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 all
the great apes. This process involved the gradual development of traits such as human
bipedalism, dexterity and complex language,[1] as well as interbreeding with other hominins (a
tribe of the African hominid subfamily),[2] indicating that human evolution was not linear but
weblike.[3][4][5][6] The study of human evolution involves several scientific disciplines,
including physical and evolutionary anthropology, paleontology, and genetics.[7][8]
Primates diverged from other mammals about 85 million years ago (mya), in the Late
Cretaceous period, with their earliest fossils appearing over 55 mya, during the Paleocene.
[9]
 Primates produced successive clades leading to the ape superfamily, which gave rise to the
hominid and the gibbon families; these diverged some 15–20 mya. African and Asian
hominids (including orangutans) diverged about 14 mya. Hominins (including
the Australopithecine and Panina subtribes) parted from the Gorillini tribe (gorillas) between 8–
9 mya; Australopithecine (including the extinct biped ancestors of humans) separated from
the Pan genus (containing chimpanzees and bonobos) 4–7 mya.[10] The Homo genus is
evidenced by the appearance of H.  habilis over 2 mya,[a] while anatomically modern
humans emerged in Africa approximately 300,000 years ago.

Before Homo[edit]
For evolutionary history before primates, see Evolution of mammals, History of life, and Timeline
of human evolution.

Early evolution of primates[edit]

See also: Evolution of primates
The evolutionary history of primates can be traced back 65 million years.[11][12][13][14][15] One of the
oldest known primate-like mammal species, the Plesiadapis, came from North America;[16][17][18][19][20]
[21]
 another, Archicebus, came from China.[22] Other similar basal primates were widespread in
Eurasia and Africa during the tropical conditions of the Paleocene and Eocene.

Notharctus tenebrosus, American Museum of Natural History, New York

David R. Begun[23] 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 lemurs of Madagascar, lorises of Southeast Asia, galagos or "bush babies" of Africa, and to
the anthropoids, which are the 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 uppermost Oligocene at Eragaleit in the
northern Great Rift Valley in Kenya, dated to 24 million years ago.[24] Its ancestry is thought to be
species related to Aegyptopithecus, Propliopithecus, and Parapithecus from the Faiyum, at
around 35 mya.[25] In 2010, Saadanius was described as a close relative of the last common
ancestor of the crown catarrhines, and tentatively dated to 29–28 mya, helping to fill an 11-
million-year gap in the fossil record.[26]
Reconstructed tailless Proconsul skeleton

In the Early Miocene, about 22 million years ago, the many kinds of arboreally adapted primitive
catarrhines from East Africa suggest a long history of prior diversification. Fossils at 20 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 million years ago
are Proconsul, Rangwapithecus, Dendropithecus, Limnopithecus, Nacholapithecus, Equatorius, 
Nyanzapithecus, Afropithecus, Heliopithecus, and Kenyapithecus, all from East Africa.
The presence of other generalized non-cercopithecids of Middle Miocene from sites far distant—
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 million years ago.
Molecular evidence indicates that the lineage of gibbons diverged from the line of great apes
some 18–12 mya, and that of orangutans (subfamily Ponginae)[b] diverged from the other great
apes at about 12 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 mya.[27]
Hominidae subfamily Homininae (African hominids) diverged from Ponginae (orangutans) about
14 mya. Hominins (including humans and the Australopithecine and Panina subtribes) parted
from the Gorillini tribe (gorillas) between 8–9 mya; Australopithecine (including the extinct biped
ancestors of humans) separated from the Pan genus (containing chimpanzees and bonobos) 4–
7 mya.[10] The Homo genus is evidenced by the appearance of H. habilis over 2 mya,[a] while
anatomically modern humans emerged in Africa approximately 300,000 years ago.

Divergence of the human clade from other great apes[edit]

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 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 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.[28] 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 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"[29] 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:
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.[28]: 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 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[edit]
Main article: Australopithecus

Reconstruction of "Lucy"

The genus Australopithecus evolved in eastern Africa around 4 million years ago before

spreading throughout the continent and eventually becoming extinct 2 million years ago. During
this time period various forms of australopiths existed, including Australopithecus
anamensis, Au. afarensis, Au. sediba, and Au. africanus. There is still some debate among
academics whether certain African hominid species of this time, such as Au. robustus and Au.
boisei, constitute members of the same genus; if so, they would be considered to be Au. robust
australopiths whilst the others would be considered Au. 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 Au. anamensis, Au. afarensis, Au.
africanus, Au. bahrelghazali, Au. garhi, and Au. sediba;
 Kenyanthropus (3–2.7 Ma), with species K. platyops;
 Paranthropus (3–1.2 Ma), with species P. aethiopicus, P. boisei, and P. robustus
A new proposed species Australopithecus deyiremeda is claimed to have been discovered living
at the same time period of Au. afarensis. There is debate if Au. deyiremeda is a new species or
is Au. afarensis.[30] Australopithecus prometheus, otherwise known as Little Foot has recently
been dated at 3.67 million years old through a new dating technique, making the
genus Australopithecus as old as afarensis.[31] 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.

Evolution of genus Homo[edit]
Main article: Homo
The earliest documented representative of the genus Homo is Homo habilis, which evolved
around 2.8 million years ago,[32] and is arguably the earliest species for which there is positive
evidence of the use of stone tools. The brains of these early hominins were about the same size
as that of a chimpanzee, although it has been suggested that this was the time in which the
human SRGAP2 gene doubled, producing a more rapid wiring of the frontal cortex. During the
next million years a process of rapid encephalization occurred, and with the arrival of Homo
erectus and Homo ergaster in the fossil record, cranial capacity had doubled to 850 cm3.[33] (Such
an increase in human brain size is equivalent to each generation having 125,000
more neurons than their parents.) It is believed that H. erectus and H. ergaster were the first to
use fire and complex tools, and were the first of the hominin line to leave Africa, spreading
throughout Africa, Asia, and Europe between 1.3 to 1.8 million years ago.

A model of the evolution of the genus Homo over the last 2 million years (vertical axis). The rapid "Out of
Africa" expansion of H. sapiens is indicated at the top of the diagram, with admixture indicated with
Neanderthals, Denisovans, and unspecified archaic African hominins. Late survival of robust
australopithecines (Paranthropus) alongside Homo until 1.2 Mya is indicated in purple.
A model of the phylogeny of H. sapiens during the Middle Paleolithic. The horizontal axis represents
geographic location; the vertical axis represents time in millions of years ago.[34] Homo heidelbergensis is
shown as diverging into Neanderthals, Denisovans and H.  sapiens. With the expansion of H.  sapiens after
200 kya, Neanderthals, Denisovans and unspecified archaic African hominins are shown as again
subsumed into the H.  sapiens lineage. In addition, admixture events in modern African populations are
indicated.

Hominin timeline
This box: 
 view
 talk
 edit
 ← Earlier apes
−10 — ← Gorilla split
– ← Chimpanzee split
Miocene
← Earliest bipedal
−9 — ← Earliest stone tools
–
← Dispersal beyond Africa
← Earliest fire / cooking
−8 — ← Earliest clothes
Pliocene ← Modern humans
–

−7 — H

– o

−6 — Pleistocene m
– i
−5 — Hominini
n
– Nakalipithecus
i
−4 — Ouranopithecus
d
– Oreopithecus
s
−3 —
Sahelanthropus
–
Orrorin
−2 —
Ardipithecus
–
Australopithecus
−1 —
Homo habilis
–
Homo erectus
0 —
Homo bodoensis
Homo sapiens
Neanderthals,Denisovans

(million years ago)

According to the recent African origin of modern humans theory, modern humans evolved in
Africa possibly from H.  heidelbergensis, H.  rhodesiensis or H.  antecessor and migrated out of
the continent some 50,000 to 100,000 years ago, gradually replacing local populations
of H. erectus, Denisova hominins, H.  floresiensis, H.  luzonensis and H.  neanderthalensis.[35][36][37][38]
[39]
 Archaic Homo sapiens, the forerunner of anatomically modern humans, evolved in the Middle
Paleolithic between 400,000 and 250,000 years ago.[40][41][42] Recent DNA evidence suggests that
several haplotypes of Neanderthal origin are present among all non-African populations, and
Neanderthals and other hominins, such as Denisovans, may have contributed up to 6% of
their genome to present-day humans, suggestive of a limited interbreeding between these
species.[43][44][45] The transition to behavioral modernity with the development of symbolic culture,
language, and specialized lithic technology happened around 50,000 years ago, according to
some anthropologists,[46] although others point to evidence that suggests that a gradual change in
behavior took place over a longer time span.[47]
Homo sapiens is the only extant species of its genus, Homo. While some (extinct) Homo species
might have been ancestors of Homo sapiens, many, perhaps most, were likely "cousins",
having speciated away from the ancestral hominin line.[48][49] There is yet no consensus as to
which of these groups should be considered a separate species and which should be a
subspecies; this may be due to the dearth of fossils or to the slight differences used to classify
species in the genus Homo.[49] The Sahara pump theory (describing an occasionally
passable "wet" Sahara desert) provides one possible explanation of the early variation in the
genus Homo.
Based on archaeological and paleontological evidence, it has been possible to infer, to some
extent, the ancient dietary practices[50] of various Homo species and to study the role of diet in
physical and behavioral evolution within Homo.[51][52][53][54][55]
Some anthropologists and archaeologists subscribe to the Toba catastrophe theory, which posits
that the supereruption of Lake Toba on Sumatran island in Indonesia some 70,000 years ago
caused global consequences,[56] killing the majority of humans and creating a population
bottleneck that affected the genetic inheritance of all humans today.[57] The genetic and
archaeological evidence for this remains in question however.[58]

H. habilis and H. gautengensis[edit]
Homo habilis lived from about 2.8[32] to 1.4 Ma. The species evolved in South and East Africa in
the Late Pliocene or Early Pleistocene, 2.5–2 Ma, when it diverged from the australopithecines
with the development of smaller molars and larger brains. One of the first known hominins, it
made tools from stone and perhaps animal bones, leading to its name homo habilis (Latin 'handy
man') bestowed by discoverer Louis Leakey. Some scientists have proposed moving this species
from Homo into Australopithecus due to the morphology of its skeleton being more adapted
to living in trees rather than walking on two legs like later hominins.[59]
In May 2010, a new species, Homo gautengensis, was discovered in South Africa.[60]

H. rudolfensis and H. georgicus[edit]
These are proposed species names for fossils from about 1.9–1.6 Ma, whose relation to Homo
habilis is not yet clear.

 Homo rudolfensis refers to a single, incomplete skull from Kenya. Scientists have

suggested that this was a specimen of Homo habilis, but this has not been
confirmed.[61]
 Homo georgicus, from Georgia, may be an intermediate form between Homo
habilis and Homo erectus,[62] or a subspecies of Homo erectus.[63]
H. ergaster and H. erectus[edit]
The first fossils of Homo erectus were discovered by Dutch physician Eugene Dubois in 1891 on
the Indonesian island of Java. He originally named the material Anthropopithecus erectus (1892–
1893, considered at this point as a chimpanzee-like fossil primate)
and Pithecanthropus erectus (1893–1894, changing his mind as of based on its morphology,
which he considered to be intermediate between that of humans and apes).[64] Years later, in the
20th century, the German physician and paleoanthropologist Franz Weidenreich (1873–1948)
compared in detail the characters of Dubois' Java Man, then named Pithecanthropus erectus,
with the characters of the Peking Man, then named Sinanthropus pekinensis. Weidenreich
concluded in 1940 that because of their anatomical similarity with modern humans it was
necessary to gather all these specimens of Java and China in a single species of the
genus Homo, the species H. erectus.[65][66]
Homo erectus lived from about 1.8 Ma to about 70,000 years ago – which would indicate that
they were probably wiped out by the Toba catastrophe; however, nearby H.  floresiensis survived
it. The early phase of H. erectus, from 1.8 to 1.25 Ma, is considered by some to be a separate
species, H. ergaster, or as H. erectus ergaster, a subspecies of H. erectus. Many
paleoanthropologists now use the term Homo ergaster for the non-Asian forms of this group, and
reserve H. erectus only for those fossils that are found in Asia and meet certain skeletal and
dental requirements which differ slightly from H. ergaster.
In Africa in the Early Pleistocene, 1.5–1 Ma, some populations of Homo habilis are thought to
have evolved larger brains and to have made more elaborate stone tools; these differences and
others are sufficient for anthropologists to classify them as a new species, Homo erectus—in
Africa.[67] The evolution of locking knees and the movement of the foramen magnum are thought
to be likely drivers of the larger population changes. This species also may have used fire to
cook meat. Richard Wrangham notes that Homo seems to have been ground dwelling, with
reduced intestinal length, smaller dentition, and "brains [swollen] to their current, horrendously
fuel-inefficient size",[68] and hypothesizes that control of fire and cooking, which released
increased nutritional value, was the key adaptation that separated Homo from tree-sleeping
Australopithecines.[69]
See also: Control of fire by early humans

H. cepranensis and H. antecessor[edit]
These are proposed as species intermediate between H. erectus and H. heidelbergensis.

 H. antecessor is known from fossils from Spain and England that are dated 1.2 Ma–
500 ka.[70][71]
 H. cepranensis refers to a single skull cap from Italy, estimated to be about 800,000
years old.[72]
H. heidelbergensis[edit]
Main article: Homo heidelbergensis
H. heidelbergensis ("Heidelberg Man") lived from about 800,000 to about 300,000 years ago.
Also proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus.[73]

H. rhodesiensis, and the Gawis cranium[edit]

 H. rhodesiensis, estimated to be 300,000–125,000 years old. Most current
researchers place Rhodesian Man within the group of Homo heidelbergensis, though
other designations such as archaic Homo sapiens and Homo sapiens
rhodesiensis have been proposed.
 In February 2006 a fossil, the Gawis cranium, was found which might possibly be a
species intermediate between H. erectus and H. sapiens or one of many
evolutionary dead ends. The skull from Gawis, Ethiopia, is believed to be 500,000–
250,000 years old. Only summary details are known, and the finders have not yet
released a peer-reviewed study. Gawis man's facial features suggest its being either
an intermediate species or an example of a "Bodo man" female.[74]
Neanderthal and Denisovan[edit]
Main articles: Neanderthal and Denisovan
Reconstruction of an elderly Neanderthal man

Homo neanderthalensis, alternatively designated as Homo sapiens neanderthalensis,[75] lived in

Europe and Asia from 400,000[76] to about 28,000 years ago.[77] There are a number of clear
anatomical differences between anatomically modern humans (AMH) and Neanderthal
specimens, many relating to the superior Neanderthal adaptation to cold environments.
Neanderthal surface to volume ratio was even lower than that among modern Inuit populations,
indicating superior retention of body heat.
Neanderthals also had significantly larger brains, as shown from brain endocasts, casting doubt
on their intellectual inferiority to modern humans. However, the higher body mass of
Neanderthals may have required larger brain mass for body control.[78] Also, recent research by
Pearce, Stringer, and Dunbar has shown important differences in brain architecture. The larger
size of the Neanderthal orbital chamber and occipital lobe suggests that they had a better visual
acuity than modern humans, useful in the dimmer light of glacial Europe.
Neanderthals may have had less brain capacity available for social functions. Inferring social
group size from endocranial volume (minus occipital lobe size) suggests that Neanderthal groups
may have been limited to 120 individuals, compared to 144 possible relationships for modern
humans. Larger social groups could imply that modern humans had less risk of inbreeding within
their clan, trade over larger areas (confirmed in the distribution of stone tools), and faster spread
of social and technological innovations. All these may have all contributed to modern Homo
sapiens replacing Neanderthal populations by 28,000 BP.[78]
Earlier evidence from sequencing mitochondrial DNA suggested that no significant gene flow
occurred between H. neanderthalensis and H. sapiens, and that the two were separate species
that shared a common ancestor about 660,000 years ago.[79][80][81] However, a sequencing of the
Neanderthal genome in 2010 indicated that Neanderthals did indeed interbreed with anatomically
modern humans c. 45,000-80,000 years ago, around the time modern humans migrated out from
Africa, but before they dispersed throughout Europe, Asia and elsewhere.[82] The genetic
sequencing of a 40,000-year-old human skeleton from Romania showed that 11% of its genome
was Neanderthal, implying the individual had a Neanderthal ancestor 4–6 generations
previously,[83] in addition to a contribution from earlier interbreeding in the Middle East. Though
this interbred Romanian population seems not to have been ancestral to modern humans, the
finding indicates that interbreeding happened repeatedly.[84]
All modern non-African humans have about 1% to 4% (or 1.5% to 2.6% by more recent data) of
their DNA derived from Neanderthals.[85][82][86] This finding is consistent with recent studies
indicating that the divergence of some human alleles dates to one Ma, although this
interpretation has been questioned.[87][88] Neanderthals and AMH Homo sapiens could have co-
existed in Europe for as long as 10,000 years, during which AMH populations exploded, vastly
outnumbering Neanderthals, possibly outcompeting them by sheer numbers.[89]
In 2008, archaeologists working at the site of Denisova Cave in the Altai
Mountains of Siberia uncovered a small bone fragment from the fifth finger of a juvenile member
of another human species, the Denisovans.[90] Artifacts, including a bracelet, excavated in the
cave at the same level were carbon dated to around 40,000 BP. As DNA had survived in the
fossil fragment due to the cool climate of the Denisova Cave, both mtDNA and nuclear DNA were
sequenced.[43][91]
While the divergence point of the mtDNA was unexpectedly deep in time,[92] the full genomic
sequence suggested the Denisovans belonged to the same lineage as Neanderthals, with the
two diverging shortly after their line split from the lineage that gave rise to modern humans.
[43]
 Modern humans are known to have overlapped with Neanderthals in Europe and the Near
East for possibly more than 40,000 years,[93] and the discovery raises the possibility that
Neanderthals, Denisovans, and modern humans may have co-existed and interbred. The
existence of this distant branch creates a much more complex picture of humankind during
the Late Pleistocene than previously thought.[91][94] Evidence has also been found that as much as
6% of the DNA of some modern Melanesians derive from Denisovans, indicating limited
interbreeding in Southeast Asia.[95][96]
Alleles thought to have originated in Neanderthals and Denisovans have been identified at
several genetic loci in the genomes of modern humans outside Africa. HLA haplotypes from
Denisovans and Neanderthal represent more than half the HLA alleles of modern Eurasians,
[45]
 indicating strong positive selection for these introgressed alleles. Corinne Simoneti at
Vanderbilt University, in Nashville and her team have found from medical records of 28,000
people of European descent that the presence of Neanderthal DNA segments may be associated
with a higher rate of depression.[97]
The flow of genes from Neanderthal populations to modern humans was not all one way. Sergi
Castellano of the Max Planck Institute for Evolutionary Anthropology reported in 2016 that while
Denisovan and Neanderthal genomes are more related to each other than they are to us,
Siberian Neanderthal genomes show more similarity to modern human genes than do European
Neanderthal populations. This suggests Neanderthal populations interbred with modern humans
around 100,000 years ago, probably somewhere in the Near East.[98]
Studies of a Neanderthal child at Gibraltar show from brain development and tooth eruption that
Neanderthal children may have matured more rapidly than Homo sapiens.[99]

H. floresiensis[edit]
Main article: Homo floresiensis

A facial reconstruction of Homo floresiensis

H. floresiensis, which lived from approximately 190,000 to 50,000 years before present (BP), has
been nicknamed the hobbit for its small size, possibly a result of insular dwarfism.[100] H.
floresiensis is intriguing both for its size and its age, being an example of a recent species of the
genus Homo that exhibits derived traits not shared with modern humans. In other words, H.
floresiensis shares a common ancestor with modern humans, but split from the modern human
lineage and followed a distinct evolutionary path. The main find was a skeleton believed to be a
woman of about 30 years of age. Found in 2003, it has been dated to approximately 18,000
years old. The living woman was estimated to be one meter in height, with a brain volume of just
380 cm3 (considered small for a chimpanzee and less than a third of the H. sapiens average of
1400 cm3).[100]
However, there is an ongoing debate over whether H. floresiensis is indeed a separate species.
[101]
 Some scientists hold that H. floresiensis was a modern H. sapiens with pathological dwarfism.
[102]
 This hypothesis is supported in part, because some modern humans who live on Flores, the
Indonesian island where the skeleton was found, are pygmies. This, coupled with pathological
dwarfism, could have resulted in a significantly diminutive human. The other major attack on H.
floresiensis as a separate species is that it was found with tools only associated with H. sapiens.
[102]

The hypothesis of pathological dwarfism, however, fails to explain additional anatomical

features that are unlike those of modern humans (diseased or not) but much like those of ancient
members of our genus. Aside from cranial features, these features include the form of bones in
the wrist, forearm, shoulder, knees, and feet. Additionally, this hypothesis fails to explain the find
of multiple examples of individuals with these same characteristics, indicating they were common
to a large population, and not limited to one individual.[101]
In 2016, fossil teeth and a partial jaw from hominins assumed to be ancestral
to H. floresiensis were discovered[103] at Mata Menge, about 74 km (46 mi) from Liang Bua. They
date to about 700,000 years ago[104] and are noted by Australian archaeologist Gerrit van den
Bergh for being even smaller than the later fossils.[105]

H. luzonensis[edit]
Main article: Homo luzonensis
A small number of specimens from the island of Luzon, dated 50,000 to 67,000 years ago, have
recently been assigned by their discoverers, based on dental characteristics, to a novel human
species, H. luzonensis.[106]

H. sapiens[edit]
Main articles: Archaic humans, Early modern human, Archaic human admixture with modern
humans, and Human §  Evolution
H. sapiens (the adjective sapiens is Latin for "wise" or "intelligent") emerged in Africa around
300,000 years ago, likely derived from H.  heidelbergensis or a related lineage.[107][108] In
September 2019, scientists reported the computerized determination, based on 260 CT scans, of
a virtual skull shape of the last common human ancestor to modern humans/H. sapiens,
representative of the earliest modern humans, and suggested that modern humans arose
between 260,000 and 350,000 years ago through a merging of populations in East and South
Africa.[109][110]
Between 400,000 years ago and the second interglacial period in the Middle Pleistocene, around
250,000 years ago, the trend in intra-cranial volume expansion and the elaboration of stone tool
technologies developed, providing evidence for a transition from H. erectus to H. sapiens. The
direct evidence suggests there was a migration of H. erectus out of Africa, then a further
speciation of H. sapiens from H. erectus in Africa. A subsequent migration (both within and out
of Africa) eventually replaced the earlier dispersed H. erectus. This migration and origin theory is
usually referred to as the "recent single-origin hypothesis" or "out of Africa"
theory. H. sapiens interbred with archaic humans both in Africa and in Eurasia, in Eurasia
notably with Neanderthals and Denisovans.[43][95]
The Toba catastrophe theory, which postulates a population bottleneck for H. sapiens about
70,000 years ago,[111] was controversial from its first proposal in the 1990s and by the 2010s had
very little support.[112] Distinctive human genetic variability has arisen as the result of the founder
effect, by archaic admixture and by recent evolutionary pressures.

Anatomical changes[edit]
Since Homo sapiens separated from its last common ancestor shared with chimpanzees, human
evolution is characterized by a number
of morphological, developmental, physiological, behavioral, and environmental changes.
[8]
 Environmental (cultural) evolution discovered much later during the Pleistocene played a
significant role in human evolution observed via human transitions between subsistence
systems.[113][8] The most significant of these adaptations are bipedalism, increased brain size,
lengthened ontogeny (gestation and infancy), and decreased sexual dimorphism. The
relationship between these changes is the subject of ongoing debate.[114] Other significant
morphological changes included the evolution of a power and precision grip, a change first
occurring in H.  erectus.[115]

Bipedalism[edit]

Bipedalism shown by a man and a woman

Bipedalism is the basic adaptation of the hominid and is considered the main cause behind a
suite of skeletal changes shared by all bipedal hominids. The earliest hominin, of presumably
primitive bipedalism, is considered to be either Sahelanthropus[116] or Orrorin, both of which arose
some 6 to 7 million years ago. The non-bipedal knuckle-walkers, the gorillas and chimpanzees,
diverged from the hominin line over a period covering the same time, so
either Sahelanthropus or Orrorin may be our last shared ancestor. Ardipithecus, a full biped,
arose approximately 5.6 million years ago.[117]
The early bipeds eventually evolved into the australopithecines and still later into the
genus Homo. There are several theories of the adaptation value of bipedalism. It is possible that
bipedalism was favored because it freed the hands for reaching and carrying food, saved energy
during locomotion,[118] enabled long-distance running and hunting, provided an enhanced field of
vision, and helped avoid hyperthermia by reducing the surface area exposed to direct sun;
features all advantageous for thriving in the new savanna and woodland environment created as
a result of the East African Rift Valley uplift versus the previous closed forest habitat.[118][119][120] A
2007 study provides support for the hypothesis that walking on two legs, or bipedalism, evolved
because it used less energy than quadrupedal knuckle-walking.[121][122] However, recent studies
suggest that bipedality without the ability to use fire would not have allowed global dispersal.
[123]
 This change in gait saw a lengthening of the legs proportionately when compared to the
length of the arms, which were shortened through the removal of the need for brachiation.
Another change is the shape of the big toe. Recent studies suggest that australopithecines still
lived part of the time in trees as a result of maintaining a grasping big toe. This was progressively
lost in habilines.
Anatomically, the evolution of bipedalism has been accompanied by a large number of skeletal
changes, not just to the legs and pelvis, but also to the vertebral column, feet and ankles, and
skull.[124] The femur evolved into a slightly more angular position to move the center of gravity
toward the geometric center of the body. The knee and ankle joints became increasingly robust
to better support increased weight. To support the increased weight on each vertebra in the
upright position, the human vertebral column became S-shaped and the lumbar
vertebrae became shorter and wider. In the feet the big toe moved into alignment with the other
toes to help in forward locomotion. The arms and forearms shortened relative to the legs making
it easier to run. The foramen magnum migrated under the skull and more anterior.[125]
The most significant changes occurred in the pelvic region, where the long downward facing iliac
blade was shortened and widened as a requirement for keeping the center of gravity stable while
walking;[27] bipedal hominids have a shorter but broader, bowl-like pelvis due to this. A drawback
is that the birth canal of bipedal apes is smaller than in knuckle-walking apes, though there has
been a widening of it in comparison to that of australopithecine and modern humans, thus
permitting the passage of newborns due to the increase in cranial size. This is limited to the
upper portion, since further increase can hinder normal bipedal movement.[126]
The shortening of the pelvis and smaller birth canal evolved as a requirement for bipedalism and
had significant effects on the process of human birth, which is much more difficult in modern
humans than in other primates. During human birth, because of the variation in size of the pelvic
region, the fetal head must be in a transverse position (compared to the mother) during entry into
the birth canal and rotate about 90 degrees upon exit.[127] The smaller birth canal became a
limiting factor to brain size increases in early humans and prompted a shorter gestation period
leading to the relative immaturity of human offspring, who are unable to walk much before 12
months and have greater neoteny, compared to other primates, who are mobile at a much earlier
age.[120] The increased brain growth after birth and the increased dependency of children on
mothers had a major effect upon the female reproductive cycle,[128] and the more frequent
appearance of alloparenting in humans when compared with other hominids.[129] Delayed human
sexual maturity also led to the evolution of menopause with one explanation, the grandmother
hypothesis, providing that elderly women could better pass on their genes by taking care of their
daughter's offspring, as compared to having more children of their own.[130][131]

Encephalization[edit]

Skulls of successive (or near-successive, depending on the source) human evolutionary ancestors,[c] up
until 'modern' Homo sapiens
* Mya – million years ago, kya – thousand years ago

Brain size and tooth size in hominins

The human species eventually developed a much larger brain than that of other primates—
typically 1,330 cm3 (81 cu in) in modern humans, nearly three times the size of a chimpanzee or
gorilla brain.[134] After a period of stasis with Australopithecus anamensis and Ardipithecus,
species which had smaller brains as a result of their bipedal locomotion,[135] the pattern
of encephalization started with Homo habilis, whose 600 cm3 (37 cu in) brain was slightly larger
than that of chimpanzees. This evolution continued in Homo erectus with 800–1,100 cm3 (49–
67 cu in), and reached a maximum in Neanderthals with 1,200–1,900 cm3 (73–116 cu in), larger
even than modern Homo sapiens. This brain increase manifested during postnatal brain growth,
far exceeding that of other apes (heterochrony). It also allowed for extended periods of social
learning and language acquisition in juvenile humans, beginning as much as 2 million years ago.
Encephalization may be due to a dependency on calorie-dense, difficult-to-acquire food.[136]
Furthermore, the changes in the structure of human brains may be even more significant than
the increase in size.[137][138][139][51]