchnocranium]] was discovered in 2002 and systematic excavations took place during May and June 2003.<ref name="Casanovasvilar-2008">{{Cite journal |last1=Casanovasvilar |first1=I |last2=Alba |first2=D |last3=Moyasola |first3=S |last4=Galindo |first4=J |last5=Cabrera |first5=L |last6=Garces |first6=M |last7=Furio |first7=M |last8=Robles |first8=J |last9=Kohler |first9=M |last10=Angelone |first10=C |date=2008 |title=Biochronological, taphonomical, and paleoenvironmental background of the fossil great ape Pierolapithecus catalaunicus (Primates, Hominidae) |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248408000948 |journal=Journal of Human Evolution |language=en |volume=55 |issue=4 |pages=589–603 |doi=10.1016/j.jhevol.2008.05.004|pmid=18691737 |bibcode=2008JHumE..55..589C |url-access=subscription }}</ref> The species was described by a team of Spanish [[paleoanthropologist]]s 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]])<ref name="Bezanson-2016">{{Cite book |url=https://onlinelibrary.wiley.com/doi/book/10.1002/9781119179313 |title=The International Encyclopedia of Primatology |date=2016-06-14 |publisher=Wiley |isbn=978-0-470-67337-9 |editor-last=Bezanson |editor-first=Michele |edition=1 |language=en |doi=10.1002/9781119179313.wbprim0216 |editor-last2=MacKinnon |editor-first2=Katherine C |editor-last3=Riley |editor-first3=Erin |editor-last4=Campbell |editor-first4=Christina J |editor-last5=Nekaris |editor-first5=K.A.I (Anna) |editor-last6=Estrada |editor-first6=Alejandro |editor-last7=Di Fiore |editor-first7=Anthony F |editor-last8=Ross |editor-first8=Stephen |editor-last9=Jones-Engel |editor-first9=Lisa E}}</ref>), discovered in December 2002. The finding was first reported in the [[Academic journal|journal]] [[Science (journal)|''Science'']] on November 19, 2004. The skeleton is of an adult male individual, composed of 83 bones that make up the splanchnocranium, both [[Maxilla|maxillae]], a complete set of cheek teeth, both canines, a right central incisor, [[Zygomatic bone|zygomatics]], [[Lacrimal bone|lacrimals]], a partial [[Frontal bone|frontal]], [[Carpal bones|carpals]], [[Metacarpal bones|metacarpals]], [[Phalanges|manual phalanges]] from two hands, [[Tarsus (skeleton)|tarsals]], [[Metatarsal bones|metatarsals]], pedal phalanges, right [[Patella|patellar]] distal [[epiphysis]], a left [[Radius (bone)|radius]], some long bone [[Diaphysis|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.<ref name="MoyaSola2004">{{Cite journal |last1=Moya-Sola |first1=S. |last2=Köhler |first2=M. |last3=Alba |first3=D. M. |last4=Casanovas-Vilar |first4=I. |last5=Galindo |first5=J. |year=2004 |title=''Pierolapithecus catalaunicus'', a New Middle Miocene Great Ape from Spain |url=http://doc.rero.ch/record/14752/files/PAL_E1879.pdf |journal=Science |volume=306 |issue=5700 |pages=1339–1344 |bibcode=2004Sci...306.1339M |doi=10.1126/science.1103094 |pmid=15550663 |s2cid=129576842}}</ref>

== Description ==
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 [[Nasal bone|nasals]] are flat and sit beneath the lower rims of the [[Orbit (anatomy)|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 cavity|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-[[Lumbar vertebrae|lumbars]], the [[Pedicles of the vertebral arches|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 [[Accessory bone|os centrale]] are unfused, the [[Triquetral bone|triquetrum]] is small and non-articulating with the [[Ulnar styloid process|ulnar styloid]], and the crevice inserting [[Meniscus (anatomy)|meniscus]] attachment and [[pisiform facet]] is distally shifted.<ref name="MoyaSola2004" />

The holotypic individual is estimated to have weighed 30 kg (66.13 lbs).<ref name="Bezanson-2016" />

== Locomotion ==
Overall, the [[adduction]] and [[supination]] capacity of the wrist, specially built thorax, [[Scapula|scapular]] shift to the back (which was inferred through the long, [[chimpanzee]]-like clavicles), and stiff lumbar vertebrae suggest that positional behavior and [[Orthograde posture|orthograde]] locomotion were emphasized. This type of movement is diagnostic for all extant [[ape]]s including [[human]]s, 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|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 [[Convergent evolution|convergence]] later and independently in the ape lineage.<ref name="MoyaSola2004" />
[[File:Pierolapithecus patella.png|thumb|248x248px|Patella of the holotype ''P. catalaunicus'']]
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.<ref name="Almécija-2009">{{Cite journal |last1=Almécija |first1=Sergio |last2=Alba |first2=David M. |last3=Moyà-Solà |first3=Salvador |date=2009 |title=Pierolapithecus and the functional morphology of Miocene ape hand phalanges: paleobiological and evolutionary implications |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248409000554 |journal=Journal of Human Evolution |language=en |volume=57 |issue=3 |pages=284–297 |doi=10.1016/j.jhevol.2009.02.008|pmid=19631964 |bibcode=2009JHumE..57..284A |url-access=subscription }}</ref> Although, the latter remains disputed.<ref>{{Cite journal |last1=Alba |first1=David M. |last2=Almécija |first2=Sergio |last3=Moyà-Solà |first3=Salvador |date=2010-07-01 |title=Locomotor inferences in Pierolapithecus and Hispanopithecus: Reply to Deane and Begun (2008) |url=https://www.sciencedirect.com/science/article/pii/S0047248410000266 |journal=Journal of Human Evolution |volume=59 |issue=1 |pages=143–149 |doi=10.1016/j.jhevol.2010.02.002 |pmid=20510436 |bibcode=2010JHumE..59..143A |issn=0047-2484|url-access=subscription }}</ref>

The patella was like extant apes in dimensions, which is typically regarded as having a mobile knee. ''Pierolapithecus'' differs from monkeys, [[hylobatid]]s, and basal [[hominoid]]s through thicker patellae. As such, a derived knee might be related to enhanced climbing, notably vertical climbing.<ref>{{Cite journal |last1=Pina |first1=Marta |last2=Almécija |first2=Sergio |last3=Alba |first3=David M. |last4=O'Neill |first4=Matthew C. |last5=Moyà-Solà |first5=Salvador |date=2014 |title=The Middle Miocene Ape Pierolapithecus catalaunicus Exhibits Extant Great Ape-Like Morphometric Affinities on Its Patella: Inferences on Knee Function and Evolution |journal=PLOS ONE |language=en |volume=9 |issue=3 |pages=e91944 |doi=10.1371/journal.pone.0091944 |issn=1932-6203 |pmc=3956854 |pmid=24637777 |bibcode=2014PLoSO...991944P |doi-access=free }}</ref> 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.<ref>{{Cite journal |last1=Hammond |first1=Ashley S. |last2=Alba |first2=David M. |last3=Almécija |first3=Sergio |last4=Moyà-Solà |first4=Salvador |date=2013 |title=Middle Miocene Pierolapithecus provides a first glimpse into early hominid pelvic morphology |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248413000742 |journal=Journal of Human Evolution |language=en |volume=64 |issue=6 |pages=658–666 |doi=10.1016/j.jhevol.2013.03.002|pmid=23545221 |bibcode=2013JHumE..64..658H |url-access=subscription }}</ref>

== Classification ==
''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-[[Hominidae|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.<ref name="MoyaSola2004" /> The large amount of homoplasy in ape locomotion creates considerable [[Taxonomy (biology)|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.<ref name="MoyaSola2004" /> These early traits would have been maintained, overlaid, and modified to suite new adaptations that occurred independently.<ref name="Almécija-2009" />

''Pierolapithecus'' and ''[[Anoiapithecus]]'' may be synonymous with ''Dryopithecus'',<ref name="Begun2015"/> but both have enough craniofacial differences to justify separation.<ref name="Alba-2010"/> Placement as a basal hominid is supported by the comparison of the thick enamel from the [[Afropithecidae|afropithecids]], which may be ancestral to apes.<ref name="Alba-2010">{{Cite journal |last1=Alba |first1=D. M. |last2=Fortuny |first2=J. |last3=Moyà-Solà |first3=S. |date=2010 |title=Enamel thickness in the Middle Miocene great apes Anoiapithecus , Pierolapithecus and Dryopithecus |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=277 |issue=1691 |pages=2237–2245 |doi=10.1098/rspb.2010.0218 |issn=0962-8452 |pmc=2880156 |pmid=20335211}}</ref> Other hypotheses are that the taxon represents a stem pongine.<ref name="Pérez de los Ríos-2012">{{Cite journal |last1=Pérez de los Ríos |first1=Miriam |last2=Moyà-Solà |first2=Salvador |last3=Alba |first3=David M. |date=2012-09-01 |title=The nasal and paranasal architecture of the Middle Miocene ape Pierolapithecus catalaunicus (primates: Hominidae): Phylogenetic implications |url=https://www.sciencedirect.com/science/article/pii/S0047248412000978 |journal=Journal of Human Evolution |volume=63 |issue=3 |pages=497–506 |doi=10.1016/j.jhevol.2012.05.012 |pmid=22819226 |bibcode=2012JHumE..63..497P |issn=0047-2484|url-access=subscription }}</ref> Rather than a full common ancestor, it has been suggested that the species may be ancestral to humans, chimpanzees and [[gorilla]]s but not [[orangutan]]s, given certain facial characteristics.<ref name="Begun2015">{{cite book |last1=Begun |first1=D. R. |title=Fossil record of Miocene hominoids. |date=2015 |publisher=Springer |isbn=978-3-642-39980-0 |edition=2nd |location=Berlin |pages=1304–1306 |ref=3}}</ref> This genus is distinguished from pongines and share traits with extant [[Homininae|hominines]], suggesting a sister relationship with ''Dryopithecus'' (possibly in a tribe called Dryopithecini, having thick and thin enamel much like ''[[Ardipithecus]]''/[[Australopithecine|australopiths]] and ''[[Pan (genus)|Pan]]''/hominins).<ref name="Sevim-Erol-2023">{{Cite journal |last1=Sevim-Erol |first1=Ayla |last2=Begun |first2=D. R. |last3=Sözer |first3=Ç Sönmez |last4=Mayda |first4=S. |last5=van den Hoek Ostende |first5=L. W. |last6=Martin |first6=R. M. G. |last7=Alçiçek |first7=M. Cihat |date=2023-08-23 |title=A new ape from Türkiye and the radiation of late Miocene hominines |journal=Communications Biology |language=en |volume=6 |issue=1 |page=842 |doi=10.1038/s42003-023-05210-5 |issn=2399-3642 |pmc=10447513 |pmid=37612372}}</ref> A reconstruction of the face of ''Pierolapithecus catalaunicus'' indicates its morphology as most consistent with a phylogenetic placement as a [[Crown group#Stem groups|stem]] [[Hominidae|hominid]].<ref>{{Cite journal|last1=Pugh |first1=K. D. |last2=Catalano |first2=S. A. |last3=Pérez de los Ríos |first3=M. |last4=Fortuny |first4=J. |last5=Shearer |first5=B. M. |last6=Vecino Gazabón |first6=A. |last7=Hammond |first7=A. S. |last8=Moyà-Solà |first8=S. |last9=Alba |first9=D. M. |last10=Almécija |first10=S. |year=2023 |title=The reconstructed cranium of ''Pierolapithecus'' and the evolution of the great ape face |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=120 |issue=44 |at=e2218778120 |doi=10.1073/pnas.2218778120 |pmid=37844214 |pmc=10622906 |bibcode=2023PNAS..12018778P }}</ref>

== Paleoecology ==
''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.<ref name="Bezanson-2016" /><ref name="Alba-2010" /> It was discovered at the locality of BCV1,<ref>{{cite journal| last1=Hammond |first1=AS |last2=Alba |first2=DM |last3=Almécija |first3=S |last4=Moyà-Solà |first4=S |year=2013 |title=Middle Miocene hominid ''Pierolapithecus'' provides insight into early hominid pelvic morphology |journal=Paleoanthropology |volume=2013 |issue=2013 |page=A16 |url=https://paleoanthropology.org/ojs/index.php/paleo/article/view/706/667 |issn=1545-0031}}</ref> which formed when the northwestern [[Mediterranean Sea|Mediterranean]] rifted to form a stretch between two mountain ranges. The proximal to distal-marginal [[Alluvium|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 [[Suidae|suid]], followed by the discovery of ''Dryopithecus fontani'', ''[[Hispanopithecus laietanus]]'', and ''[[Sivapithecus|Sivapithecus occidentalis]]'' in the area. From the area hailing ''Pierolapithecus'' specifically was explored from the 1950s&ndash;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).<ref name="Casanovasvilar-2008" />

The [[fauna]] comprises the ''Pierolapithecus'', [[Megaherbivore|megaherbivores]] like elephants, and various others like carnivores, [[Artiodactyl|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 [[Insectivore|insectivores]], arboreal [[Dormouse|dormice]], and [[Flying squirrel|flying squirrels]] support a humid environment, and the open woodlands of other sites would have made hominid occupation impossible.<ref name="Casanovasvilar-2008" />

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.<ref>{{cite journal |last1=Roegl |first1=Fred |date=1999 |title=Mediterranean and Paratethys. Facts and hypotheses of an Oligocene to Miocene paleogeography (short overview) |journal=Geol. Carpathica |volume=50 |pages=339–349}}</ref>

''Pierolapithecus'', like many other Miocene apes, is predicted to have been [[Polygyny|polygynous]] by its low second-to-fourth digit ratios, which are reflective of high prenatal androgen effects and correlated with polygyny in apes.<ref>{{Cite journal |last=Nelson |first=Emma |last2=Rolian |first2=Campbell |last3=Cashmore |first3=Lisa |last4=Shultz |first4=Susanne |date=3 November 2010 |title=Digit ratios predict polygyny in early apes, Ardipithecus , Neanderthals and early modern humans but not in Australopithecus |url=https://royalsocietypublishing.org/doi/10.1098/rspb.2010.1740 |journal=[[Proceedings of the Royal Society B: Biological Sciences]] |language=en |volume=278 |issue=1711 |pages=1556–1563 |doi=10.1098/rspb.2010.1740 |issn=0962-8452 |pmc=3081742 |pmid=21047863 |access-date=10 July 2024}}</ref>

== See also ==
{{Portal|Paleontology|Primates
}}
{{columnslist|colwidth=25em|
* {{annotated link|Anoiapithecus|''Anoiapithecus''}}
* {{annotated link|Chororapithecus|''Chororapithecus''}}
* {{annotated link|Griphopithecus|''Griphopithecus''}}
* {{annotated link|Nakalipithecus|''Nakalipithecus''}}
* {{annotated link|Samburupithecus|''Samburupithecus''}}
}}

==References==
{{Reflist}}

== External links ==
{{Commons category|Pierolapithecus catalaunicus|''Pierolapithecus catalaunicus''}}
* [http://news.bbc.co.uk/1/hi/sci/tech/4014351.stm BBC news: 'Original' great ape discovered]
* [https://www.science.org/doi/abs/10.1126/science.1103094 Research article and comments]

{{Haplorhini|Ho.}}
{{Taxonbar|from1=Q21446402|from2=Q132728}}
[[Category:Hominidae]]
[[Category:Prehistoric mammals of Europe]]
[[Category:Prehistoric apes]]
[[Category:Miocene primates of Europe]]
[[Category:Transitional fossils]]
[[Category:Fossil taxa described in 2004]]
[[Category:Monotypic prehistoric primate genera]]