Editing Paranthropus (section)

==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>