Editing Paranthropus (section)

==Taxonomy==
===Species===
{{Multiple image|align=right|direction=vertical|width=150|image1=Original of Paranthropus robustus Face.jpg|image2=Australophithecus boisei (cast), Olduvai Gorge - Springfield Science Museum - Springfield, MA - DSC03368.JPG|image3=Paranthropus aethiopicus.JPG|footer=From top to bottom, ''P. robustus'' ([[SK 48]]), ''P. boisei'' ([[OH 5]]) and ''P. aethiopicus'' ([[KNM WT 17000]])}}
<mapframe text="Selected Paranthropus sites from the [https://www.roceeh.uni-tuebingen.de/roadweb ROAD database] (CC BY-SA 4.0 ROCEEH)" width="250", height="300">
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====''P. robustus''====
The genus ''Paranthropus'' was first erected by Scottish-South African [[palaeontologist]] [[Robert Broom]] in 1938, with the [[type species]] ''[[Paranthropus robustus|P. robustus]]''.<ref name=Broom1938/> "''Paranthropus''" derives from [[Ancient Greek]] παρα ''para'' beside or alongside; and άνθρωπος ''ánthropos'' man.<ref>{{cite web|url=https://www.merriam-webster.com/dictionary/Paranthropus|title=''Paranthropus''|publisher=Merriam–Webster Dictionary|access-date=19 December 2019}}</ref> The [[type specimen]], a male braincase, [[TM 1517]], was discovered by schoolboy Gert Terblanche at the [[Kromdraai fossil site]], about {{cvt|70|km}} southwest of [[Pretoria]], South Africa.<ref name=Broom1938>{{cite journal|first=R.|last=Broom|author-link=Robert Broom|year=1938|title=The Pleistocene Anthropoid Apes of South Africa|journal=Nature|volume=142|issue=3591|pages=377–379|doi=10.1038/142377a0|bibcode=1938Natur.142..377B|doi-access=free}}</ref> By 1988, at least six individuals were unearthed in around the same area, now known as the [[Cradle of Humankind]].<ref name=Constantino2004/>

In 1948, at [[Swartkrans]] Cave, in about the same vicinity as Kromdraai, Broom and South African palaeontologist [[John Talbot Robinson]] described ''P. crassidens'' based on a subadult jaw, SK 6. He believed later ''Paranthropus'' were morphologically distinct from earlier ''Paranthropus'' in the cave—that is, the Swartkrans ''Paranthropus'' were [[reproductively isolated]] from Kromdraai ''Paranthropus'' and the former eventually [[speciate]]d.<ref>{{cite journal|first=R.|last=Broom|year=1948|title=Another new type of fossil ape-man|journal=Nature|volume=162|issue=4132|page=57|doi=10.1038/163057a0|pmid=18106151|s2cid=4126221|doi-access=free}}</ref> By 1988, several specimens from Swartkrans had been placed into ''P. crassidens''. However, this has since been synonymised with ''P. robustus'' as the two populations do not seem to be very distinct.<ref name=Constantino2004/>

====''P. boisei''====
In 1959, ''[[P. boisei]]'' was discovered by [[Mary Leakey]] at [[Olduvai Gorge]], [[Tanzania]] (specimen [[OH 5]]). Her husband [[Louis Leakey|Louis]] named it ''Zinjanthropus boisei'' because he believed it differed greatly from ''Paranthropus'' and ''Australopithecus''. The name derives from "[[Zanj|Zinj]]", an ancient Arabic word for the coast of East Africa, and "boisei", referring to their financial benefactor [[Charles Watson Boise]].<ref name=Leakey1959>{{cite journal|first=L.|last=Leakey|author-link=Louis Leakey|year=1959|title=A New Fossil Skull from Olduvai|journal=Nature|volume=184|issue=4685|pages=491–493|url=https://www.academia.edu/616307|doi=10.1038/184491a0|bibcode=1959Natur.184..491L|s2cid=4217460}}</ref> However, this genus was rejected at Mr. Leakey's presentation before the 4th Pan-African Congress on Prehistory, as it was based on a single specimen.<ref>{{cite book|last=Morell|first=V.|year=2011|title=Ancestral Passions: The Leakey Family and the Quest for Humankind's Beginnings|publisher=Touchstone|page=193|isbn=978-1-4391-4387-2}}</ref> The discovery of the [[Peninj Mandible]] made the Leakeys reclassify their species as ''Australopithecus (Zinjanthropus) boisei'' in 1964,<ref>{{cite journal|last1=Leakey|first1=L. S. B.|author-link=Louis Leakey|last2=Leakey|first2=M. B.|author-link2=Mary Leakey|year=1964|title=Recent discoveries of fossil hominidsin Tanganyika, at Olduvai and near Lake Natron|journal=Nature|volume=202|issue=4927|pages=5–7|doi=10.1038/202005a0|pmid=14166721|bibcode=1964Natur.202....5L|s2cid=4162123}}</ref> but in 1967, South African palaeoanthropologist [[Phillip V. Tobias]] subsumed it into ''Australopithecus'' as ''A. boisei''. However, as more specimens were found, the combination ''Paranthropus boisei'' became more popular.<ref>{{cite journal|first=B.|last=Wood|year=2005|title=A tale of two taxa|journal=Transactions of the Royal Society of South Africa|volume=60|issue=2|pages=91–94|doi=10.1080/00359190509520483|bibcode=2005TRSSA..60...91W |s2cid=83659439}}</ref>

It is debated whether the wide range of variation in jaw size indicates simply [[sexual dimorphism]] or a grounds for identifying a new species. It could be explained as [[groundmass]] filling in cracks naturally formed after death, inflating the perceived size of the bone.<ref>{{cite journal|first1=N.|last1=Silverman|first2=B.|last2=Richmond|first3=B.|last3=Wood|year=2001|title=Testing the taxonomic integrity of ''Paranthropus boisei sensu stricto''|journal=American Journal of Physical Anthropology|volume=115|issue=2|pages=167–178|doi=10.1002/ajpa.1066|pmid=11385603}}</ref><ref name=Constantino2007/><ref name=Wood2007/> ''P. boisei'' also has a notably wide range of variation in skull anatomy, but these features likely have no taxonomic bearing.<ref>{{cite journal|first1=B.|last1=Wood|first2=D.|last2=Lieberman|author-link2=Daniel Lieberman|year=2001|title=Craniodental variation in ''Paranthropus boisei'': a developmental and functional perspective|journal=American Journal of Physical Anthropology|volume=116|issue=1|pages=13–25|doi=10.1002/ajpa.1097|pmid=11536113|url=https://scholar.harvard.edu/files/dlieberman/files/2001a.pdf}}</ref>

====''P. aethiopicus''====
[[File:Paranthropus Africa.jpg|thumb|Locations of ''Paranthropus'' finds]]
In 1968, French palaeontologists [[Camille Arambourg]] and [[Yves Coppens]] described "''Paraustralopithecus aethiopicus''" based on a toothless mandible from the [[Shungura Formation]], Ethiopia (Omo 18).<ref>{{cite journal|last1=Arambourg|first1=C.|author-link=Camille Arambourg|first2=Y.|last2=Coppens|author-link2=Yves Coppens|year=1968|title=Sur la decouverte dans le Pleistocene inferieur de la valle de l'Omo (Ethiopie) d'une mandibule d'Australopithecien|language=fr|trans-title=On the discovery in the Lower Pleistocene Omo Valley (Ethiopia) of an Australopithecine Mandible|journal=Comptes Rendus des Séances de l'Académie des Sciences|volume=265|pages=589–590}}</ref> In 1976, American anthropologist [[Francis Clark Howell]] and Breton anthropologist [[Yves Coppens]] reclassified it as ''A. africanus''.<ref name=Ferguson1989/> In 1986, after the discovery of the skull [[KNM WT 17000]] by English anthropologist [[Alan Walker (anthropologist)|Alan Walker]] and [[Richard Leakey]] classified it into ''Paranthropus'' as ''[[P. aethiopicus]]''.<ref>{{cite journal|first1=A.|last1=Walker|author-link=Alan Walker (anthropologist)|first2=R. E.|last2=Leakey|author2-link=Richard Leakey|first3=J. M.|last3=Harris|first4=F. H.|last4=Brown|year=1986|title=2.5-Myr Australopithecus boisei from west of Lake Turkana, Kenya|journal=Nature|volume=322|issue=6079|pages=517–522|doi=10.1038/322517a0|bibcode=1986Natur.322..517W|s2cid=4270200}}</ref> There is debate whether this is synonymous with ''P. boisei'',<ref name=Constantino2007/> the main argument for separation being the skull seems less adapted for chewing tough vegetation.<ref name=Wood2007/><ref name=Wood2000/>

In 1989, palaeoartist and zoologist [[Walter Ferguson]] reclassified KNM WT 17000 into a new species, ''walkeri'', because he considered the skull's species designation questionable as it comprised the skull whereas the holotype of ''P. aethiopicus'' comprised only the mandible.<ref name=Ferguson1989>{{cite journal|first=W. W.|last=Ferguson|author-link=Walter Ferguson|year=1989|title=A New Species of the Genus ''Australopithecus'' (Primates: Hominidae) from Plio/Pleistocene Deposits West of Lake Turkana in Kenya|journal=Primates|volume=30|issue=2|pages=223–232|doi=10.1007/BF02381307|s2cid=28642451}}</ref> Ferguson's classification is almost universally ignored,<ref>{{cite book|url=https://books.google.com/books?id=440TmWXToLAC&pg=PT298|first=B.|last=Wood|title=Wiley-Blackwell Encyclopedia of Human Evolution|publisher=John Wiley & Sons|pages=298–299|isbn=978-1-4443-4247-5|year=2011}}</ref> and is considered to be synonymous with ''P. aethiopicus''.<ref>{{cite book|first1=R.|last1=Leakey|author-link=Richard Leakey|first2=R.|last2=Lewin|author-link2=Roger Lewin|year=1993|title=Origins Reconsidered: In Search of what Makes Us Human|url=https://books.google.com/books?id=KQULA9F9--IC&pg=PA362|publisher=Anchor Books|pages=132–133|isbn=978-0-385-46792-6}}</ref>

====Others====

In 2015, Ethiopian palaeoanthropologist [[Yohannes Haile-Selassie]] and colleagues described the 3.5–3.2 Ma ''[[A. deyiremeda]]'' based on three jawbones from the [[Afar Region]], Ethiopia. They noted that, though it shares many similarities with ''Paranthropus'', it may not have been closely related because it lacked enlarged molars which characterize the genus.<ref>{{cite journal|first1=Y.|last1=Haile-Selassie|author-link=Yohannes Haile-Selassie|first2=L.|last2=Gilbert|first3=S. M.|last3=Melillo|display-authors=et al.|year=2015|title=New species from Ethiopia further expands Middle Pliocene hominin diversity|journal=Nature|volume=521|issue=14448|pages=483–488|doi=10.1038/nature14448|pmid=26017448|url=https://afanporsaber.com/wp-content/uploads/2017/09/New-species-from-Ethiopia-further-expands-Middle-Pliocene-hominin-diversity.pdf|bibcode=2015Natur.521..483H|s2cid=4455029}}</ref> Nonetheless, in 2018, independent researcher Johan Nygren recommended moving it to ''Paranthropus'' based on dental and presumed dietary similarity.<ref name=Nygren>{{cite journal|first=J.|last=Nygren|year=2018|title=The speciation of ''Australopithecus'' and ''Paranthropus'' was caused by introgression from the ''Gorilla'' lineage|journal=PeerJ Preprints|volume=6|page=e27130v3|url=https://www.biorxiv.org/content/biorxiv/early/2018/10/05/434894.full.pdf|doi=10.7287/peerj.preprints.27130v3|bibcode=2018arXiv180806307N|arxiv=1808.06307|s2cid=52054499 |doi-access=free }}</ref>

===Validity===
In 1951, American anthropologists [[Sherwood Washburn]] and [[Bruce D. Patterson]] were the first to suggest that ''Paranthropus'' should be considered a [[junior synonym]] of ''Australopithecus'' as the former was only known from fragmentary remains at the time, and dental differences were too minute to serve as justification.<ref>{{cite journal|first1=S. L.|last1=Washburn|author-link=Sherwood Washburn|first2=B. D.|last2=Patterson|year=1951|title=Evolutionary Importance of the South African 'Man-apes'|journal=Nature|volume=167|issue=4251|pages=650–651|doi=10.1038/167650a0|pmid=14826894|bibcode=1951Natur.167..650W|s2cid=4207075}}</ref> In face of calls for subsumation, Leakey<ref name=Leakey1959/> and Robinson<ref>{{cite journal|first=J. T.|last=Robinson|author-link=John Talbot Robinson|year=1965|title=''Homo 'habilis''{{'}} and the Australopithecines|journal=Nature|volume=205|issue=4967|pages=121–124|doi=10.1038/205121a0|bibcode=1965Natur.205..121R|s2cid=4196031}}</ref> continued defending its validity. Various other authors were still unsure until more complete remains were found.<ref name=Constantino2004>{{cite book|first1=P. J.|last1=Constantino|first2=B. A.|last2=Wood|year=2004|chapter=''Paranthropus'' Paleobiology|title=Miscelanea en Homenaje a Emiliano Aguirre|volume=III|series=Paleoantropologia|publisher=Museo Arqueológico Regional|url=https://mds.marshall.edu/cgi/viewcontent.cgi?article=1028&=&context=bio_sciences_faculty&=&sei-redir=1&referer=https%253A%252F%252Fscholar.google.com%252Fscholar%253Fhl%253Den%2526as_sdt%253D0%25252C44%2526q%253Dparanthropus%252Baethiopicus%2526oq%253DParanthropus#search=%22paranthropus%20aethiopicus%22}}</ref> ''Paranthropus'' is sometimes classified as a [[subgenus]] of ''Australopithecus''.<ref>{{cite journal|last1=Cela-Conde|first1=C. J.|author-link=Camilo José Cela Conde|last2=Ayala|first2=F. J.|author-link2=Francisco J. Ayala|year=2003|title=Genera of the human lineage|journal=Proceedings of the National Academy of Sciences|volume=100|issue=13|pages=7684–7689|bibcode=2003PNAS..100.7684C|doi=10.1073/pnas.0832372100|pmc=164648|pmid=12794185|doi-access=free}}</ref>

[[File:Paranthropus robustus top (University of Zurich).JPG|thumb|upright|''P. robustus'' ([[SK 48]])]]
There is currently no clear consensus on the validity of ''Paranthropus''. The argument rests upon whether the genus is [[monophyletic]]—is composed of a common ancestor and all of its descendants—and the argument against monophyly (that the genus is [[paraphyletic]]) says that ''P. robustus'' and ''P. boisei'' evolved similar gorilla-like heads independently of each other by coincidence ([[convergent evolution]]), as chewing adaptations in hominins evolve very rapidly and multiple times at various points in the family tree ([[homoplasy]]).<ref name=Wood2007>{{cite journal|first1=B.|last1=Wood|first2=J.|last2=Constantino|year=2007|title=''Paranthropus boisei'': Fifty Years of Evidence and Analysis|journal=Yearbook of Physical Anthropology|volume=50|pages=106–132|doi=10.1002/ajpa.20732|pmid=18046746|url=https://mds.marshall.edu/bio_sciences_faculty/37|doi-access=free}}</ref> In 1999, a chimp-like [[ulna]] forearm bone was assigned to ''P. boisei'', the first discovered ulna of the species, which was markedly different from ''P. robustus'' ulnae, which could suggest paraphyly.<ref name=McHenry2007>{{cite journal|first1=H. M.|last1=McHenry|author-link=Henry McHenry (anthropologist)|first2=C. C.|last2=Brown|first3=L. J.|last3=McHenry|year=2007|title=Fossil hominin ulnae and the forelimb of ''Paranthropus''|journal=American Journal of Physical Anthropology|volume=134|issue=2|pages=209–218|doi=10.1002/ajpa.20656|pmid=17596856}}</ref>

===Evolution===
''P. aethiopicus'' is the earliest member of the genus, with the oldest remains, from the Ethiopian [[Omo Kibish Formation]], dated to 2.6 mya at the end of the [[Pliocene]]. It is sometimes regarded as the direct ancestor of ''P. boisei'' and ''P. robustus''.<ref name=Constantino2007/> It is possible that ''P. aethiopicus'' evolved even earlier, up to 3.3 mya, on the expansive Kenyan floodplains of the time.<ref name=Joordens2019>{{cite journal|first1=J. C. A.|last1=Joordens|first2=C. S.|last2=Feibel|first3=H. B.|last3=Vonhof|first4=A. S.|last4=Schulp|first5=D.|last5=Kroon|year=2019|title=Relevance of the eastern African coastal forest for early hominin biogeography|journal=Journal of Human Evolution|volume=131|pages=176–202|pmid=31182201|doi=10.1016/j.jhevol.2019.03.012|doi-access=free|bibcode=2019JHumE.131..176J |hdl=20.500.11820/6c1ee960-79ba-45df-9e12-3350c768a497|hdl-access=free}}</ref> The oldest ''P. boisei'' remains date to about 2.3 mya from [[Malema]], Malawi.<ref name=Constantino2007>{{cite journal|first1=P. J.|last1=Constantino|first2=B. A.|last2=Wood|year=2007|title=The Evolution of ''Zinjanthropus boisei''|journal=Evolutionary Anthropology|volume=16|issue=2|pages=49–62|doi=10.1002/evan.20130|s2cid=53574805|url=https://mds.marshall.edu/cgi/viewcontent.cgi?article=1027&context=bio_sciences_faculty|url-access=subscription}}</ref> ''P. boisei'' changed remarkably little over its nearly one-million-year existence.<ref>{{cite journal|first1=B.|last1=Wood|first2=C.|last2=Wood|first3=L.|last3=Konigsberg|year=1994|title=''Paranthropus boisei'': an example of evolutionary stasis?|journal=American Journal of Physical Anthropology|volume=95|issue=2|pages=117–136|doi=10.1002/ajpa.1330950202|pmid=7802091}}</ref> ''Paranthropus'' had spread into South Africa by 2 mya with the earliest ''P. robustus'' remains.<ref name=Wood2000>{{cite journal|first1=B.|last1=Wood|first2=B. G.|last2=Richmond|year=2000|title=Human evolution: taxonomy and paleobiology|journal=Journal of Anatomy|volume=192|issue=Pt 1 |pages=34–38<!--only citing these pages-->|pmc=1468107|pmid=10999270|doi=10.1046/j.1469-7580.2000.19710019.x}}</ref><ref name=Stammers2018/><ref name=Herries2009/>

It is sometimes suggested that ''Paranthropus'' and ''Homo'' are [[sister taxa]], both evolving from ''[[Australopithecus]]''. This may have occurred during a drying trend 2.8–2.5 mya in the [[Great Rift Valley]], which caused the retreat of woodland environments in favor of open savanna, with forests growing only along rivers and lakes. ''Homo'' evolved in the former, and ''Paranthropus'' in the latter [[riparian]] environment.<ref name=Joordens2019/><ref>{{cite journal|first1=O.|last1=Kullmer|first2=O.|last2=Sandrock|first3=F.|last3=Schrenk|first4=T. G.|last4=Bromage|year=1999|title=The Malawi Rift: Biogeography, Ecology and Coexistence of Homo and Paranthropus|journal=Anthropologie|volume=37|issue=3|pages=221–231|jstor=26294888}}</ref><ref>{{cite journal|first1=R.|last1=Bobe|first2=A. K.|last2=Behrensmeyer|first3=R. E.|last3=Chapman|title=Faunal change, environmental variability and late Pliocene hominin evolution|journal=Journal of Human Evolution|volume=42|issue=4|pages=475–497|doi=10.1006/jhev.2001.0535|pmid=11908957|year=2002|bibcode=2002JHumE..42..475B |s2cid=26032638}}</ref> However, the classifications of ''Australopithecus'' species is problematic.<ref name=Parins2019/>

[[File:Paranthropus boisei face (University of Zurich).JPG|thumb|upright|''P. boisei'' [[OH 5]]]]
[[Evolutionary tree]] according to a 2019 study:<ref name=Parins2019>{{cite journal|first1=C.|last1=Parins-Fukuchi|first2=E.|last2=Greiner|first3=L. M.|last3=MacLatchy|first4=D. C.|last4=Fisher|year=2019|title=Phylogeny, ancestors and anagenesis in the hominin fossil record|journal=Paleobiology|volume=45|issue=2|pages=378–393|doi=10.1017/pab.2019.12|bibcode=2019Pbio...45..378P |s2cid=196659329|url=https://www.biorxiv.org/content/biorxiv/early/2018/10/05/434894.full.pdf}}</ref>
{{clade| style=font-size:85%;line-height:85%
|label1=[[Hominini]]
|1={{clade
 |1=[[Chimpanzee]]
 |label2=''[[Sahelanthropus]]''
 |2={{clade
  |1={{clade
   |1=''[[Ardipithecus]]''
   |label2=''[[A. anamensis]]''
   |2=''[[A. afarensis]]''
   }}
  |2={{clade
   |label1='''''Paranthropus'''''
   |1={{clade
    |1=''[[P. aethiopicus]]''
    |2={{clade
     |1=''[[P. boisei]]''
     |2=''[[Paranthropus robustus|P. robustus]]''
     }}
    }}
   |2={{clade
    |label2=''[[A. garhi]]''
    |1=''[[Australopithecus africanus|A. africanus]]''
    |2={{clade
     |1=''[[H. floresiensis]]''
     |2={{clade
      |1=''[[A. sediba]]''
      |2={{clade
       |1=''[[H. habilis]]''
       |2=Other ''[[Homo]]''
       }}
      }}
     }}
    }}
}}}}}}}}