Editing Human evolution (section)

== Before ''Homo'' ==
{{For| evolutionary history before primates|Evolution of mammals|History of life|Timeline of human evolution}}

=== Early evolution of primates ===
{{See also|Evolution of primates}}
The evolutionary history of primates can be traced back 65&nbsp;million years.{{sfn|Maxwell|1984|p=296}}<ref>{{cite journal |last1=Zhang |first1=Rui |last2=Wang |first2=Yin-Qiu |last3=Su |first3=Bing |date=July 2008 |title=Molecular Evolution of a Primate-Specific microRNA Family |journal=Molecular Biology and Evolution |volume=25 |issue=7 |pages=1493–1502 |doi=10.1093/molbev/msn094 |issn=0737-4038 |pmid=18417486 |doi-access=free}}</ref><ref>{{cite journal |last=Willoughby |first=Pamela R. |date=2005 |title=Palaeoanthropology and the Evolutionary Place of Humans in Nature |url= http://escholarship.org/uc/item/92w669xb |journal=International Journal of Comparative Psychology |volume=18 |issue=1 |pages=60–91 |doi=10.46867/IJCP.2005.18.01.02 |issn=0889-3667 |access-date=April 27, 2015 |archive-date=January 17, 2012 |archive-url= https://web.archive.org/web/20120117154844/http://escholarship.org/uc/item/92w669xb |url-status=live |doi-access=free}}</ref>{{sfn|Martin|2001|pp=12032–12038}}<ref>{{cite journal |last1=Tavaré |first1=Simon |author1-link=Simon Tavaré |last2=Marshall |first2=Charles R. |last3=Will |first3=Oliver |last4=Soligo |first4=Christophe |last5=Martin |first5=Robert D. |author4-link=Robert D. Martin |display-authors=3 |date=April 18, 2002 |title=Using the fossil record to estimate the age of the last common ancestor of extant primates |journal=[[Nature (journal)|Nature]] |volume=416 |issue=6882 |pages=726–729 |doi=10.1038/416726a |issn=0028-0836 |pmid=11961552 |bibcode=2002Natur.416..726T |s2cid=4368374}}</ref> One of the oldest known primate-like mammal species, the ''[[Plesiadapis]]'', came from North America;<ref>{{cite journal |last=Rose |first=Kenneth D. |date=1994 |title=The earliest primates |journal=Evolutionary Anthropology: Issues, News, and Reviews |volume=3 |issue=5 |pages=159–173 |doi=10.1002/evan.1360030505 |s2cid=85035753 |issn=1060-1538}}</ref><ref>{{cite web |url= http://alltheworldsprimates.org/John_Fleagle_Public.aspx |title=Primate Evolution |last1=Fleagle |first1=John |author1-link=John G. Fleagle |last2=Gilbert |first2=Chris |date=2011 |editor1-last=Rowe |editor1-first=Noel |editor2-last=Myers |editor2-first=Marc |website=All The World's Primates |publisher=Primate Conservation |location=Charlestown, Rhode Island |access-date=April 27, 2015 |archive-date=May 12, 2015 |archive-url= https://web.archive.org/web/20150512013618/http://www.alltheworldsprimates.org/john_fleagle_public.aspx |url-status=live}}</ref><ref>{{cite news |last=Roach |first=John |date=March 3, 2008 |title=Oldest Primate Fossil in North America Discovered |url= http://news.nationalgeographic.com/news/2008/03/080303-american-primate.html |work=National Geographic News |location=Washington, DC |publisher=[[National Geographic Society]] |access-date=April 27, 2015 |archive-date=October 16, 2012 |archive-url= https://web.archive.org/web/20121016193314/http://news.nationalgeographic.com/news/2008/03/080303-american-primate.html |url-status=dead}}</ref><ref>{{cite news |last=McMains |first=Vanessa |date=December 5, 2011 |title=Found in Wyoming: New fossils of oldest American primate |url= http://gazette.jhu.edu/2011/12/05/found-in-wyoming-new-fossils-of-oldest-american-primate/ |newspaper=The Gazette |location=Baltimore |publisher=[[Johns Hopkins University]] |access-date=April 27, 2015 |archive-date=January 16, 2019 |archive-url= https://web.archive.org/web/20190116200750/https://gazette.jhu.edu/2011/12/05/found-in-wyoming-new-fossils-of-oldest-american-primate/ |url-status=live}}</ref><ref>{{cite news |last=Caldwell |first=Sara B. |date=May 19, 2009 |title=Missing link found, early primate fossil 47 million years old |location=Toronto |work=Digital Journal |url= http://www.digitaljournal.com/article/272808 |access-date=April 27, 2015 |archive-date=July 22, 2015 |archive-url= https://web.archive.org/web/20150722061001/http://www.digitaljournal.com/article/272808 |url-status=live}}</ref><ref>{{cite news |last=Watts |first=Alex |date=May 20, 2009 |title=Scientists Unveil Missing Link In Evolution |url= http://news.sky.com/home/world-news/article/15284582 |access-date=April 27, 2015 |work=[[Sky News]] Online |location=London |publisher=[[Sky (United Kingdom)|BSkyB]] |archive-url= https://web.archive.org/web/20110728005247/http://news.sky.com/home/world-news/article/15284582 |archive-date=July 28, 2011}}</ref> another, ''[[Archicebus]]'', came from [[China]].<ref name="NYT-20130605">{{cite news |last=Wilford |first=J. N. |title=Palm-size fossil resets primates' clock, scientists say |url= https://www.nytimes.com/2013/06/06/science/palm-size-fossil-resets-primates-clock-scientists-say.html |archive-url= https://ghostarchive.org/archive/20220101/https://www.nytimes.com/2013/06/06/science/palm-size-fossil-resets-primates-clock-scientists-say.html |archive-date=January 1, 2022 |url-access=limited |date=June 5, 2013 |work=[[The New York Times]] |access-date=June 5, 2013}}{{cbignore}}</ref> Other similar basal primates were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and [[Eocene]].

[[File:Notharctus tenebrosus AMNH.jpg|thumb|''[[Notharctus tenebrosus]]'', [[American Museum of Natural History]], New York]]

David R. Begun<ref name="Kordos-p17">{{cite journal |last1=Kordos |first1=László |last2=Begun |first2=David R. |date=January 2001 |title=Primates from Rudabánya: Allocation of specimens to individuals, sex and age categories |journal=Journal of Human Evolution |volume=40 |issue=1 |pages=17–39 |doi=10.1006/jhev.2000.0437 |issn=0047-2484 |pmid=11139358 |bibcode=2001JHumE..40...17K}}</ref> concluded that early primates flourished in Eurasia and that a lineage leading to the African apes and humans, including to ''[[Dryopithecus]]'', migrated south from Europe or Western Asia into Africa. The surviving tropical population of primates—which is seen most completely in the Upper Eocene and lowermost [[Oligocene]] fossil beds of the [[Faiyum]] depression southwest of [[Cairo]]—gave rise to all extant primate species, including the [[lemur]]s of [[Madagascar]], [[loris]]es of Southeast Asia, [[galago]]s or "bush babies" of Africa, and to the [[Simian|anthropoids]], which are the [[New World monkey|Platyrrhines]] or New World monkeys, the [[Catarrhines]] or Old World monkeys, and the great apes, including humans and other hominids.

The earliest known catarrhine is ''[[Kamoyapithecus]]'' from the uppermost Oligocene at Eragaleit in the northern [[Great Rift Valley, Kenya|Great Rift Valley]] in Kenya, dated to 24&nbsp;million years ago.{{sfn|Cameron|2004|p=[https://books.google.com/books?id=fnqzb4_UVfkC&pg=PA76 76]}} Its ancestry is thought to be species related to ''[[Aegyptopithecus]]'', ''[[Propliopithecus]]'', and ''[[Parapithecus]]'' from the Faiyum, at around 35&nbsp;mya.{{sfn|Wallace|2004|p=[https://books.google.com/books?id=YoyQQEOKGVQC&pg=PA240 240]}} In 2010, ''[[Saadanius]]'' was described as a close relative of the last common ancestor of the [[Crown group|crown]] catarrhines, and tentatively dated to 29–28&nbsp;mya, helping to fill an 11-million-year gap in the fossil record.<ref name="2010Zalmout">{{cite journal |last1=Zalmout |first1=Iyad S. |last2=Sanders |first2=William J. |author2-link=William J. Sanders |last3=MacLatchy |first3=Laura M. |last4=Gunnell |first4=Gregg F. |last5=Al-Mufarreh |first5=Yahya A. |last6=Ali |first6=Mohammad A. |last7=Nasser |first7=Abdul-Azziz H. |last8=Al-Masari |first8=Abdu M. |last9=Al-Sobhi |first9=Salih A. |last10=Nadhra |first10=Ayman O. |last11=Matari |first11=Adel H. |last12=Wilson |first12=Jeffrey A. |last13=Gingerich |first13=Philip D. |date=July 15, 2010 |title=New Oligocene primate from Saudi Arabia and the divergence of apes and Old World monkeys |journal=[[Nature (journal)|Nature]] |volume=466 |issue=7304 |pages=360–364 |bibcode=2010Natur.466..360Z |doi=10.1038/nature09094 |issn=0028-0836 |pmid=20631798 |s2cid=205220837 |display-authors=3}}</ref>

[[File:Proconsul skeleton reconstitution (University of Zurich).JPG|thumb|Reconstructed tailless ''[[Proconsul (mammal)|Proconsul]]'' skeleton]]

In the [[Early Miocene]], about 22&nbsp;million years ago, the many kinds of [[Arboreal locomotion|arboreally]]-adapted (tree-dwelling) primitive catarrhines from East Africa suggest a long history of prior diversification. [[Fossil|Fossils]] at 20&nbsp;million years ago include fragments attributed to ''[[Victoriapithecus]]'', the earliest Old World monkey. Among the genera thought to be in the [[ape]] lineage leading up to 13&nbsp;million years ago are ''[[Proconsul (mammal)|Proconsul]]'', ''[[Rangwapithecus]]'', ''[[Dendropithecus]]'', ''[[Limnopithecus]]'', ''[[Nacholapithecus]]'', ''[[Equatorius]]'', ''[[Nyanzapithecus pickfordi|Nyanzapithecus]]'', ''[[Afropithecus]]'', ''Heliopithecus'', and ''[[Kenyapithecus]]'', all from East Africa.

The presence of other generalized non-cercopithecids of [[Middle Miocene]] from sites far distant, such as ''[[Otavipithecus]]'' from cave deposits in Namibia, and ''[[Pierolapithecus]]'' and ''[[Dryopithecus]]'' from France, Spain and Austria, is evidence of a wide diversity of forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the Early and Middle Miocene. The youngest of the [[Miocene]] hominoids, ''[[Oreopithecus]]'', is from coal beds in Italy that have been dated to 9&nbsp;million years ago.

Molecular evidence indicates that the lineage of gibbons diverged from the line of great apes some 18–12&nbsp;mya, and that of orangutans (subfamily [[Ponginae]]){{efn|Not to be confused with [[Pongidae]], an obsolete family which grouped together [[orangutan]]s, [[gorilla]]s and [[chimpanzee]]s to separate them from humans}} diverged from the other great apes at about 12&nbsp;million years; there are no fossils that clearly document the ancestry of gibbons, which may have originated in a so-far-unknown Southeast Asian hominoid population, but fossil proto-orangutans may be represented by ''[[Sivapithecus]]'' from India and ''[[Griphopithecus]]'' from Turkey, dated to around 10&nbsp;mya.{{sfn|Srivastava|2009|p=[https://books.google.com/books?id=kCerOsM8XMwC&pg=PA87 87]}}

Hominidae subfamily [[Homininae]] (African hominids) diverged from Ponginae (orangutans) about 14&nbsp;mya. Hominins (including humans and the Australopithecine and [[Panina]] subtribes) parted from the [[Gorillini]] tribe (gorillas) between 8 and 9&nbsp;mya; Australopithecine (including the extinct biped ancestors of humans) separated from the ''Pan'' genus (containing chimpanzees and bonobos) 4–7&nbsp;mya.<ref name=":4" /> The ''Homo'' genus is evidenced by the appearance of ''H.&nbsp;habilis'' over 2&nbsp;mya,{{efn|name=habilis}} while anatomically modern humans emerged in Africa approximately 300,000 years ago.

=== Divergence of the human clade from other great apes ===
[[File:016 Alpha male chimpanzee walking at Kibale forest National Park Photo by Giles Laurent.jpg|thumb|upright|Genetic data suggest that [[chimpanzee]]s and humans may have diverged from a common ancestor between 9 and 7 million years ago.<ref>{{cite news |title=Scientists observe chimpanzees using human-like warfare tactic |url=https://www.reuters.com/science/scientists-observe-chimpanzees-using-human-like-warfare-tactic-2023-11-02/ |work=Reuters |date=3 November 2023}}</ref>]]
Species close to the last common ancestor of gorillas, chimpanzees and humans may be represented by ''[[Nakalipithecus]]'' fossils found in Kenya and ''[[Ouranopithecus]]'' found in Greece. Molecular evidence suggests that between 8 and 4&nbsp;million years ago, first the gorillas, and then the chimpanzees (genus ''Pan'') split off from the line leading to the humans. Human DNA is approximately 98.4% identical to that of chimpanzees when comparing single nucleotide polymorphisms (see [[human evolutionary genetics]]). The fossil record, however, of gorillas and chimpanzees is limited; both poor preservation – rain forest soils tend to be acidic and dissolve bone – and [[sampling bias]] probably contribute to this problem.

Other hominins probably adapted to the drier environments outside the equatorial belt; and there they encountered antelope, hyenas, dogs, pigs, elephants, horses, and others. The equatorial belt contracted after about 8&nbsp;million years ago, and there is very little fossil evidence for the split—thought to have occurred around that time—of the hominin lineage from the lineages of gorillas and chimpanzees. The earliest fossils argued by some to belong to the human lineage are ''Sahelanthropus tchadensis'' (7 Ma) and ''Orrorin tugenensis'' (6 Ma), followed by ''Ardipithecus'' (5.5–4.4 Ma), with species ''Ar. kadabba'' and ''[[Ar. ramidus]]''.

It has been argued in a study of the life history of ''Ar. ramidus'' that the species provides evidence for a suite of anatomical and behavioral adaptations in very early hominins unlike any species of extant great ape.<ref name="aramidus">{{Cite journal |last1=Clark |first1=G. |last2=Henneberg |first2=M. |title=The life history of Ardipithecus ramidus: A heterochronic model of sexual and social maturation |journal=[[Anthropological Review]] |volume=78 |issue=2 |pages=109–132 |date=June 2015 |doi=10.1515/anre-2015-0009 |s2cid=54900467 |doi-access=free}}</ref> This study demonstrated affinities between the skull morphology of ''Ar. ramidus'' and that of infant and juvenile chimpanzees, suggesting the species evolved a juvenalised or [[paedomorphic]] craniofacial morphology via [[Heterochrony|heterochronic]] dissociation of growth trajectories. It was also argued that the species provides support for the notion that very early hominins, akin to bonobos (''Pan paniscus'') the less aggressive species of the genus ''Pan'', may have evolved via the process of [[self-domestication]]. Consequently, arguing against the so-called "chimpanzee referential model"<ref name="sayers" /> the authors suggest it is no longer tenable to use chimpanzee (''Pan troglodytes'') social and mating behaviors in models of early hominin social evolution. When commenting on the absence of aggressive canine morphology in ''Ar. ramidus'' and the implications this has for the evolution of hominin social psychology, they wrote:

{{blockquote|Of course ''Ar. ramidus'' differs significantly from bonobos, bonobos having retained a functional canine honing complex. However, the fact that ''Ar. ramidus'' shares with bonobos reduced sexual dimorphism, and a more paedomorphic form relative to chimpanzees, suggests that the developmental and social adaptations evident in bonobos may be of assistance in future reconstructions of early hominin social and sexual psychology. In fact the trend towards increased maternal care, female mate selection and self-domestication may have been stronger and more refined in ''Ar. ramidus'' than what we see in bonobos.{{r|aramidus|page1=128}}}}

The authors argue that many of the basic human adaptations evolved in the ancient forest and woodland ecosystems of late [[Miocene]] and early [[Pliocene]] Africa. Consequently, they argue that humans may not represent evolution from a chimpanzee-like ancestor as has traditionally been supposed. This suggests many modern human adaptations represent [[Phylogeny|phylogenetically]] deep traits and that the behavior and morphology of chimpanzees may have evolved subsequent to the split with the common ancestor they share with humans.

=== Genus ''Australopithecus'' ===
{{Main|Australopithecus}}
[[File:Bienvenida al Museo de Lucy.jpg|thumb|upright|Reconstruction of "[[Lucy (Australopithecus)|Lucy]]"]]
The genus ''[[Australopithecus]]'' evolved in eastern Africa around 4&nbsp;million years ago before spreading throughout the continent and eventually becoming extinct 2&nbsp;million years ago. During this time period various forms of australopiths existed, including ''[[Australopithecus anamensis]]'', ''[[Australopithecus afarensis|A. afarensis]]'', ''[[Australopithecus sediba|A. sediba]]'', and ''[[Australopithecus africanus|A. africanus]]''. There is still some debate among academics whether certain African hominid species of this time, such as ''[[Australopithecus robustus|A. robustus]]'' and ''[[Australopithecus boisei|A. boisei]]'', constitute members of the same genus; if so, they would be considered to be "robust australopiths" while the others would be considered "gracile australopiths". However, if these species do indeed constitute their own genus, then they may be given their own name, ''Paranthropus''.
* ''[[Australopithecus]]'' (4–1.8 Ma), with species ''[[Australopithecus anamensis|A. anamensis]]'', ''[[Australopithecus afarensis|A. afarensis]]'', ''[[Australopithecus africanus|A. africanus]]'', ''[[Australopithecus bahrelghazali|A. bahrelghazali]]'', ''[[Australopithecus garhi|A. garhi]]'', and ''[[Australopithecus sediba|A. sediba]]'';
* ''[[Kenyanthropus]]'' (3–2.7 Ma), with species ''[[Kenyanthropus|K. platyops]]'';
* ''[[Paranthropus]]'' (3–1.2 Ma), with species ''[[Paranthropus aethiopicus|P. aethiopicus]]'', ''[[Paranthropus boisei|P. boisei]]'', and ''[[Paranthropus robustus|P. robustus]]''
A new proposed species ''[[Australopithecus deyiremeda]]'' is claimed to have been discovered living at the same time period of ''[[Australopithecus afarensis|A. afarensis]]''. There is debate whether ''[[Australopithecus deyiremeda|A. deyiremeda]]'' is a new species or is ''[[Australopithecus afarensis|A. afarensis]]''.<ref>{{cite news |last=Zimmer |first=Carl |author-link=Carl Zimmer |date=May 27, 2015 |title=The Human Family Tree Bristles With New Branches |url= https://www.nytimes.com/2015/06/02/science/adding-branches-to-the-human-family-tree.html |archive-url= https://ghostarchive.org/archive/20220101/https://www.nytimes.com/2015/06/02/science/adding-branches-to-the-human-family-tree.html |archive-date=January 1, 2022 |url-access=limited |newspaper=The New York Times |access-date=May 30, 2015}}{{cbignore}}</ref> ''Australopithecus prometheus'', otherwise known as [[Little Foot]] has recently been dated at 3.67&nbsp;million years old through a new dating technique, making the genus ''Australopithecus'' as old as ''afarensis''.<ref>Gardner., Elizabeth K.; Purdue University (April 1, 2015). "New instrument dates old skeleton before 'Lucy'; 'Little Foot' 3.67 million years old". Science Daily. Retrieved April 3, 2015.</ref> Given the opposable big toe found on Little Foot, it seems that the specimen was a good climber. It is thought given the night predators of the region that he built a nesting platform at night in the trees in a similar fashion to chimpanzees and gorillas.
[[File:Chimp's Nest.jpg|thumb|Chimpanzee nest. Later [[hominin]]s may have developed [[Niche construction|niche creating]] shelter-building traditions from such earlier [[Nest-building in primates|nest-building practices]].]]