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The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''
Homo ''Homo'' () is the genus that emerged in the (otherwise extinct) genus ''Australopithecus'' that encompasses the extant species ''Homo sapiens'' ( modern humans), plus several extinct species classified as either ancestral to or closely relate ...
'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''
Gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
'' (gorillas). The term was originally introduced by
Camille Arambourg Camille Arambourg ( February 3, 1885– November 19, 1969) was a French vertebrate paleontologist. He conducted extensive field work in North Africa. In the 1950s he argued against the prevailing model of Neanderthals as brutish and simian. Du ...
(1948). Arambourg combined the categories of ''Hominina'' and ''Simiina'' due to Gray (1825) into his new subtribe. Traditionally,
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative th ...
s,
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
s and
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s were grouped together as pongids. Since Gray's classification, evidence has accumulated from genetic
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
confirming that
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
s, chimpanzees, and gorillas are more closely related to each other than to the orangutan. The former pongids were reassigned to the subfamily
Hominidae The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the ea ...
("great
ape Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its siste ...
s"), which already included humans, but the details of this reassignment remain contested; within Hominini, not every source excludes gorillas, and not every source includes chimpanzees. Humans are the only extant species in the
Australopithecine Australopithecina or Hominina is a subtribe in the tribe Hominini. The members of the subtribe are generally ''Australopithecus'' (cladistically including the genus, genera ''Homo'', ''Paranthropus'', and ''Kenyanthropus''), and it typically in ...
branch (subtribe), which also contains many extinct close relatives of humans.


Terminology and definition

Concerning membership, when Hominini is taken to exclude ''Pan'', Panini ("panins") may refer to the tribe containing ''Pan'' as its only genus. Or perhaps place ''Pan'' with other
dryopithecine Dryopithecini is an extinct tribe of Eurasian and African great apes that are believed to be close to the ancestry of gorillas, chimpanzees and humans. Members of this tribe are known as dryopithecines. Taxonomy * Tribe Dryopithecini † ** ''K ...
genera, making the whole tribe or subtribe of Panini or Panina together. Minority dissenting nomenclatures include ''Gorilla'' in Hominini and ''Pan'' in ''Homo'' (Goodman et al. 1998), or both ''Pan'' and ''Gorilla'' in ''Homo'' (Watson et al. 2001). By convention, the adjectival term "hominin" (or nominalized "hominins") refers to the tribe Hominini, whereas the members of the subtribe Hominina (and thus all archaic human species) are referred to as "homininian" ("homininians"). This follows the proposal by Mann and Weiss (1996), which presents tribe Hominini as including both ''Pan'' and ''Homo'', placed in separate subtribes. The genus '' Pan'' is referred to subtribe
Panina The genus ''Pan'' consists of two extant species: the chimpanzee and the bonobo. Taxonomically, these two ape species are collectively termed panins; however, both species are more commonly referred to collectively using the generalized term ...
, and genus ''Homo'' is included in the subtribe Hominina (see
below Below may refer to: *Earth *Ground (disambiguation) *Soil *Floor *Bottom (disambiguation) Bottom may refer to: Anatomy and sex * Bottom (BDSM), the partner in a BDSM who takes the passive, receiving, or obedient role, to that of the top or ...
). However, there is an alternative convention which uses "hominin" to exclude members of Panina, i.e. either just for ''Homo'' or for both human and australopithecine species. This alternative convention is referenced in e.g. Coyne (2009) and in Dunbar (2014). Potts (2010) in addition uses the name Hominini in a different sense, as excluding ''Pan'', and uses "hominins" for this, while a separate tribe (rather than subtribe) for chimpanzees is introduced, under the name Panini. In this recent convention, ''contra'' Arambourg, the term "hominin" is applied to ''Homo'', ''Australopithecus'', '' Ardipithecus'', and others that arose after the split from the line that led to chimpanzees (see cladogram below); that is, they distinguish fossil members on the human side of the split, as "hominins", from those on the chimpanzee side, as "not hominins" (or "non-hominin hominids").


Cladogram

This cladogram shows the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
Hominoidea and its descendent clades, focused on the division of Hominini (omitting detail on clades not ancestral to Hominini). The
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Hominidae ("hominids") comprises the
tribes The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to conflic ...
Ponginae (including
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s), Gorillini (including
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
s) and Hominini, the latter two forming the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
of Homininae. Hominini is divided into Panina (
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative th ...
s) and Australopithecina (australopithecines). The Hominina (humans) are usually held to have emerged within the Australopithecina (which would roughly correspond to the alternative definition of Hominini according to the alternative definition which excludes ''Pan''). Genetic analysis combined with fossil evidence indicates that hominoids diverged from the
Old World monkey Old World monkey is the common English name for a family of primates known taxonomically as the Cercopithecidae (). Twenty-four genera and 138 species are recognized, making it the largest primate family. Old World monkey genera include baboons ...
s about 25 million years ago (Mya), near the Oligocene-Miocene boundary. The
most recent common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
s (MRCA) of the subfamilies Homininae and Ponginae lived about 15 million years ago. The most well-known fossil genus of Ponginae is ''Sivapithecus'', consisting of several species from 12.5 million to 8.5 million years ago. It differs from orangutans in dentition and postcranial morphology. In the following cladogram, the approximate time the clades radiated newer clades is indicated in millions of years ago (Mya).


Evolutionary history

Both ''Sahelanthropus'' and ''Orrorin'' existed during the estimated duration of the ancestral chimpanzee–human speciation events, within the range of eight to four million years ago (Mya). Very few fossil specimens have been found that can be considered directly ancestral to genus '' Pan''. News of the first fossil chimpanzee, found in Kenya, was published in 2005. However, it is dated to very recent times—between 545 and 284 thousand years ago. The divergence of a "proto-human" or "pre-human" lineage separate from ''Pan'' appears to have been a process of complex
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
-
hybridization Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...
rather than a clean split, taking place over the period of anywhere between 13 Mya (close to the age of the tribe Hominini itself) and some 4 Mya. Different chromosomes appear to have split at different times, with broad-scale hybridization activity occurring between the two emerging lineages as late as the period 6.3 to 5.4 Mya, according to Patterson et al. (2006), This research group noted that one hypothetical late hybridization period was based in particular on the similarity of
X chromosome The X chromosome is one of the two sex-determining chromosomes (allosomes) in many organisms, including mammals (the other is the Y chromosome), and is found in both males and females. It is a part of the XY sex-determination system and XO sex-d ...
s in the proto-humans and stem chimpanzees, suggesting that the final divergence was even as recent as 4 Mya. Wakeley (2008) rejected these hypotheses; he suggested alternative explanations, including selection pressure on the X chromosome in the ancestral populations prior to the chimpanzee–human last common ancestor (CHLCA). Most DNA studies find that humans and ''Pan'' are 99% identical, but one study found only 94% commonality, with some of the difference occurring in
non-coding DNA Non-coding DNA (ncDNA) sequences are components of an organism's DNA that do not encode protein sequences. Some non-coding DNA is transcribed into functional non-coding RNA molecules (e.g. transfer RNA, microRNA, piRNA, ribosomal RNA, and regul ...
. It is most likely that the australopithecines, dating from 4.4 to 3 Mya, evolved into the earliest members of genus ''Homo''. In the year 2000, the discovery of ''
Orrorin tugenensis ''Orrorin tugenensis'' is a postulated early species of Homininae, estimated at and discovered in 2000. It is not confirmed how ''Orrorin'' is related to modern humans. Its discovery was used to argue against the hypothesis that australopithecin ...
'', dated as early as 6.2 Mya, briefly challenged critical elements of that hypothesis, as it suggested that ''Homo'' did not in fact derive from australopithecine ancestors. All the listed fossil genera are evaluated for: # probability of being ancestral to ''Homo'', and # whether they are more closely related to ''Homo'' than to any other living primate—two traits that could identify them as hominins. Some, including ''
Paranthropus ''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 sy ...
'', '' Ardipithecus'', and ''
Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene and Early Pleistocene. The genus ''Homo'' (which includes modern humans) emerged within ''Australopithecus'', as sister to e.g. ''Australopi ...
'', are broadly thought to be ancestral and closely related to ''Homo''; others, especially earlier genera, including ''
Sahelanthropus ''Sahelanthropus tchadensis'' is an extinct species of the Homininae (African apes) dated to about , during the Miocene epoch. The species, and its genus ''Sahelanthropus'', was announced in 2002, based mainly on a partial cranium, nicknamed ''T ...
'' (and perhaps ''
Orrorin ''Orrorin tugenensis'' is a postulated early species of Homininae, estimated at and discovered in 2000. It is not confirmed how ''Orrorin'' is related to modern humans. Its discovery was used to argue against the hypothesis that australopitheci ...
''), are supported by one community of scientists but doubted by another.


List of known hominin species

''Extant species are in bold.'' *''
Sahelanthropus tchadensis ''Sahelanthropus tchadensis'' is an extinct species of the Homininae (African apes) dated to about , during the Miocene epoch. The species, and its genus ''Sahelanthropus'', was announced in 2002, based mainly on a partial cranium, nicknamed ''T ...
'' *''
Orrorin tugenensis ''Orrorin tugenensis'' is a postulated early species of Homininae, estimated at and discovered in 2000. It is not confirmed how ''Orrorin'' is related to modern humans. Its discovery was used to argue against the hypothesis that australopithecin ...
'' *''
Ardipithecus kadabba ''Ardipithecus kadabba'' is the scientific classification given to fossil remains "known only from teeth and bits and pieces of skeletal bones", originally estimated to be 5.8 to 5.2 million years old, and later revised to 5.77 to 5.54 million ye ...
'' *'' Ardipithecus ramidus'' *'' Australopithecus anamensis'' *''
Australopithecus afarensis ''Australopithecus afarensis'' is an extinct species of australopithecine which lived from about 3.9–2.9 million years ago (mya) in the Pliocene of East Africa. The first fossils were discovered in the 1930s, but major fossil finds would not ...
'' *''
Australopithecus deyiremeda ''Australopithecus deyiremeda'' is an extinct species of australopithecine from Woranso–Mille, Afar Region, Ethiopia, about 3.5 to 3.3 million years ago during the Pliocene. Because it is known only from three partial jawbones, it is unclea ...
'' *'' Australopithecus garhi'' *''
Kenyanthropus platyops ''Kenyanthropus'' is a hominin genus identified from the Lomekwi site by Lake Turkana, Kenya, dated to 3.3 to 3.2 million years ago during the Middle Pliocene. It contains one species, ''K. platyops'', but may also include the 2 million year ol ...
'' *''
Australopithecus africanus ''Australopithecus africanus'' is an extinct species of australopithecine which lived between about 3.3 and 2.1 million years ago in the Late Pliocene to Early Pleistocene of South Africa. The species has been recovered from Taung, Sterkfonte ...
'' *''
Australopithecus sediba ''Australopithecus sediba'' is an extinct species of australopithecine recovered from Malapa Cave, Cradle of Humankind, South Africa. It is known from a partial juvenile skeleton, the holotype MH1, and a partial adult female skeleton, the para ...
'' *''
Paranthropus aethiopicus ''Paranthropus aethiopicus'' is an extinct species of robust australopithecine from the Late Pliocene to Early Pleistocene of East Africa about 2.7–2.3 million years ago. However, it is much debated whether or not ''Paranthropus'' is an invali ...
'' *''
Paranthropus boisei ''Paranthropus boisei'' is a species of australopithecine from the Early Pleistocene of East Africa about 2.5 to 1.15 million years ago. The holotype specimen, OH 5, was discovered by palaeoanthropologist Mary Leakey in 1959, and described by h ...
'' *''
Paranthropus robustus ''Paranthropus robustus'' is a species of robustness (morphology), robust australopithecine from the Early Pleistocene, Early and possibly Middle Pleistocene, Middle Pleistocene of the Cradle of Humankind, South Africa, about 2.27 to 0.87 (or, mo ...
'' *''
Pan troglodytes The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of Hominidae, great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close r ...
'' *''
Pan paniscus The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee and less often the dwarf chimpanzee or gracile chimpanzee, is an endangered great ape and one of the two species making up the genus '' Pan,'' the other being the com ...
'' *''
Homo habilis ''Homo habilis'' ("handy man") is an extinct species of archaic human from the Early Pleistocene of East and South Africa about 2.31 million years ago to 1.65 million years ago (mya). Upon species description in 1964, ''H. habilis'' was highly ...
'' *''
Homo rudolfensis ''Homo rudolfensis'' is an extinct species of archaic human from the Early Pleistocene of East Africa about 2 million years ago (mya). Because ''H. rudolfensis'' coexisted with several other hominins, it is debated what specimens can be confide ...
'' *'' Homo ergaster'' *''
Homo erectus ''Homo erectus'' (; meaning "upright man") is an extinct species of archaic human from the Pleistocene, with its earliest occurrence about 2 million years ago. Several human species, such as '' H. heidelbergensis'' and '' H. antecessor' ...
'' *''
Homo antecessor ''Homo antecessor'' (Latin "pioneer man") is an Extinction, extinct species of archaic human recorded in the Spanish Archaeological Site of Atapuerca, Sierra de Atapuerca, a productive archaeological site, from 1.2 to 0.8 million years ago durin ...
'' *'' Homo heidelbergensis'' *'' Homo naledi'' *'' Homo neanderthalensis'' *''
Homo denisova The Denisovans or Denisova hominins ) are an extinct species or subspecies of archaic human that ranged across Asia during the Lower and Middle Paleolithic. Denisovans are known from few physical remains and consequently, most of what is known ...
'' *''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
'' *'' Homo floresiensis'' *'' Homo luzonensis''


See also

*
History of hominoid taxonomy Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its sister g ...
*
Human evolution Human evolution is the evolutionary process within the history of primates that led to the emergence of ''Homo sapiens'' as a distinct species of the hominid family, which includes the great apes. This process involved the gradual development of ...
* Human taxonomy *
Human timeline The timeline of human evolution outlines the major events in the evolutionary lineage of the modern human species, ''Homo sapiens'', throughout the history of life, beginning some 4 billion years ago down to recent evolution within ''H. sapiens ...
*
List of human evolution fossils The following tables give an overview of notable finds of Hominini, hominin fossils and Skeleton, remains relating to human evolution, beginning with the formation of the tribe Hominini (the divergence of the Chimpanzee–human last common ancest ...


References


External links


Human Timeline (Interactive)
Smithsonian,
National Museum of Natural History The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with 7 ...
(August 2016). {{Taxonbar, from=Q107588 Homininae Human evolution Mammal tribes Taxa named by Camille Arambourg Taxa described in 1948