Editing Paranthropus

{{good article}}
{{short description|Contested extinct genus of hominins}}
{{Italic title}}
{{Automatic taxobox
| name = ''Paranthropus''
| fossil_range = [[Piacenzian]]–[[Chibanian]], {{fossilrange|2.9|1.2}}
| image = Smac Paläolithikum 013.jpg
| image_caption = Skull of ''[[P. boisei]]'' (MGL 95211)
| display_parents = 3
| taxon = Paranthropus
| authority = [[Robert Broom|Broom]], 1938
| type_species = †''[[Paranthropus robustus]]''
| type_species_authority = [[Robert Broom|Broom]], 1938
| subdivision_ranks = [[Species]]
| subdivision = *†''[[P. aethiopicus]]''
*†''[[Paranthropus robustus|P. robustus]]''
*†''[[P. boisei]]''
*†''[[Paranthropus capensis|P. capensis]]''
| synonyms = *''Paraustralopithecus''<br/><small>[[Camille Arambourg|Arambourg]] and [[Yves Coppens|Coppens]], 1968</small>
*''Zinjanthropus''<br/><small>[[Mary Leakey|Leakey]], 1959</small>
}}

'''''Paranthropus''''' is a [[genus]] of extinct [[hominin]] which contains two widely accepted [[species]]: ''[[Paranthropus robustus|P. robustus]]'' and ''[[P. boisei]]''. However, the validity of ''Paranthropus'' is contested, and it is sometimes considered to be [[synonym (taxonomy)|synonymous]] with ''[[Australopithecus]]''. They are also referred to as the '''robust australopithecines'''. They lived between approximately 2.9 and 1.2 million years ago (mya) from the end of the [[Pliocene]] to the [[Middle Pleistocene]].

''Paranthropus'' is characterised by robust [[skulls]], with a prominent [[gorilla]]-like [[sagittal crest]] along the midline—which suggest strong chewing muscles—and broad, [[herbivorous]] teeth used for grinding. However, they likely preferred soft food over tough and hard food. Typically, ''Paranthropus'' species were generalist feeders, but while ''P. robustus'' was likely an [[omnivore]], ''P. boisei'' seems to have been herbivorous, possibly preferring abundant [[bulbotuber]]s. Paranthropoids were [[bipeds]]. Despite their robust heads, they had comparatively small bodies. Average weight and height are estimated to be {{cvt|40|kg}} at {{cvt|132|cm|ft|0}} for ''P. robustus'' males, {{cvt|50|kg}} at {{cvt|137|cm|ftin|0}} for ''P. boisei'' males, {{cvt|32|kg}} at {{cvt|110|cm|ftin|0}} for ''P. robustus'' females, and {{cvt|34|kg}} at {{cvt|124|cm|ftin|0}} for ''P. boisei'' females.

They were possibly [[polygamous]] and [[patrilocal]], but there are no modern analogues for australopithecine societies. They are associated with bone tools and contested as the earliest evidence of fire usage. They typically inhabited woodlands, and coexisted with some early human species, namely ''[[Australopithecus africanus|A. africanus]]'', ''[[H. habilis]]'' and ''[[H. erectus]]''. They were preyed upon by the large carnivores of the time, specifically crocodiles, leopards, [[sabertoothed cat]]s and hyenas.

==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]]''
       }}
      }}
     }}
    }}
}}}}}}}}

==Description==
===Skull===
''Paranthropus'' had a massively built, tall and flat skull, with a prominent [[gorilla]]-like [[sagittal crest]] along the midline which anchored large [[temporalis muscle]]s used in chewing.<ref name=Wood2004/> Like other australopithecines, ''Paranthropus'' exhibited sexual dimorphism, with males notably larger than females.<ref name=Wood2000/><ref name=McHenry1991a/><ref name=McHenry1991b/> They had large [[molar tooth|molars]] with a relatively thick [[tooth enamel]] coating ([[post-canine megadontia]]),<ref>{{cite journal|first1=A. J.|last1=Olejniczak|first2=T. M.|last2=Smith|first3=M. M.|last3=Skinner|display-authors=et al.|year=2008|title=Three-dimensional molar enamel distribution and thickness in ''Australopithecus'' and ''Paranthropus''|journal=Biology Letters|volume=4|issue=4|pages=406–410|doi=10.1098/rsbl.2008.0223|pmid=18522924|pmc=2610159}}</ref> and comparatively small [[incisor]]s (similar in size to modern [[human]]s),<ref>{{cite journal|first1=P. S.|last1=Ungar|first2=F. E.|last2=Grine|year=1991|title=Incisor size and wear in ''Australopithecus africanus'' and ''Paranthropus robustus''|journal=Journal of Human Evolution|volume=20|issue=4|pages=313–340|doi=10.1016/0047-2484(91)90013-L|bibcode=1991JHumE..20..313U }}</ref> possibly adaptations to processing abrasive foods.<ref name=Williams2015>{{cite journal|first=F. L.|last=Williams|year=2015|title=Dietary proclivities of ''Paranthropus robustus'' from Swartkrans, South Africa|journal=Anthropological Review|volume=78|issue=1|pages=1–19|doi=10.1515/anre-2015-0001|doi-access=free}}</ref><ref name=Wood2012>{{cite journal|first1=B.|last1=Wood|first2=K.|last2=Schroer|year=2012|title=Reconstructing the Diet of an Extinct Hominin Taxon: The Role of Extant Primate Models|journal=International Journal of Primatology|volume=33|issue=3|pages=716–742|doi=10.1007/s10764-012-9602-7|s2cid=15983306}}</ref> The teeth of ''P. aethiopicus'' developed faster than those of ''P. boisei''.<ref>{{cite journal|first=F. V.|last=Ramirez-Rozzi|year=1993|title=Tooth development in East African ''Paranthropus''|journal=Journal of Human Evolution|volume=24|issue=6|pages=429–454|doi=10.1006/jhev.1993.1030|bibcode=1993JHumE..24..429R }}</ref>

''Paranthropus'' had adaptations to the skull to resist large bite loads while feeding, namely the expansive [[squamosal suture]]s.<ref>{{cite journal|first1=C.|last1=Dzialo|first2=S. A.|last2=Wood|first3=M.|last3=Berthaume|display-authors=et al.|year=2013|title=Functional implications of squamosal suture size in ''Paranthropus boisei''|journal=American Journal of Physical Anthropology|volume=153|issue=2|pages=260–268|doi=10.1002/ajpa.22427|pmid=24242913}}</ref> The notably thick [[palate]] was once thought to have been an adaptation to resist a high bite force, but is better explained as a byproduct of facial lengthening and nasal anatomy.<ref>{{cite journal|first=M. A.|last=McCollum|year=1998|title=Palatal thickening and facial form in ''Paranthropus'': Examination of alternative developmental models|journal=American Journal of Physical Anthropology|volume=103|issue=3|pages=375–392|doi=10.1002/(SICI)1096-8644(199707)103:3<375::AID-AJPA7>3.0.CO;2-P|pmid=9261500|doi-access=free}}</ref>

In ''P. boisei'', the [[temporomandibular joint|jaw hinge]] was adapted to grinding food side-to-side (rather than up-and-down in modern humans), which is better at processing the [[starch]]y abrasive foods that likely made up the bulk of its diet. ''P. robustus'' may have chewed in a front-to-back direction instead, and had less exaggerated (less [[synapomorphy and apomorphy|derived]]) anatomical features than ''P. boisei'' as it perhaps did not require them with this kind of chewing strategy. This may have also allowed ''P. robustus'' to better process tougher foods.<ref>{{cite journal|first1=K.|last1=Kupczik|first2=V.|last2=Toro-Ibacache|first3=G. A.|last3=Macho|year=2018|title=On the relationship between maxillary molar root shape and jaw kinematics in ''Australopithecus africanus'' and ''Paranthropus robustus''|journal=Royal Society Open Science|volume=5|issue=8|page=180825|doi=10.1098/rsos.180825|pmc=6124107|pmid=30225074|bibcode=2018RSOS....580825K}}</ref>

The braincase volume averaged about {{cvt|500|cm3}}, comparable to gracile australopithecines, but smaller than ''Homo''.<ref name=Du2018>{{cite journal|first1=A.|last1=Du|first2=A. M.|last2=Zipkin|first3=K. G.|last3=Hatala|display-authors=et al.|year=2018|title=Pattern and process in hominin brain size evolution are scale-dependent|journal=Proceedings of the Royal Society B|volume=285|issue=1873|pages=20172738|doi=10.1098/rspb.2017.2738|pmc=5832710|pmid=29467267}}</ref> Modern human brain volume averages {{cvt|1270|cm3}} for men and {{cvt|1130|cm3}} for women.<ref>{{cite journal|first1=J. S.|last1=Allen|first2=H.|last2=Damasio|first3=T. J.|last3=Grabowski|year=2002|title=Normal neuroanatomical variation in the human brain: an MRI-volumetric study|journal=American Journal of Physical Anthropology|volume=118|issue=4|pages=341–358|doi=10.1002/ajpa.10092|pmid=12124914|s2cid=21705705}}</ref>

===Limbs and locomotion===
Unlike ''P. robustus'', the forearms of ''P. boisei'' were heavily built, which might suggest habitual [[suspensory behaviour]] as in [[orangutan]]s and [[gibbon]]s.<ref name=McHenry2007/><ref name=Dominguez2013/><ref name=Lague2019/> A ''P. boisei'' [[shoulder blade]] indicates long [[infraspinatus muscle]]s, which is also associated with suspensory behavior.<ref>{{cite journal|first1=D. J.|last1=Green|first2=H.|last2=Chirchir|first3=E.|last3=Mbua|year=2018|title=Scapular anatomy of ''Paranthropus boisei'' from Ileret, Kenya|journal=Journal of Human Evolution|volume=125|pages=181–192|doi=10.1016/j.jhevol.2017.06.013|pmid=30502893|doi-access=free|bibcode=2018JHumE.125..181G }}</ref> A ''P. aethiopicus'' ulna, on the other hand, shows more similarities to ''Homo'' than ''P. boisei''.<ref name=Lague2019>{{cite journal|first1=M. R.|last1=Lague|first2=H.|last2=Chirchir|first3=D. J.|last3=Green|first4=E.|last4=Mbua|year=2019|title=Humeral anatomy of the KNM-ER 47000 upper limb skeleton from Ileret, Kenya: Implications for taxonomic identification|journal=Journal of Human Evolution|volume=126|pages=24–38|doi=10.1016/j.jhevol.2018.06.011|pmid=30583842|s2cid=58607106|url=https://www.researchgate.net/publication/329486548|doi-access=free|bibcode=2019JHumE.126...24L }}</ref>

''Paranthropus'' were [[biped]]s, and their hips, legs and feet resemble ''A. afarensis'' and modern humans.<ref>{{cite journal |vauthors=Wood B, Richmond BG |title=Human evolution: taxonomy and paleobiology |journal=Journal of Anatomy |volume=197 |issue=1 |pages=19–60 |date=July 2000 |pmid=10999270 |pmc=1468107 |doi=10.1046/j.1469-7580.2000.19710019.x}}</ref><ref>{{cite journal|first1=T. M.|last1=Ryan|first2=K. J.|last2=Carlson|first3=A. D.|last3=Gordon|display-authors=et al.|year=2018|title=Human-like hip joint loading in ''Australopithecus africanus'' and ''Paranthropus robustus''|journal=Journal of Human Anthropology|volume=121|pages=12–24|doi=10.1016/j.jhevol.2018.03.008|pmid=29706230|s2cid=14060188|url=https://www.repository.cam.ac.uk/handle/1810/277557|doi-access=free|bibcode=2018JHumE.121...12R }}</ref> The pelvis is similar to ''A. afarensis'', but the hip joints are smaller in ''P. robustus''. The physical similarity implies a similar walking gait.<ref>{{cite journal |vauthors=Macchiarelli R, Bondioli L, Galichon V, Tobias PV |title=Hip bone trabecular architecture shows uniquely distinctive locomotor behaviour in South African australopithecines |journal=Journal of Human Evolution |volume=36 |issue=2 |pages=211–32 |date=February 1999 |pmid=10068067 |doi=10.1006/jhev.1998.0267|bibcode=1999JHumE..36..211M }}</ref> Their modern-humanlike big toe indicates a modern-humanlike foot posture and range of motion, but the more distal ankle joint would have inhibited the modern human toe-off [[bipedal gait cycle|gait cycle]]. By 1.8 mya, ''Paranthropus'' and ''[[H. habilis]]'' may have achieved about the same grade of bipedality.<ref>{{cite journal|first1=R. L.|last1=Susman|first2=T. M.|last2=Brain|year=1988|title=New first metatarsal (SKX 5017) from Swartkrans and the gait of ''Paranthropus robustus''|journal=American Journal of Physical Anthropology|volume=77|issue=1|pages=7–15|doi=10.1002/ajpa.1330770103|pmid=3189526}}</ref>

===Height and weight===
In comparison to the large, robust head, the body was rather small. Average weight for ''P. robustus'' may have been {{cvt|40|kg}} for males and {{cvt|32|kg}} for females;<ref name=Wood2000/> and for ''P. boisei'' {{cvt|50|kg}} for males and {{cvt|34|kg}} for females.<ref name=Wood2000/> At Swartkrans Cave Members 1 and 2, about 35% of the ''P. robustus'' individuals are estimated to have weighed {{cvt|28|kg}}, 22% about {{cvt|43|kg}}, and the remaining 43% bigger than the former but less than {{cvt|54|kg}}. At Member 3, all individuals were about {{cvt|45|kg}}.<ref name=McHenry1991a/> Female weight was about the same in contemporaneous ''H. erectus'', but male ''H. erectus'' were on average {{cvt|13|kg|1}} heavier than ''P. robustus'' males.<ref name=Susman2001>{{cite journal|first1=R. L.|last1=Susman|first2=D.|last2=de Ruiter|first3=C. K.|last3=Brain|year=2001|title=Recently identified postcranial remains of ''Paranthropus'' and Early ''Homo'' from Swartkrans Cave, South Africa|journal=Journal of Human Evolution|volume=41|issue=6|pages=607–629|doi=10.1006/jhev.2001.0510|pmid=11782111|bibcode=2001JHumE..41..607S |s2cid=26326715}}</ref> ''P. robustus'' sites are oddly dominated by small adults, which could be explained as heightened predation or mortality of the larger males of a group.<ref name=Braga2017>{{cite journal|first1=J.|last1=Braga|first2=J. F.|last2=Thackeray|first3=L.|last3=Bruxelles|first4=J.|last4=Dumoncel|first5=J.-P.|last5=Fourvel|year=2017|title=Stretching the time span of hominin evolution at Kromdraai (Gauteng, South Africa): Recent discoveries|journal=Comptes Rendus Palevol|volume=16|issue=1|pages=58–70|doi=10.1016/j.crpv.2016.03.003|bibcode=2017CRPal..16...58B |doi-access=free}}</ref> The largest-known ''Paranthropus'' individual was estimated at {{cvt|54|kg}}.<ref name=McHenry1991a>{{cite journal|first=H. M.|last=McHenry|author-link=Henry McHenry (anthropologist)|year=1991|title=Petite bodies of the "robust" australopithecines|journal=American Journal of Physical Anthropology|volume=86|issue=4|pages=445–454|doi=10.1002/ajpa.1330860402}}</ref>

According to a 1991 study, based on [[femur]] length and using the dimensions of modern humans, male and female ''P. robustus'' are estimated to have stood on average {{cvt|132|and|110|cm|ftin|0}}, respectively, and ''P. boisei'' {{cvt|137|and|124|cm|ftin|0}}. However, the latter estimates are problematic as there were no positively identified male ''P. boisei'' femurs at the time.<ref name=McHenry1991b>{{cite journal|first=H. M.|last=McHenry|author-link=Henry McHenry (anthropologist)|year=1991|title=Femoral lengths and stature in Plio-Pleistocene hominids|journal=American Journal of Physical Anthropology|volume=85|issue=2|pages=149–158|doi=10.1002/ajpa.1330850204|pmid=1882979}}</ref> In 2013, a 1.34 Ma male ''P. boisei'' partial skeleton was estimated to be at least {{cvt|156|cm|ftin|0}} and {{cvt|50|kg}}.<ref name=Dominguez2013>{{cite journal|first1=M.|last1=Domínguez-Rodrigo|first2=T. R.|last2=Rayne|first3=E.|last3=Baquedano|display-authors=et al.|year=2013|title=First Partial Skeleton of a 1.34-Million-Year-Old ''Paranthropus boisei'' from Bed II, Olduvai Gorge, Tanzania|journal=PLOS ONE|volume=8|issue=12|page=e80347|pmc=3855051|pmid=24339873|doi=10.1371/journal.pone.0080347|bibcode=2013PLoSO...880347D|doi-access=free}}</ref>

===Pathology===
''Paranthropus'' seems to have had notably high rates of [[pitting enamel hypoplasia]] (PEH), where [[tooth enamel]] formation is spotty instead of mostly uniform. In ''P. robustus'', about 47% of [[baby teeth]] and 14% of adult teeth were affected, in comparison to about 6.7% and 4.3%, respectively, in any other tested hominin species. The condition of these holes covering the entire tooth is consistent with the modern human ailment [[amelogenesis imperfecta]]. However, since circular holes in enamel coverage are uniform in size, only present on the [[molar teeth]], and have the same severity across individuals, the PEH may have been a genetic condition. It is possible that the [[coding region|coding-DNA]] concerned with thickening enamel also left them more vulnerable to PEH.<ref>{{cite journal|first1=I.|last1=Towle|first2=J. D.|last2=Irish|year=2019|title=A probable genetic origin for pitting enamel hypoplasia on the molars of ''Paranthropus robustus''|journal=Journal of Human Evolution|volume=129|pages=54–61|doi=10.1016/j.jhevol.2019.01.002|pmid=30904040|bibcode=2019JHumE.129...54T |s2cid=85502058|url=http://researchonline.ljmu.ac.uk/id/eprint/10289/1/Towle_Irish_JHE%202019.pdf}}</ref>

There have been 10 identified cases of [[tooth decay|cavities]] in ''P. robustus'', indicating a rate similar to modern humans. A molar from [[Drimolen]], South Africa, showed a cavity on the [[tooth root]], a rare occurrence in fossil [[great ape]]s. In order for cavity-creating bacteria to reach this area, the individual would have had to have also presented either [[alveolar process#Alveolar bone loss|alveolar resportion]], which is commonly associated with [[gum disease]]; or super-eruption of teeth which occurs when teeth become worn down and have to erupt a bit more in order to maintain a proper bite, and this exposed the root. The latter is most likely, and the exposed root seems to have caused [[hypercementosis]] to anchor the tooth in place. The cavity seems to have been healing, which may have been caused by a change in diet or [[oral microbiology|mouth microbiome]], or the loss of the adjacent molar.<ref name=Towle2019>{{cite journal|first1=I.|last1=Towle|first2=A.|last2=Riga|first3=J. D.|last3=Irish|display-authors=et al.|year=2019|title=Root caries on a ''Paranthropus robustus'' third molar from Drimolen|journal=American Journal of Physical Anthropology|volume=170|issue=2|pages=319–323|doi=10.1002/ajpa.23891|pmid=31265762|s2cid=195786562|url=https://www.biorxiv.org/content/biorxiv/early/2019/04/11/573964.full.pdf}}</ref>

==Palaeobiology==
===Diet===
[[File:Paranthropus boisei side (University of Zurich).JPG|thumb|left|''P. boisei'' [[OH 5]]]]
It was once thought ''P. boisei'' cracked open nuts with its powerful teeth, giving OH 5 the nickname "Nutcracker Man". However, like gorillas, ''Paranthropus'' likely preferred soft foods, but would consume tough or hard food during leaner times, and the powerful jaws were used only in the latter situation.<ref>{{cite journal|first1=P. S.|last1=Ungar|first2=F. E.|last2=Grine|first3=M. F.|last3=Teaford|year=2008|title=Dental Microwear and Diet of the Plio-Pleistocene Hominin ''Paranthropus boisei''|journal=PLOS ONE|volume=3|issue=4|page=e2044|doi=10.1371/journal.pone.0002044|pmc=2315797|pmid=18446200|bibcode=2008PLoSO...3.2044U|doi-access=free}}</ref> In ''P. boisei'', thick enamel was more likely used to resist abrasive gritty particles rather than to minimize chipping while eating hard foods.<ref>{{cite journal|first1=D.|last1=Rabenold|first2=O. M.|last2=Pearson|year=2011|title=Abrasive, Silica Phytoliths and the Evolution of Thick Molar Enamel in Primates, with Implications for the Diet of ''Paranthropus boisei''|journal=PLOS ONE|volume=6|issue=12|page=e28379|doi=10.1371/journal.pone.0028379|pmc=3233556|pmid=22163299|bibcode=2011PLoSO...628379R|doi-access=free}}</ref> In fact, there is a distinct lack of tooth fractures which would have resulted from such activity.<ref name=":1">{{Cite journal|last1=Towle|first1=Ian|last2=Irish|first2=Joel D.|last3=Groote|first3=Isabelle De|date=2017|title=Behavioral inferences from the high levels of dental chipping in ''Homo naledi''|journal=American Journal of Physical Anthropology|language=en|volume=164|issue=1|pages=184–192|doi=10.1002/ajpa.23250|pmid=28542710|s2cid=24296825 |issn=1096-8644|url=http://researchonline.ljmu.ac.uk/6367/3/Towle%20et%20al%20Homo%20naledi%20chipping%20%282%29.pdf}}</ref><ref>{{Cite journal|year=2005|title=Hominins, sedges, and termites: new carbon isotope data from the Sterkfontein valley and Kruger National Park|journal=Journal of Human Evolution|language=en|volume=48|issue=3|pages=301–312|doi=10.1016/j.jhevol.2004.11.008|pmid=15737395|last1=Sponheimer|first1=M.|last2=Lee-Thorp|first2=J.|last3=De Ruiter|first3=D.|last4=Codron|first4=D.|last5=Codron|first5=J.|last6=Baugh|first6=A. T.|last7=Thackeray|first7=F.|bibcode=2005JHumE..48..301S |citeseerx=10.1.1.421.8468}}</ref>

''Paranthropus'' were generalist feeders, but diet seems to have ranged dramatically with location. The South African ''P. robustus'' appears to have been an omnivore, with a diet similar to contemporaneous ''Homo''<ref name=Wood2004>{{cite journal|first1=B.|last1=Wood|first2=D.|last2=Strait|year=2004|title=Patterns of resource use in early ''Homo'' and ''Paranthropus''|journal=Journal of Human Evolution|volume=46|issue=2|pages=119–162|doi=10.1016/j.jhevol.2003.11.004|pmid=14871560|bibcode=2004JHumE..46..119W }}</ref> and nearly identical to the later ''[[H. ergaster]]'',<ref name=Lee2000>{{cite journal|first1=J.|last1=Lee-Thorp|first2=J. F.|last2=Thackeray|first3=N. V.|last3=der Merwe|year=2000|title=The hunters and the hunted revisited|journal=Journal of Human Evolution|volume=39|issue=6|pages=565–576|doi=10.1006/jhev.2000.0436|pmid=11102267|bibcode=2000JHumE..39..565L }}</ref> and subsisted on mainly [[C4 carbon fixation|C4]] savanna plants and [[C3 carbon fixation|C3]] forest plants, which could indicate either seasonal shifts in diet or seasonal migration from forest to savanna. In leaner times it may have fallen back on brittle food. It likely also consumed seeds<ref name=Sponheimer2006>{{cite journal|first1=M.|last1=Sponheimer|first2=B. H.|last2=Passey|first3=D. J.|last3=de Ruiter|display-authors=et al.|year=2006|title=Isotopic Evidence for Dietary Variability in the Early Hominin ''Paranthropus robustus''|journal=Science|volume=314|issue=5801|pages=980–982|doi=10.1126/science.1133827|pmid=17095699|bibcode=2006Sci...314..980S|s2cid=22291574}}</ref><ref>{{cite journal|first1=P. J.|last1=Constantino|first2=O.|last2=Borrero-Lopez|first3=B. R.|last3=Lawn|year=2018|title=Mechanisms of Tooth Damage in ''Paranthropus'' Dietary Reconstruction|journal= Biosurface and Biotribology|volume=4|issue=3|pages=73–78|doi=10.1049/bsbt.2018.0017|doi-access=free}}</ref> and possibly [[tuber]]s or [[termite]]s.<ref name=Backwell2001/> A high cavity rate could indicate [[honey]] consumption.<ref name=Towle2019/>

The East African ''P. boisei'', on the other hand, seems to have been largely herbivorous and fed on C4 plants. Its powerful jaws allowed it to consume a wide variety of different plants,<ref name=Wood2012/><ref name=Cerling2011>{{cite journal|last1=Cerling|first1=T. E.|last2=Mbua|first2=E.|last3=Kirera|first3=F. M.|display-authors=et al.|title=Diet of ''Paranthropus boisei'' in the early Pleistocene of East Africa |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=108 |issue=23 |pages=9337–41 |year= 2011  |pmid=21536914 |pmc=3111323 |doi=10.1073/pnas.1104627108 |bibcode=2011PNAS..108.9337C|doi-access=free}}</ref> though it may have largely preferred nutrient-rich [[bulbotuber]]s as these are known to thrive in the well-watered woodlands it is thought to have inhabited. Feeding on these, ''P. boisei'' may have been able to meet its daily caloric requirements of approximately 9,700 kJ after about 6 hours of foraging.<ref name=Macho2014>{{cite journal|first=G. M.|last=Macho|year=2014|title=Baboon Feeding Ecology Informs the Dietary Niche of ''Paranthropus boisei''|journal=PLOS ONE|volume=9|issue=1|page=e84942|doi=10.1371/journal.pone.0084942|pmc=3885648|pmid=24416315|bibcode=2014PLoSO...984942M|doi-access=free}}</ref>

Juvenile ''P. robustus'' may have relied more on tubers than adults, given the elevated levels of [[strontium]] compared to adults in teeth from Swartkrans Cave, which, in the area, was most likely sourced from tubers. [[Dentin]] exposure on juvenile teeth could indicate early weaning, or a more abrasive diet than adults which wore away the [[cementum]] and [[tooth enamel|enamel]] coatings, or both. It is also possible juveniles were less capable of removing grit from dug-up food rather than purposefully seeking out more abrasive foods.<ref name=Williams2015/>

===Technology===
[[File:DNH7.jpg|thumb|left|upright|[[DNH 7]], the most complete ''P. robustus'' skull known until the description of DNH155 in 2020<ref name=Stammers2018/>]]

Oldowan toolkits were uncovered at an excavation site on the Homa Peninsula in western Kenya. Stone tools called "oldowan toolkits" are used to pound and shape other rocks or plant materials. These tools are thought to be between 2.6 and 3 million years old. The stone tools were found near Paranthropus teeth.<ref>{{cite web|url=https://www.msn.com/en-us/money/smallbusiness/stone-tools-used-by-hippo-eating-human-ancestors-are-millions-of-years-old-researchers-say/ar-AA17iMEE?ocid=Peregrine&cvid=5d8b2955d48b41618f23d23f75d09dc3|title=Stone Tools Used By Hippo-Eating Human Ancestors Are Millions Of Years Old, Researchers Say|website=[[MSN]] |date=2023-02-09}}</ref>

Bone tools dating between 2.3 and 0.6 mya have been found in abundance in Swartkrans,<ref name=Backwell2001/> Kromdraai and [[Drimolen]] caves, and are often associated with ''P. robustus''. Though ''Homo'' is also known from these caves, their remains are comparatively scarce to ''Paranthropus'', making ''Homo''-attribution unlikely. The tools also cooccur with ''Homo''-associated [[Oldawan]] and possibly [[Acheulian]] stone tool [[industry (archaeology)|industries]]. The bone tools were typically sourced from the [[diaphysis|shaft]] of [[long bone]]s from medium- to large-sized mammals, but tools made sourced from [[mandible]]s, [[rib]]s and horn cores have also been found. Bone tools have also been found at Oldawan Gorge and directly associated with ''P. boisei'', the youngest dating to 1.34 mya, though a great proportion of other bone tools from here have ambiguous attribution. Stone tools from Kromdraai could possibly be attributed to ''P. robustus'', as no ''Homo'' have been found there yet.<ref name=Stammers2018/>

The bone tools were not manufactured or purposefully shaped for a task. However, since the bones display no weathering (and were not scavenged randomly), and there is a preference displayed for certain bones, raw materials were likely specifically hand-picked. This could indicate a similar cognitive ability to contemporary Stone Age ''Homo''.<ref name=Stammers2018>{{cite journal|first1=R. C.|last1=Stammers|first2=M.|last2=Caruana|first3=A. I. R.|last3=Herries|year=2018|title=The first bone tools from Kromdraai and stone tools from Drimolen, and the place of bone tools in the South African Earlier Stone Age|journal= Quaternary International|volume=495|pages=87–101|doi=10.1016/j.quaint.2018.04.026|bibcode=2018QuInt.495...87S|s2cid=135196415|doi-access=free}}</ref>

Bone tools may have been used to cut or process vegetation,<ref name=Susman1988>{{cite journal |last=Susman|first=R. L. |title=Hand of ''Paranthropus robustus'' from Member 1, Swartkrans: fossil evidence for tool behavior |journal=Science |volume=240 |issue=4853 |pages=781–784 |year= 1988  |pmid=3129783 |doi=10.1126/science.3129783|bibcode=1988Sci...240..781S }}</ref> or dig up [[tuber]]s or [[termite]]s,<ref name=Stammers2018/><ref name=Backwell2001>{{cite journal|first1=L. R.|last1=Backwell|first2=F.|last2=d'Errico|year=2001|title=Evidence of termite foraging by Swartkrans early hominids|journal=Proceedings of the National Academy of Sciences|volume=98|issue=4|pages=1358–1363|doi=10.1073/pnas.021551598|pmid=11171955|pmc=29261|doi-access=free}}</ref> The form of ''P. robustus'' incisors appear to be intermediate between ''H. erectus'' and modern humans, which could indicate less food processing done by the teeth due to preparation with simple tools.<ref name=Williams2015/>

Burnt bones were also associated with the inhabitants of Swartkrans, which could indicate some of the earliest fire usage.<ref name=Brain1988>{{cite journal|first1=C. K.|last1=Brain|first2=A.|last2=Sillent|year=1988|title=Evidence from the Swartkrans cave for the earliest use of fire|journal=Nature|volume=336|issue=6198|pages=464–466|doi=10.1038/336464a0|bibcode=1988Natur.336..464B|s2cid=4318364}}</ref> However, these bones were found in Member 3, where ''Paranthropus'' remains are rarer than ''H. erectus'', and it is also possible the bones were burned in a wildfire and washed into the cave as it is known the bones were not burned onsite.<ref name=Pickering>{{cite journal|last=Pickering|first=T. R.|year=2012|title=What's new is old: comments on (more) archaeological evidence of one-million-year-old fire from South Africa|journal=South African Journal of Science|volume=108|issue=5–6|pages=1–2|doi=10.4102/sajs.v108i5/6.1250|url=https://www.researchgate.net/publication/262749300|doi-access=free}}</ref><ref>{{cite journal|first1=J. A. J.|last1=Gowlett|first2=R. W.|last2=Wrangham|year=2013|title=Earliest fire in Africa: towards the convergence of archaeological evidence and the cooking hypothesis|journal=Azania: Archaeological Research in Africa|volume=48|issue=1|pages=16–17<!--only citing these pages-->|doi=10.1080/0067270X.2012.756754|s2cid=163033909}}</ref>

===Social structure===
[[File:Paranthropus aethiopicus face (University of Zurich) blackbckgr.JPG|thumb|upright|''P. aethiopicus'' [[KNM WT 17000]]]]
Given the marked anatomical and physical differences with modern great apes, there may be no modern analogue for australopithecine societies, so comparisons drawn with modern primates will not be entirely accurate.<ref name=Copeland2011/><ref name=Kaszycka2016/>

''Paranthropus'' had pronounced [[sexual dimorphism]], with males notably larger than females, which is commonly correlated with a male-dominated [[polygamous]] society. ''P. robustus'' may have had a harem society similar to modern forest-dwelling [[silverback gorilla]]s, where one male has exclusive breeding rights to a group of females, as male-female size disparity is comparable to gorillas (based on facial dimensions), and younger males were less robust than older males (delayed maturity is also exhibited in gorillas).<ref>{{cite journal|first1=C. A.|last1=Lockwood|first2=C. G.|last2=Menter|first3=J.|last3=Moggi-Cecchi|first4=A. W.|last4=Keyser|year=2007|title=Extended male growth in a fossil hominin species|journal=Science|volume=318|issue=5855|pages=1443–1446|doi=10.1126/science.1149211|pmid=18048687|bibcode=2007Sci...318.1443L|s2cid=32900905|url=http://doc.rero.ch/record/15464/files/PAL_E2851.pdf }}</ref>

However, if ''P. robustus'' preferred a savanna habitat, a multi-male society would have been more productive to better defend the troop from predators in the more exposed environment, much like savanna [[baboon]]s. Further, among primates, delayed maturity is also exhibited in the [[rhesus monkey]] which has a multi-male society, and may not be an accurate indicator of social structure.<ref name=Kaszycka2016>{{cite journal|first=K. A.|last=Kaszycka|year=2016|title=''Australopithecus robustus'' societies - one-male or multimale?|journal=South African Journal of Science|volume=112|issue=1–2|pages=124–131|doi=10.17159/sajs.2016/20150165|doi-access=free}}</ref>

A 2011 [[strontium isotope]] study of ''P. robustus'' teeth from the [[dolomite (rock)|dolomite]] [[Sterkfontein]] Valley found that, like other [[hominin]]s, but unlike other great apes, ''P. robustus'' females were more likely to leave their place of birth ([[patrilocal]]). This also discounts the plausibility of a harem society, which would have resulted in a [[matrilocal]] society due to heightened male–male competition. Males did not seem to have ventured very far from the valley, which could either indicate small home ranges, or that they preferred dolomitic landscapes due to perhaps cave abundance or factors related to vegetation growth.<ref name=Copeland2011>{{cite journal|first1=S. R.|last1=Copeland|first2=M.|last2=Sponheimmer|first3=D. J.|last3=de Ruiter|first4=J.|last4=Lee-Thorp|year=2011|title=Strontium isotope evidence for landscape use by early hominins|journal=Nature|volume=474|issue=7349|pages=76–78|doi=10.1038/nature10149|pmid=21637256|s2cid=205225222}}</ref>

===Life history===
Dental development seems to have followed about the same timeframe as it does in modern humans and most other hominins, but, since ''Paranthropus'' molars are markedly larger, rate of [[tooth eruption]] would have been accelerated.<ref name=Wood2007/><ref>{{cite journal|first=M. C.|last=Dean|year=1985|title=The eruption pattern of the permanent incisors and first permanent molars in ''Australopithecus (Paranthropus) robustus''|journal=American Journal of Physical Anthropology|volume=67|issue=3|pages=251–257|doi=10.1002/ajpa.1330670310|pmid=3933358}}</ref> Their life history may have mirrored that of gorillas as they have the same brain volume,<ref>{{cite journal|first1=J.|last1=Kelley|first2=G. T.|last2=Schwartz|year=2012|title=Life-History Inference in the Early Hominins ''Australopithecus'' and ''Paranthropus''|journal=International Journal of Primatology|volume=33|issue=6|pages=1332–1363|doi=10.1007/s10764-012-9607-2|s2cid=16288970}}</ref> which (depending on the subspecies) reach physical maturity from 12–18 years and have birthing intervals of 40–70 months.<ref>{{cite journal|first1=M. M.|last1=Robbins|first2=A. M.|last2=Robbins|year=2018|title=Variation in the social organization of gorillas: Life history and socioecological perspectives|journal=Evolutionary Anthropology|volume=27|issue=5|pages=218–233|doi=10.1002/evan.21721|pmid=30325554|s2cid=53100488|url=https://www.eva.mpg.de/documents/Wiley-Blackwell/Robbins_Variation_EvolAnthrop_2018_3007627.pdf}}</ref>

==Palaeoecology==
===Habitat===
[[File:Upper Omo River Valley, Ethiopia (12562049655).jpg|thumb|upright|Modern-day [[Omo River]] Valley]]
It is generally thought that ''Paranthropus'' preferred to inhabit wooded, riverine landscapes.<ref name=Cerling2011/> The teeth of ''Paranthropus'', ''H. habilis'' and ''[[H. erectus]]'' are all known from various overlapping beds in East Africa, such as at Olduvai Gorge<ref>{{cite journal|first=R. J.|last=Clarke|year=2012|title=A ''Homo habilis'' maxilla and other newly-discovered hominid fossils from Olduvai Gorge, Tanzania|journal=Journal of Human Evolution|volume=63|issue=2|pages=418–428|doi=10.1016/j.jhevol.2011.11.007|pmid=22561056|bibcode=2012JHumE..63..418C }}</ref> and the [[Turkana Basin]].<ref name="Lague2019"/> ''P. robustus'' and ''H. erectus'' also appear to have coexisted.<ref name=Susman2001/><ref name=Pickering/>

''P. boisei'', known from the Great Rift Valley, may have typically inhabited wetlands along lakes and rivers, wooded or arid [[shrubland]]s, and semiarid woodlands,<ref name=Cerling2011/> though their presence in the savanna-dominated Malawian [[Chiwondo Beds]] implies they could tolerate a range of habitats.<ref name=Bocherens>{{cite journal|first1=H.|last1=Bocherens|first2=O.|last2=Sandrock|first3=O.|last3=Kullmer|first4=F.|last4=Schrenk|year=2011|title=Hominin palaeoecology in late Pliocene Malawi: first insights from isotopes (<sup>13</sup>C, <sup>18</sup>O) in mammal teeth|journal=South African Journal of Science|volume=107|issue=3–4|pages=1–6|doi=10.4102/sajs.v107i3/4.331|doi-access=free}}</ref> During the Pleistocene, there seem to have been coastal and montane forests in Eastern Africa. More expansive river valleys—namely the [[Omo River]] Valley—may have served as important refuges for forest-dwelling creatures. Being cut off from the forests of Central Africa by a savanna corridor, these East African forests would have promoted high rates of [[endemism]], especially during times of climatic volatility.<ref>{{cite journal|first=R.|last=Bobe|year=2006|title=The evolution of arid ecosystems in eastern Africa|journal=Journal of Arid Environments|volume=66|issue=3|pages=564–584|doi=10.1016/j.jaridenv.2006.01.010|bibcode=2006JArEn..66..564B}}</ref>

The Cradle of Humankind, the only area ''P. robustus'' is known from, was mainly dominated by the [[springbok]] ''Antidorcas recki'', but other antelope, [[Giraffidae|giraffes]] and [[Elephantidae|elephants]] were also seemingly abundant megafauna. Other known primates are early ''Homo'', the [[hamadryas baboon]], and the extinct [[colobine]] monkey ''[[Cercopithecoides williamsi]]''.<ref name=Adams>{{cite journal|first1=J. W.|last1=Adams|first2=D. S.|last2=Rovinsky|first3=A. I. R.|last3=Herries|first4=C. G.|last4=Menter|year=2016|title=Macromammalian faunas, biochronology and palaeoecology of the early Pleistocene Main Quarry hominin-bearing deposits of the Drimolen Palaeocave System, South Africa|journal=PeerJ|volume=4|page=e1941|pmid=27114884|pmc=4841245|doi=10.7717/peerj.1941 |doi-access=free }}</ref>

===Predators===

The left foot of a ''P. boisei'' specimen (though perhaps actually belonging to ''H. habilis'') from Olduvai Gorge seems to have been bitten off by a crocodile,<ref name=Njau2012>{{cite journal|first1=J. K.|last1=Njau|first2=R. J.|last2=Blumenschine|year=2012|title=Crocodylian and mammalian carnivore feeding traces on hominid fossils from FLK 22 and FLK NN 3, Plio-Pleistocene, Olduvai Gorge, Tanzania|journal=Journal of Human Evolution|volume=63|issue=2|pages=408–417|doi=10.1016/j.jhevol.2011.05.008|pmid=21937084|bibcode=2012JHumE..63..408N }}</ref> possibly ''[[Crocodylus anthropophagus]]'',<ref name=Brochu2010>{{cite journal |first1=C. A.|last1=Brochu|first2=J.|last2=Njau|first3=R. J.|last3=Blumenschine|first4=L. D.|last4=Densmore |year=2010 |title=A New Horned Crocodile from the Plio-Pleistocene Hominid Sites at Olduvai Gorge, Tanzania |journal=PLOS ONE |volume=5 |issue=2 |pages=e9333 |doi=10.1371/journal.pone.0009333 |pmid=20195356 |pmc=2827537|bibcode=2010PLoSO...5.9333B|doi-access=free}}</ref> and another's leg shows evidence of leopard predation.<ref name=Njau2012/> Other likely Olduvan predators of great apes include the [[hunting hyena]] ''Chasmaporthetes nitidula'', and the [[sabertoothed cat]]s ''[[Dinofelis]]'' and ''[[Megantereon]]''.<ref name=Lee2000/> The carnivore assemblage at the Cradle of Humankind comprises the two sabertooths, and the hyena ''[[Lycyaenops silberbergi]]''.<ref name=Adams/>

Male ''P. robustus'' appear to have had a higher mortality rate than females. It is possible that males were more likely to be kicked out of a group, and these lone males had a higher risk of predation.<ref name=Kaszycka2016/>

===Extinction===
It was once thought that ''Paranthropus'' had become a specialist feeder, and were inferior to the more adaptable tool-producing ''Homo'', leading to their extinction, but this has been called into question.<ref name=Wood2004/><ref name=Lee2000/><ref name=Sponheimer2006/><ref name=Backwell2001/><ref name=Susman1988/> However, smaller brain size may have been a factor in their extinction along with gracile australopithecines.<ref name=Du2018/> ''P. boisei'' may have died out due to an arid trend starting 1.45 mya, causing the retreat of woodlands, and more competition with savanna baboons and ''Homo'' for alternative food resources.<ref name=Macho2014/>

South African ''Paranthropus'' appear to have outlasted their East African counterparts.<ref name=Herries2009/> The youngest record of ''P. boisei'' comes from [[Konso]], Ethiopia about 1.4 mya; however, there are no East African sites dated between 1.4 and 1 mya, so it may have persisted until 1 mya.<ref name=Wood2007/> ''P. robustus'', on the other hand, was recorded in [[Swartkrans]] until Member 3 dated to 1–0.6 mya (the [[Middle Pleistocene]]), though more likely the younger side of the estimate.<ref name=Herries2009>{{cite journal|first1=A. I. R.|last1=Herries|first2=D.|last2=Curnoe|first3=J. W.|last3=Adams|year=2009|title=A multi-disciplinary seriation of early ''Homo'' and ''Paranthropus'' bearing palaeocaves in southern Africa|journal=Quaternary International|volume=202|issue=1–2|pages=14–28|doi=10.1016/j.quaint.2008.05.017|bibcode=2009QuInt.202...14H}}</ref>

==See also==
{{Portal|Paleontology}}
{{div col|colwidth=30}}
* ''[[Australopithecus]]''
* ''[[Ardipithecus]]''
* ''[[Graecopithecus]]''
* ''[[Orrorin]]''
* ''[[Sahelanthropus]]''
{{div col end}}

==References==
{{Reflist}}

==Further reading==

*{{cite book|first=F. E.|last=Grine|year=2007|title=Evolutionary History of the Robust Australopithecines|publisher=Transaction Publishers|isbn=978-0-202-36596-1}}
*{{cite journal |last1=Wood |first1=Bernard |last2=Williams |first2=Alexis |title=Meet Your Exotic, Extinct Close Relative: For a million years our likely ancestors in eastern Africa lived alongside creatures so peculiar that scientists today still struggle to make sense of them. |journal=American Scientist |date=2020 |volume=108 |issue=6 |page=348 |doi=10.1511/2020.108.6.348 |s2cid=241348079 |url=https://www.americanscientist.org/article/meet-your-exotic-extinct-close-relative|url-access=subscription }}

==External links==
{{wiktionary}}
{{Wikispecies}}
{{Commons category|Paranthropus}}
{{Wikibooks|Introduction to Paleoanthropology}}
* [http://gurche.com/homo-floresiensis-1-1 Reconstructions of ''P. boisei''] by [[John Gurche]]
* {{cite web
 | url = http://www.mnh.si.edu/anthro/humanorigins/ha/a_tree.html
 | title = Early Human Phylogeny
 | publisher = [[Smithsonian Institution]]
 | access-date = 2006-09-14
 | archive-date = 2005-11-02
 | archive-url = https://web.archive.org/web/20051102090342/http://www.mnh.si.edu/anthro/humanorigins/ha/a_tree.html
 | url-status = dead
 }}
* [http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive Human Timeline (Interactive)] – [[Smithsonian Institution|Smithsonian]], [[National Museum of Natural History]] (August 2016).

{{Human Evolution}}
{{Haplorhini|Ho.}}
{{Taxonbar|from=Q111463}}

[[Category:Paranthropus| ]]
[[Category:Prehistoric primate genera]]
[[Category:Pliocene primates]]
[[Category:Pleistocene primates]]
[[Category:Pleistocene extinctions]]
[[Category:Cenozoic mammals of Africa]]
[[Category:Pleistocene genus extinctions]]
[[Category:Fossil taxa described in 1938]]
[[Category:Cradle of Humankind fauna]]