(? =more field data needed)
(† =extinct =fossil)
Hominini ("hominins") form a taxonomic tribe of the subfamily
Hominini includes genus
Homo (humans), but
Gorilla (gorillas). There is at present (as of
2018[update]) no consensus on whether it should include genus Pan (the
chimpanzees), the question being closely tied to the complex
speciation process connecting humans and chimpanzees and the
development of bipedalism in proto-humans.
The tribe was originally introduced by Gray (1824), long before any
details on the speciation of Pan and
Homo were known. Gray's tribe
Hominini by definition includes both Pan and Homo. This definition is
still adhered to in the proposal by Mann and Weiss (1996), which
Hominini into three subtribes,
Panina (containing Pan),
Hominina ("homininans", containing
Homo "humans"), and
Australopithecina (containing several extinct "australopithecine"
Hominini is taken to exclude Pan. In this case, Panini
("panins", Delson 1977) may be used to refer to the tribe
containing Pan as its only genus.
Minority dissenting views include
Hominini and Pan in Homo
(Goodman et al. 1998), or both Pan and
Homo (Watson et al.
1 Terminology and definition
3 Evolutionary history
4 See also
6 External links
Terminology and definition
By convention, the adjectival term "hominin" (or nominalized
"hominins") refers to the tribe Hominini, while the members of the
Hominina subtribe (and thus all archaic human species) are referred to
as "hominan" ("hominans"). 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
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
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 Gray, the term hominin is
applied all species of genus Homo, as well as to species of the
ancestral genera 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
This cladogram shows the clade of superfamily
Hominoidea and its
descendent clades, focussed on the division of
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
Australopithecina (australopithicenes). 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. In the following cladogram, the
approximate time the clades radiated newer clades indicated in
millions of years ago (Mya).
Hominoidea (20.4 Mya)
Paranthropus robustus (†2)
Australopithecus garhi (†2.5)
Further information: Chimpanzee–human last common ancestor
Model of the phylogeny of
Hominini over the past 10 million years.
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 a of "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 million years ago (close to the age of the
itself) and some 4 million years ago. 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 the final divergence 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).
DNA studies find that humans and Pan are 99% identical,
but one study found only 94% commonality, with some of the difference
occurring in noncoding DNA.
It is most likely that the australopithecines, dating from 3 to 4.4
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
All the listed fossil genera are evaluated for: 1) probability of
being ancestral to Homo, and 2) whether they are more closely related
Homo than to any other living primate—two traits that could
identify them as hominins. 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.
History of hominoid taxonomy
List of human evolution fossils
List of human evolution fossils (with images)
^ Fuss, J; Spassov, N; Begun, DR; Böhme, M (2017). "Potential hominin
Graecopithecus from the Late Miocene of Europe". PLoS
ONE. 12 (5).
^ a b Mann, Alan; Mark Weiss (1996). "Hominoid Phylogeny and Taxonomy:
a consideration of the molecular and Fossil Evidence in an Historical
Perspective". Molecular Phylogenetics and Evolution. 5 (1): 169–181.
doi:10.1006/mpev.1996.0011. PMID 8673284.
^ Delson, Journal of
Human Evolution 6 (1977), p. 450.
^ a b Potts (2010). “What Does It Mean to Be Human?”, pp. 34.
ISBN 978-1-4262-0606-1. National Geographic Society, Washington.
^ a b "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."Dunbar, Robin (2014).
Human evolution. ISBN 9780141975313.
^ B. Wood (2010). "Reconstructing human evolution: Achievements,
challenges, and opportunities". Proceedings of the National Academy of
Sciences. 107: 8902–8909. Bibcode:2010PNAS..107.8902W.
doi:10.1073/pnas.1001649107. PMC 3024019 .
^ a b Coyne, Jerry A. (2009) Why Evolution Is True, pp.197-208, 244,
248. ISBN 978-0-670-02053-9(hc),
ISBN 978-0-14-311664-6(pbk). Penguin Books Ltd, London.
"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)
^ Bradley, B. J. (2006). "Reconstructing Phylogenies and Phenotypes: A
Molecular View of
Human Evolution". Journal of Anatomy. 212 (4):
337–353. doi:10.1111/j.1469-7580.2007.00840.x. PMC 2409108 .
^ Wood and Richmond.; Richmond, BG (2000). "
Human evolution: taxonomy
and paleobiology". Journal of Anatomy. 197 (Pt 1): 19–60.
doi:10.1046/j.1469-7580.2000.19710019.x. PMC 1468107 .
PMID 10999270. 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.
^ "Fossils May Pinpoint Critical Split Between Apes and Monkeys".
redOrbit.com. 15 May 2013.
^ 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. >Taylor, C. (2011). "Old men of the woods". Palaeos.
^ McBrearty, Sally; Nina G. Jablonski (2005). "First fossil
chimpanzee". Nature. 437 (7055): 105–108.
^ Patterson N, Richter DJ, Gnerre S, Lander ES, Reich D (June 2006).
"Genetic evidence for complex speciation of humans and chimpanzees".
Nature. 441 (7097): 1103–8. Bibcode:2006Natur.441.1103P.
doi:10.1038/nature04789. PMID 16710306.
^ Wakeley J (March 2008). "Complex speciation of humans and
chimpanzees". Nature. 452 (7184): E3–4; discussion E4.
PMID 18337768. "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."
^ Mary-Claire King (1973) Protein polymorphisms in chimpanzee and
human evolution, Doctoral dissertation, University of California,
^ Wong, Kate (1 September 2014). "Tiny Genetic Differences between
Humans and Other Primates Pervade the Genome". Scientific
^ Minkel JR (2006-12-19). "Humans and Chimps: Close But Not That
Close". Scientific American.
^ Coyne, Jerry A. (2009) Why Evolution Is True, pp.202-204.
ISBN 978-0-670-02053-9(hc), ISBN 978-0-14-311664-6(pbk).
Penguin Books Ltd, London. "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
^ Cameron, D. W. (2003). "Early hominin speciation at the
Plio/Pleistocene transition". HOMO - Journal of Comparative Human
Biology. 54 (1): 1–28. doi:10.1078/0018-442x-00057.
^ Potts, Richard and Sloan, Christopher. “What Does It Mean to Be
Human?”, pp. 38-39. ISBN 978-1-4262-0606-1. National Geographic
^ Reynolds, Sally C; Gallagher, Andrew (2012-03-29). African Genesis:
Perspectives on Hominin Evolution. ISBN 9781107019959. :
"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."
^ Potts, Richard and Sloan, Christopher. “What Does It Mean to Be
Human?”, pp. 31-42. ISBN 978-1-4262-0606-1. National Geographic
^ Brunet, Michel; Guy, F; Pilbeam, D; MacKaye, H. T.; Likius, A;
Ahounta, D; Beauvilain, A; Blondel, C; Bocherens, H; Boisserie, JR; De
Bonis, L; Coppens, Y; Dejax, J; Denys, C; Duringer, P; Eisenmann, V;
Fanone, G; Fronty, P; Geraads, D; Lehmann, T; Lihoreau, F; Louchart,
A; Mahamat, A; Merceron, G; Mouchelin, G; Otero, O; Pelaez Campomanes,
P; Ponce De Leon, M; Rage, J. C.; et al. (July 2002), "A new hominid
from the Upper Miocene of Chad, Central Africa", Nature, 418 (6894):
145–151, doi:10.1038/nature00879, PMID 12110880, Sahelanthropus
is the oldest and most primitive known member of the hominid clade,
close to the divergence of hominids and chimpanzees.
^ Wolpoff, Milford; Senut, Brigitte; Pickford, Martin; Hawks, John
(October 2002), "
Sahelanthropus or 'Sahelpithecus'?", Nature, 419
(6907): 581–582, Bibcode:2002Natur.419..581W, doi:10.1038/419581a,
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.
Yohannes Haile-Selassiea,b,1, Stephanie M. Melilloc, and Denise F.
Sud, "The Pliocene hominin diversity conundrum: Do more fossils mean
less clarity?" PNAS June 7, 2016 vol. 113 no. 23, 6364-6371, doi:
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