|Two hominins: A human (Homo sapiens) holding a chimpanzee (Pan troglodytes)|
|Tribe:|| Hominini |
Arambourg, 1948 
| Homo |
The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera Homo (humans) and Pan (chimpanzees and bonobos) and in standard usage excludes the genus Gorilla (gorillas).
The term was originally introduced by Camille Arambourg (1948). Arambourg combined the categories of Hominina and Simiina due to Gray (1825) into his new subtribe.
Traditionally, chimpanzees, gorillas and orangutans were grouped together as pongids. Since Gray's classification, evidence has accumulated from genetic phylogeny confirming that humans, chimpanzees, and gorillas are more closely related to each other than to the orangutan.  The former pongids were reassigned to the subfamily Hominidae ("great apes"), 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 branch (subtribe), which also contains many extinct close relatives of humans.
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 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, and genus Homo is included in the subtribe Hominina (see below). 
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"). 
This cladogram shows the clade of superfamily Hominoidea and its descendent clades, focused on the division of Hominini (omitting detail on clades not ancestral to Hominini). The family Hominidae ("hominids") comprises the tribes Ponginae (including orangutans), Gorillini (including gorillas) and Hominini, the latter two forming the subfamily of Homininae. Hominini is divided into Panina (chimpanzees) 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 monkeys about 25 million years ago (Mya), near the Oligocene-Miocene boundary.  The most recent common ancestors (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).
|Hominoidea (20.4 Mya)|
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-hybridization 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 chromosomes 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.  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 , 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:
Some, including Paranthropus , Ardipithecus , and Australopithecus , are broadly thought to be ancestral and closely related to Homo;  others, especially earlier genera, including Sahelanthropus (and perhaps Orrorin ), are supported by one community of scientists but doubted by another.  
Extant species are in bold.
Ardipithecus is a genus of an extinct hominine that lived during the Late Miocene and Early Pliocene epochs in the Afar Depression, Ethiopia. Originally described as one of the earliest ancestors of humans after they diverged from the chimpanzees, the relation of this genus to human ancestors and whether it is a hominin is now a matter of debate. Two fossil species are described in the literature: A. ramidus, which lived about 4.4 million years ago during the early Pliocene, and A. kadabba, dated to approximately 5.6 million years ago. Initial behavioral analysis indicated that Ardipithecus could be very similar to chimpanzees, however more recent analysis based on canine size and lack of canine sexual dimorphism indicates that Ardipithecus was characterised by reduced aggression, and that they more closely resemble bonobos.
Homininae, also called "African hominids" or "African apes", is a subfamily of Hominidae. It includes two tribes, with their extant as well as extinct species: 1) the tribe Hominini ―and 2) the tribe Gorillini (gorillas). Alternatively, the genus Pan is sometimes considered to belong to its own third tribe, Panini. Homininae comprises all hominids that arose after orangutans split from the line of great apes. The Homininae cladogram has three main branches, which lead to gorillas, and to humans and chimpanzees via the tribe Hominini and subtribes Hominina and Panina. There are two living species of Panina and two living species of gorillas, but only one extant human species. Traces of extinct Homo species, including Homo floresiensis have been found with dates as recent as 40,000 years ago. Organisms in this subfamily are described as hominine or hominines.
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 old Homo rudolfensis, or K. rudolfensis. Before its naming in 2001, Australopithecus afarensis was widely regarded as the only australopithecine to exist during the Middle Pliocene, but Kenyanthropus evinces a greater diversity than once acknowledged. Kenyanthropus is most recognisable by an unusually flat face and small teeth for such an early hominin, with values on the extremes or beyond the range of variation for australopithecines in regard to these features. Multiple australopithecine species may have coexisted by foraging for different food items, which may be reason why these apes anatomically differ in features related to chewing.
Orrorin tugenensis is a postulated early species of Homininae, estimated at 6.1 to 5.7 million years ago 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 australopithecines are human ancestors, although this remains the most prevalent hypothesis of human evolution as of 2012.
Paranthropus is a genus of extinct hominin which contains two widely accepted species: P. robustus and P. boisei. However, the validity of Paranthropus is contested, and it is sometimes considered to be 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.
Australopithecus is a genus of early hominins that existed in Africa during the Pliocene and Early Pleistocene. The genera Homo, Paranthropus, and Kenyanthropus evolved from some Australopithecus species. Australopithecus is a member of the subtribe Australopithecina, which sometimes also includes Ardipithecus, though the term "australopithecine" is sometimes used to refer only to members of Australopithecus. Species include A. garhi, A. africanus, A. sediba, A. afarensis, A. anamensis, A. bahrelghazali and A. deyiremeda. Debate exists as to whether some Australopithecus species should be reclassified into new genera, or if Paranthropus and Kenyanthropus are synonymous with Australopithecus, in part because of the taxonomic inconsistency.
Sahelanthropus tchadensis is an extinct species of the Homininae dated to about 7 million years ago, during the Miocene epoch. The species, and its genus Sahelanthropus, was announced in 2002, based mainly on a partial cranium, nicknamed Toumaï, discovered in northern Chad.
Apes are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia, which together with its sister group Cercopithecidae form the catarrhine clade, cladistically making them monkeys. Apes do not have tails due to a mutation of the TBXT gene. In traditional and non-scientific use, the term "ape" can include tailless primates taxonomically considered Cercopithecidae, and is thus not equivalent to the scientific taxon Hominoidea. There are two extant branches of the superfamily Hominoidea: the gibbons, or lesser apes; and the hominids, or great apes.
Pongidae, or the pongids is an obsolete primate taxon containing chimpanzees, gorillas and orangutans. By this definition pongids were also called "great apes". This taxon is not used today but is of historical significance. The great apes are currently classified as Hominidae. This entry addresses the old usage of pongid.
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 take place until the 1970s. From 1972 to 1977, the International Afar Research Expedition—led by anthropologists Maurice Taieb, Donald Johanson and Yves Coppens—unearthed several hundreds of hominin specimens in Hadar, Ethiopia, the most significant being the exceedingly well-preserved skeleton AL 288-1 ("Lucy") and the site AL 333. Beginning in 1974, Mary Leakey led an expedition into Laetoli, Tanzania, and notably recovered fossil trackways. In 1978, the species was first described, but this was followed by arguments for splitting the wealth of specimens into different species given the wide range of variation which had been attributed to sexual dimorphism. A. afarensis probably descended from A. anamensis and is hypothesised to have given rise to Homo, though the latter is debated.
Homo is the genus that emerged in the genus Australopithecus that encompasses the extant species Homo sapiens, plus several extinct species classified as either ancestral to or closely related to modern humans, most notably H. erectus and H. neanderthalensis. The genus emerged with the appearance of H. habilis just over 2 million years ago. Homo, together with the genus Paranthropus, is probably sister to Australopithecus africanus, which itself had previously split from the lineage of Pan, the chimpanzees.
Paleoanthropology or paleo-anthropology is a branch of paleontology and anthropology which seeks to understand the early development of anatomically modern humans, a process known as hominization, through the reconstruction of evolutionary kinship lines within the family Hominidae, working from biological evidence and cultural evidence.
Australopithecus anamensis is a hominin species that lived approximately between 4.2 and 3.8 million years ago and is the oldest known Australopithecus species, living during the Plio-Pleistocene era.
Human taxonomy is the classification of the human species within zoological taxonomy. The systematic genus, Homo, is designed to include both anatomically modern humans and extinct varieties of archaic humans. Current humans have been designated as subspecies Homo sapiens sapiens, differentiated, according to some, from the direct ancestor, Homo sapiens idaltu.
Australopithecina or Hominina is a subtribe in the tribe Hominini. The members of the subtribe are generally Australopithecus, and it typically includes the earlier Ardipithecus, Orrorin, Sahelanthropus, and Graecopithecus. All these closely related species are now sometimes collectively termed australopiths or homininians. They are the extinct, close relatives of humans and, with the extant genus Homo, comprise the human clade. Members of the human clade, i.e. the Hominini after the split from the chimpanzees, are now called Hominina.
The Middle Awash is a paleoanthropological research area in the Afar Region along the Awash River in Ethiopia's Afar Depression. It is a unique natural laboratory for the study of human origins and evolution and a number of fossils of the earliest hominins, particularly of the Australopithecines, as well as some of the oldest known Olduwan stone artifacts, have been found at the site—all of late Miocene, the Pliocene, and the very early Pleistocene times, that is, about 5.6 million years ago (mya) to 2.5 mya. It is broadly thought that the divergence of the lines of the earliest humans (hominins) and of chimpanzees (hominids) was completed near the beginning of that time range, or sometime between seven and five mya. However, the larger community of scientists provide several estimates for periods of divergence that imply a greater range for this event, see CHLCA: human-chimpanzee split.
Ardipithecus ramidus is a species of australopithecine from the Afar region of Early Pliocene Ethiopia 4.4 million years ago (mya). A. ramidus, unlike modern hominids, has adaptations for both walking on two legs (bipedality) and life in the trees (arboreality). However, it would not have been as efficient at bipedality as humans, nor at arboreality as non-human great apes. Its discovery, along with Miocene apes, has reworked academic understanding of the chimpanzee-human last common ancestor from appearing much like modern day chimpanzees, orangutans and gorillas to being a creature without a modern anatomical cognate.
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 ; Gorilla ; Pan ; and Homo, of which only modern humans remain.
The chimpanzee–human last common ancestor (CHLCA) is the last common ancestor shared by the extant Homo (human) and Pan genera of Hominini. Due to complex hybrid speciation, it is not currently possible to give a precise estimate on the age of this ancestral population. While "original divergence" between populations may have occurred as early as 13 million years ago (Miocene), hybridization may have been ongoing until as recently as 4 million years ago (Pliocene).
Changes to the dental morphology and jaw are major elements of hominid evolution. These changes were driven by the types and processing of food eaten. The evolution of the jaw is thought to have facilitated encephalization, speech, and the formation of the uniquely human chin.
However, overwhelming genetic evidence has since demonstrated that humans, chimpanzees, and gorillas are much more closely related to each other than to the orangutan ... Thus, there is no genetic support for grouping the great apes together in a distinct group from humans. For this reason, many researchers now place all species of great ape and human within a single family, Hominidae – making them all proper 'hominids'.
Conventionally, taxonomists now refer to the great ape family (including humans) as 'hominids', while all members of the lineage leading to modern humans that arose after the split with the [Homo-Pan] LCA are referred to as 'hominins'. The older literature used the terms hominoids and hominids respectively.
Anthropologists apply the term hominin to all the species on the "human" side of our family tree after it split from the branch that became modern chimps." (p.197)
Thus human evolution is the study of the lineage, or clade, comprising species more closely related to modern humans than to chimpanzees. Its stem species is the so-called 'common hominin ancestor', and its only extant member is Homo sapiens. This clade contains all the species more closely related to modern humans than to any other living primate. Until recently, these species were all subsumed into a family, Hominidae, but this group is now more usually recognised as a tribe, the Hominini.
Patterson et al. suggest that the apparently short divergence time between humans and chimpanzees on the X chromosome is explained by a massive interspecific hybridization event in the ancestry of these two species. However, Patterson et al. do not statistically test their own null model of simple speciation before concluding that speciation was complex, and—even if the null model could be rejected—they do not consider other explanations of a short divergence time on the X chromosome. These include natural selection on the X chromosome in the common ancestor of humans and chimpanzees, changes in the ratio of male-to-female mutation rates over time, and less extreme versions of divergence with gene flow. I therefore believe that their claim of hybridization is unwarranted.
After A. afarensis, the fossil record shows a confusing melange of gracile australopithecine species lasting up to about two million years ago. … [T]he late australopithecines, already bipedal, were beginning to show changes in teeth, skull, and brain that presage modern humans. It is very likely that the lineage that gave rise to modern humans included at least one of these species.
The discovery of Orrorin has ... radically modified interpretations of human origins and the environmental context in which the African apes/hominoid transition occurred, although ... the less likely hypothesis of derivation of Homo from the australopithecines still holds primacy in the minds of most palaeoanthropologists.
Sahelanthropus is the oldest and most primitive known member of the hominid clade, close to the divergence of hominids and chimpanzees.
Sahelanthropus tchadensis is an enigmatic new Miocene species, whose characteristics are a mix of those of apes and Homo erectus and which has been proclaimed by Brunet et al. to be the earliest hominid. However, we believe that features of the dentition, face and cranial base that are said to define unique links between this Toumaï specimen and the hominid clade are either not diagnostic or are consequences of biomechanical adaptations. To represent a valid clade, hominids must share unique defining features, and Sahelanthropus does not appear to have been an obligate biped.