From Wikipedia, the free encyclopedia
Human evolution is the
evolutionary process that led to the emergence of
anatomically modern humans, beginning with the
evolutionary history of
primates – in particular
genus Homo – and leading to the emergence of
Homo sapiens as a distinct species of the
hominid family, the great apes. This process involved the gradual development of traits such as
human bipedalism and
language.
[1]
The study of human evolution involves many scientific disciplines, including
physical anthropology,
primatology,
archaeology,
paleontology,
neurobiology,
ethology,
linguistics,
evolutionary psychology,
embryology and
genetics.
[2] Genetic studies show that primates diverged from other
mammals about
85 million years ago, in the
Late Cretaceous period, and the earliest
fossils appear in the
Paleocene, around
55 million years ago.
[3]
Within the
Hominoidea (apes) superfamily, the
Hominidae family diverged from the
Hylobatidae (gibbon) family some 15–20 million years ago; African great apes (subfamily
Homininae) diverged from
orangutans (
Ponginae) about
14 million years ago; the
Hominini tribe (humans,
Australopithecines and other extinct biped genera, and
chimpanzee) parted from the
Gorillini tribe (gorillas) between
9 million years ago and
8 million years ago; and, in turn, the subtribes
Hominina (humans and biped ancestors) and
Panina (chimps) separated about
7.5 million years ago to
4 million years ago.
[4]
Anatomical changes
Human evolution from its first separation from the
last common ancestor of humans and chimpanzees is characterized by a number of
morphological,
developmental,
physiological, and
behavioral changes. 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.
[5][page needed] Other significant morphological changes included the evolution of a
power and precision grip, a change first occurring in
H. erectus.
[6]
Bipedalism
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[7] or
Orrorin,
both of which arose some 6 to 7 million years ago. The non-bipedal
knuckle-walkers, the gorilla and chimpanzee, diverged from the hominin
line over a period covering the same time, so either of
Sahelanthropus or
Orrorin may be our
last shared ancestor.
Ardipithecus, a full biped, arose somewhat later.
[citation needed]
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,
[8]
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.
[9][8][10]
A new study provides support for the hypothesis that walking on two
legs, or bipedalism, evolved because it used less energy than
quadrupedal knuckle-walking.
[11][12] However, recent studies suggest that bipedality without the
ability to use fire would not have allowed global dispersal.
[13]
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.
[14] 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.
[15]
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;
[16]
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, permitting
the passage of newborns due to the increase in cranial size but this is
limited to the upper portion, since further increase can hinder normal
bipedal movement.
[17]
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.
[18]
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.
[10]
The increased brain growth after birth and the increased dependency of
children on mothers had a big effect upon the female reproductive cycle,
[19] and the more frequent appearance of
alloparenting in humans when compared with other hominids.
[20] Delayed human sexual maturity also led to the evolution of
menopause
with one explanation 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.
[21]
Encephalization
Brain size and tooth size in hominins.
The human species eventually developed a much larger brain than that of other primates—typically 1,330 cm
3 (81 cu in) in modern humans, nearly three times the size of a chimpanzee or gorilla brain.
[22] After a period of stasis with
Australopithecus anamensis and
Ardipithecus, species which had smaller brains as a result of their bipedal locomotion,
[23] the pattern of
encephalization started with
Homo habilis, whose 600 cm
3 (37 cu in) brain was slightly larger than that of chimpanzees. This evolution continued in
Homo erectus with 800–1,100 cm
3 (49–67 cu in), and reached a maximum in Neanderthals with 1,200–1,900 cm
3 (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.
Furthermore, the changes in the structure of
human brains may be even more significant than the increase in size.
[24][25][26][27]
The size and shape of the skull changed over time. The leftmost, and largest, is a replica of a modern human skull.
The
temporal lobes, which contain centers for language processing, have increased disproportionately, as has the
prefrontal cortex, which has been related to complex decision-making and moderating social behavior.
[22] Encephalization has been tied to increased meat in the diet,
[28][29][30] and the development of cooking,
[31] and it has been proposed that intelligence increased as a response to an increased necessity for
solving social problems as human society became more complex.
[32]
Changes in skull morphology, such as smaller mandibles and mandible
muscle attachments, allowed more room for the brain to grow.
[33]
The increase in volume of the
neocortex also included a rapid increase in size of the
cerebellum. Its function has traditionally been associated with balance and fine motor control, but more recently with
speech and
cognition.
The great apes, including hominids, had a more pronounced cerebellum
relative to the neocortex than other primates. It has been suggested
that because of its function of sensory-motor control and learning
complex muscular actions, the cerebellum may have underpinned human
technological adaptations, including the preconditions of speech.
[34][35][36][37]
The immediate survival advantage of encephalization is difficult to discern, as the major brain changes from
Homo erectus to
Homo heidelbergensis
were not accompanied by major changes in technology. It has been
suggested that the changes were mainly social, including increased
empathic abilities
[38][39] and increases in size of social groups
[40][41][42]
Sexual dimorphism
The reduced degree of
sexual dimorphism in humans is visible primarily in the reduction of the male
canine tooth
relative to other ape species (except gibbons) and reduced brow ridges
and general robustness of males. Another important physiological change
related to sexuality in humans was the evolution of
hidden estrus. Humans and
bonobos
are the only apes in which the female is fertile year round and in
which no special signals of fertility are produced by the body (such as
genital swelling during estrus).
[citation needed]
Nonetheless, humans retain a degree of sexual dimorphism in the
distribution of body hair and subcutaneous fat, and in the overall size,
males being around 15% larger than females. These changes taken
together have been interpreted as a result of an increased emphasis on
pair bonding as a possible solution to the requirement for increased parental investment due to the prolonged infancy of offspring.
Ulnar opposition
The ulnar opposition – the contact between the
thumb and the tip of the
little finger of the same hand – is unique to
anatomically modern humans.
[43][44] In other primates the thumb is short and unable to touch the little finger.
[43] The ulnar opposition facilitates the precision grip and power grip of the human hand, underlying all the skilled manipulations.
Other changes
A
number of other changes have also characterized the evolution of
humans, among them an increased importance on vision rather than smell; a
smaller gut; loss of body hair; evolution of sweat glands; a change in
the shape of the dental arcade from being u-shaped to being parabolic;
development of a
chin (found in
Homo sapiens alone); development of
styloid processes; and the development of a
descended larynx.
History of study
Before Darwin
The word
homo, the name of the biological genus to which humans belong, is
Latin for "human". It was chosen originally by
Carl Linnaeus in his classification system. The word "human" is from the Latin
humanus, the adjectival form of
homo. The Latin "homo" derives from the
Indo-European root *
dhghem, or "earth".
[45] Linnaeus and other scientists of his time also considered the great apes to be the closest relatives of humans based on
morphological and
anatomical similarities.
Darwin
The possibility of linking humans with earlier apes by descent became clear only after 1859 with the publication of
Charles Darwin's
On the Origin of Species,
in which he argued for the idea of the evolution of new species from
earlier ones. Darwin's book did not address the question of human
evolution, saying only that "Light will be thrown on the origin of man
and his history."
[46]
The first debates about the nature of human evolution arose between
Thomas Henry Huxley and
Richard Owen.
Huxley argued for human evolution from apes by illustrating many of the
similarities and differences between humans and apes, and did so
particularly in his 1863 book
Evidence as to Man's Place in Nature. However, many of Darwin's early supporters (such as
Alfred Russel Wallace and
Charles Lyell) did not initially agree that the origin of the mental capacities and the moral sensibilities of humans could be explained by
natural selection, though this later changed. Darwin applied the theory of evolution and
sexual selection to humans when he published
The Descent of Man in 1871.
[47]
First fossils
A major problem in the 19th century was the lack of
fossil intermediaries. Neanderthal remains were discovered in a limestone quarry in 1856, three years before the publication of
On the Origin of Species,
and Neanderthal fossils had been discovered in Gibraltar even earlier,
but it was originally claimed that these were human remains of a
creature suffering some kind of illness.
[48] Despite the 1891 discovery by
Eugène Dubois of what is now called
Homo erectus at
Trinil,
Java, it was only in the 1920s when such fossils were discovered in Africa, that intermediate species began to accumulate.
[citation needed] In 1925,
Raymond Dart described
Australopithecus africanus.
[49] The
type specimen was the
Taung Child,
an australopithecine infant which was discovered in a cave. The child's
remains were a remarkably well-preserved tiny skull and an
endocast of the brain.
Although the brain was small (410 cm
3), its shape was rounded, unlike that of chimpanzees and
gorillas, and more like a modern human brain. Also, the specimen showed short
canine teeth, and the position of the
foramen magnum (the hole in the skull where the spine enters) was evidence of
bipedal
locomotion. All of these traits convinced Dart that the Taung Child was
a bipedal human ancestor, a transitional form between apes and humans.
The East African fossils—and Homo naledi in South Africa
During the 1960s and 1970s, hundreds of fossils were found in East Africa in the regions of the
Olduvai Gorge and
Lake Turkana. The driving force of these searches was the Leakey family, with
Louis Leakey and his wife
Mary Leakey, and later their son
Richard and daughter-in-law
Meave—all successful and world-renowned fossil hunters and
paleoanthropologists. From the fossil beds of Olduvai and Lake Turkana they amassed specimens of the early hominins: the
australopithecines and
Homo species, and even
Homo erectus.
These finds cemented Africa as the cradle of humankind. In the late 1970s and the 1980s,
Ethiopia emerged as the new hot spot of
paleoanthropology after
"Lucy", the most complete fossil member of the species
Australopithecus afarensis, was found in 1974 by
Donald Johanson near
Hadar in the desertic
Afar Triangle
region of northern Ethiopia. Although the specimen had a small brain,
the pelvis and leg bones were almost identical in function to those of
modern humans, showing with certainty that these hominins had walked
erect.
[50] Lucy was classified as a new species,
Australopithecus afarensis, which is thought to be more closely related to the genus
Homo
as a direct ancestor, or as a close relative of an unknown ancestor,
than any other known hominid or hominin from this early time range;
see terms "hominid" and "hominin".
[51] (The specimen was nicknamed "Lucy" after
the Beatles' song "
Lucy in the Sky with Diamonds", which was played loudly and repeatedly in the camp during the excavations.
[52]) The
Afar Triangle area would later yield discovery of many more hominin fossils, particularly those uncovered or described by teams headed by
Tim D. White in the 1990s, including
Ardipithecus ramidus and
Ardipithecus kadabba.
[53]
In 2013, fossil skeletons of
Homo naledi, an
extinct species of
hominin assigned (provisionally) to the
genus Homo, were found in the
Rising Star Cave system, a site in
South Africa's
Cradle of Humankind region in
Gauteng province near
Johannesburg.
[54][55] As of September 2015, fossils of at least fifteen individuals, amounting to 1550 specimens, have been excavated from the cave.
[55] The species is characterized by a body mass and stature similar to small-bodied human populations, a smaller
endocranial volume similar to
Australopithecus, and a
cranial morphology (skull shape) similar to early
Homo species. The skeletal anatomy combines primitive features known from
australopithecines
with features known from early hominins. The individuals show signs of
having been deliberately disposed of within the cave near the time of
death. The fossils have not yet been dated.
[56]
The genetic revolution
The genetic revolution in studies of human evolution started when
Vincent Sarich and
Allan Wilson measured the strength of immunological cross-reactions of
blood serum albumin between pairs of creatures, including humans and African apes (chimpanzees and gorillas).
[57]
The strength of the reaction could be expressed numerically as an
immunological distance, which was in turn proportional to the number of
amino acid
differences between homologous proteins in different species. By
constructing a calibration curve of the ID of species' pairs with known
divergence times in the fossil record, the data could be used as a
molecular clock to estimate the times of divergence of pairs with poorer or unknown fossil records.
In their seminal 1967 paper in
Science, Sarich and Wilson estimated the divergence time of humans and apes as four to five million years ago,
[57]
at a time when standard interpretations of the fossil record gave this
divergence as at least 10 to as much as 30 million years. Subsequent
fossil discoveries, notably "Lucy", and reinterpretation of older fossil
materials, notably
Ramapithecus, showed the younger estimates to be correct and validated the albumin method.
Progress in
DNA sequencing, specifically
mitochondrial DNA (mtDNA) and then
Y-chromosome DNA (Y-DNA) advanced the understanding of human origins.
[58][9][59] Application of the
molecular clock principle revolutionized the study of
molecular evolution.
On the basis of a separation from the orangutan between 10 and 20
million years ago, earlier studies of the molecular clock suggested that
there were about 76 mutations per generation that were not inherited by
human children from their parents; this evidence supported the
divergence time between hominins and chimps noted above. However, a 2012
study in Iceland of 78 children and their parents suggests a mutation
rate of only 36 mutations per generation; this datum extends the
separation between humans and chimps to an earlier period greater than 7
million years ago (
Ma).
Additional research with 226 offspring of wild chimp populations in 8
locations suggests that chimps reproduce at age 26.5 years, on average;
which suggests the human divergence from chimps occurred between 7 and
13 million years ago. And these data suggest that
Ardipithecus (4.5 Ma),
Orrorin (6 Ma) and
Sahelanthropus (7 Ma) all may be on the hominid
lineage, and even that the separation may have occurred outside the
East African Rift region.
Furthermore, analysis of the two species' genes in 2006 provides
evidence that after human ancestors had started to diverge from
chimpanzees, interspecies mating between "proto-human" and
"proto-chimps" nonetheless occurred regularly enough to change certain
genes in the new
gene pool:
- A new comparison of the human and chimp genomes suggests that after
the two lineages separated, they may have begun interbreeding... A
principal finding is that the X chromosomes of humans and chimps appear to have diverged about 1.2 million years more recently than the other chromosomes.
The research suggests:
- There were in fact two splits between the human and chimp lineages,
with the first being followed by interbreeding between the two
populations and then a second split. The suggestion of a hybridization
has startled paleoanthropologists, who nonetheless are treating the new genetic data seriously.[60]
The quest for the earliest hominin
In
the 1990s, several teams of paleoanthropologists were working
throughout Africa looking for evidence of the earliest divergence of the
hominin lineage from the great apes. In 1994, Meave Leakey discovered
Australopithecus anamensis. The find was overshadowed by Tim D. White's 1995 discovery of
Ardipithecus ramidus, which pushed back the fossil record to
4.2 million years ago.
In 2000,
Martin Pickford and
Brigitte Senut discovered, in the
Tugen Hills of
Kenya, a 6-million-year-old bipedal hominin which they named
Orrorin tugenensis. And in 2001, a team led by
Michel Brunet discovered the skull of
Sahelanthropus tchadensis which was dated as
7.2 million years ago, and which Brunet argued was a bipedal, and therefore a hominid—that is, a hominin (
cf Hominidae;
terms "hominids" and hominins).
Human dispersal
A global mapping model of human
migration, based from divergence
Timescale (
ka) indicated by colours.
called it) that emphasizes
back-and-forth gene flow among
Different models for the beginning of the present human species.
Anthropologists in the 1980s were divided regarding some details of reproductive barriers and migratory dispersals of the genus
Homo. Subsequently, genetics has been used to investigate and resolve these issues. According to the
Sahara pump theory evidence suggests that genus
Homo have migrated out of Africa at least three and possibly four times (e.g.
Homo erectus,
Homo heidelbergensis and two or three times for
Homo sapiens). Recent evidence suggests these dispersals are closely related to fluctuating periods of climate change.
[63]
Recent evidence suggests that humans may have left Africa half a
million years earlier than previously thought. A joint Franco-Indian
team has found human artifacts in the Siwalk Hills north of New Delhi
dating back at least 2.6 million years. This is earlier than the
previous earliest finding of genus
Homo at
Dmanisi, in
Georgia,
dating to 1.85 million years. Although controversial, tools found at a
Chinese cave strengthen the case that humans used tools as far back as
2.48 million years ago.
[64]
This suggests that the Asian "Chopper" tool tradition, found in Java
and northern China may have left Africa before the appearance of the
Acheulian hand axe.
Dispersal of modern Homo sapiens
Up until the genetic evidence became available there were two dominant models for the dispersal of modern humans. The
multiregional hypothesis proposed that the genus
Homo
contained only a single interconnected population as it does today (not
separate species), and that its evolution took place worldwide
continuously over the last couple of million years. This model was
proposed in 1988 by
Milford H. Wolpoff.
[65][66] In contrast the "out of Africa" model proposed that modern
H. sapiens speciated in Africa recently (that is, approximately 200,000 years ago) and the subsequent migration through
Eurasia resulted in nearly complete replacement of other
Homo species. This model has been developed by
Chris B. Stringer and Peter Andrews.
[67][68]
Sequencing mtDNA and Y-DNA sampled from a wide range of indigenous
populations revealed ancestral information relating to both male and
female genetic heritage, and strengthened the Out of Africa theory and
weakened the views of Multiregional Evolutionism.
[69] Aligned in genetic tree differences were interpreted as supportive of a recent single origin.
[70]
Analyses have shown a greater diversity of DNA patterns throughout
Africa, consistent with the idea that Africa is the ancestral home of
mitochondrial Eve and
Y-chromosomal Adam, and that modern human dispersal out of Africa has only occurred over the last 55,000 years.
[71]
"Out of Africa" has thus gained much support from research using female mitochondrial DNA and the male
Y chromosome.
After analysing genealogy trees constructed using 133 types of mtDNA,
researchers concluded that all were descended from a female African
progenitor, dubbed
Mitochondrial Eve. "Out of Africa" is also supported by the fact that mitochondrial genetic diversity is highest among African populations.
[72]
A broad study of African genetic diversity, headed by
Sarah Tishkoff, found the
San people had the greatest genetic diversity among the 113 distinct populations sampled, making them one of 14 "
ancestral population clusters". The research also located a possible origin of modern human migration in south-western Africa, near the coastal border of
Namibia and
Angola.
[73] The fossil evidence was insufficient for archaeologist
Richard Leakey to resolve the debate about exactly where in Africa modern humans first appeared.
[74] Studies of
haplogroups in
Y-chromosomal DNA and
mitochondrial DNA have largely supported a recent African origin.
[75] All the evidence from autosomal DNA also predominantly supports a Recent African origin. However, evidence for
archaic admixture in modern humans, both in Africa and later, throughout Eurasia has recently been suggested by a number of studies.
[76]
Recent sequencing of
Neanderthal[77] and
Denisovan[78] genomes shows that some admixture with these populations has occurred. Modern humans outside Africa have 2–4% Neanderthal
alleles in their genome, and some
Melanesians
have an additional 4–6% of Denisovan alleles. These new results do not
contradict the "out of Africa" model, except in its strictest
interpretation, although they make the situation more complex. After
recovery from a
genetic bottleneck that could possibly be due to the
Toba supervolcano catastrophe,
a fairly small group left Africa and later briefly interbred on three
separate occasions with Neanderthals, probably in the middle-east, on
the Eurasian steppe or even in North Africa before their departure.
Their still predominantly African descendants spread to populate the
world. A fraction in turn interbred with Denisovans, probably in
south-east Asia, before populating Melanesia.
[79] HLA haplotypes of Neanderthal and Denisova origin have been identified in modern Eurasian and
Oceanian populations.
[80] The Denisovan
EPAS1 gene has also been found in Tibetan populations.
[81]
There are still differing theories on whether there was a single
exodus from Africa or several. A multiple dispersal model involves the
Southern Dispersal theory,
[82]
which has gained support in recent years from genetic, linguistic and
archaeological evidence. In this theory, there was a coastal dispersal
of modern humans from the
Horn of Africa
crossing the Bab el Mandib to Yemen at a lower sea level around 70,000
years ago. This group helped to populate Southeast Asia and Oceania,
explaining the discovery of early human sites in these areas much
earlier than those in the
Levant.
[82] This group seems to have been dependent upon marine resources for their survival.
Stephen Oppenheimer
has proposed a second wave of humans may have later dispersed through
the Persian Gulf oases, and the Zagros mountains into the Middle East.
Alternatively it may have come across the
Sinai Peninsula
into Asia, from shortly after 50,000 yrs BP, resulting in the bulk of
the human populations of Eurasia. It has been suggested that this second
group possibly possessed a more sophisticated "big game hunting" tool
technology and was less dependent on coastal food sources than the
original group. Much of the evidence for the first group's expansion
would have been destroyed by the rising sea levels at the end of each
glacial maximum.
[82]
The multiple dispersal model is contradicted by studies indicating that
the populations of Eurasia and the populations of Southeast Asia and
Oceania are all descended from the same mitochondrial DNA L3
lineages, which support a single migration out of Africa that gave rise to all non-African populations.
[83]
Stephen Oppenheimer, on the basis of the early date of Badoshan
Iranian Aurignacian, suggests that this second dispersal, may have
occurred with a pluvial period about 50,000 years before the present,
with modern human big-game hunting cultures spreading up the Zagros
Mountains, carrying modern human genomes from Oman, throughout the
Persian Gulf, northward into Armenia and Anatolia, with a variant
travelling south into Israel and to Cyrenicia.
[84]
Evidence
The evidence on which scientific accounts of human evolution are based comes from many fields of
natural science.
The main source of knowledge about the evolutionary process has
traditionally been the fossil record, but since the development of
genetics beginning in the 1970s, DNA analysis has come to occupy a place
of comparable importance. The studies of ontogeny,
phylogeny and especially
evolutionary developmental biology
of both vertebrates and invertebrates offer considerable insight into
the evolution of all life, including how humans evolved. The specific
study of the origin and life of humans is
anthropology, particularly paleoanthropology which focuses on the study of human
prehistory.
[85]
Evidence from molecular biology
The closest living relatives of humans are bonobos and chimpanzees (both genus
Pan) and gorillas (genus
Gorilla).
[86]
With the sequencing of both the human and chimpanzee genome, as of 2012
estimates of the similarity between their DNA sequences range between
95% and 99%.
[86][87][88] By using the technique called the
molecular clock
which estimates the time required for the number of divergent mutations
to accumulate between two lineages, the approximate date for the split
between lineages can be calculated.
The gibbons (family Hylobatidae) and then orangutans (genus
Pongo)
were the first groups to split from the line leading to the hominins,
including humans—followed by gorillas, and, ultimately, by the
chimpanzees (genus
Pan). The splitting date between hominin and chimpanzee lineages is placed by some between
4 to 8 million years ago, that is, during the
Late Miocene.
[89][90][91] Speciation, however, appears to have been unusually drawn-out. Initial divergence occurred sometime between
7 to 13 million years ago,
but ongoing hybridization blurred the separation and delayed complete
separation during several millions of years. Patterson (2006) dated the
final divergence at
5 to 6 million years ago.
[92]
Genetic evidence has also been employed to resolve the question of whether there was any
gene flow between early modern humans and Neanderthals,
and to enhance our understanding of the early human migration patterns
and splitting dates. By comparing the parts of the genome that are not
under natural selection and which therefore accumulate mutations at a
fairly steady rate, it is possible to reconstruct a genetic tree
incorporating the entire human species since the last shared ancestor.
Each time a certain mutation (
single-nucleotide polymorphism)
appears in an individual and is passed on to his or her descendants a
haplogroup is formed including all of the descendants of the individual
who will also carry that mutation. By comparing mitochondrial DNA which
is inherited only from the mother, geneticists have concluded that the
last female common ancestor whose genetic marker is found in all modern
humans, the so-called mitochondrial Eve, must have lived around 200,000
years ago.
Genetics
Human evolutionary genetics studies how one
human genome differs from the other, the evolutionary past that gave rise to it, and its current effects. Differences between genomes have
anthropological,
medical and
forensic implications and applications. Genetic data can provide important insight into human evolution.
Evidence from the fossil record
Replica of fossil skull of
Homo ergaster (African
Homo erectus). Fossil number Khm-Heu 3733 discovered in 1975 in Kenya.
There is little fossil evidence for the divergence of the gorilla, chimpanzee and hominin lineages.
[93] The earliest fossils that have been proposed as members of the hominin lineage are
Sahelanthropus tchadensis dating from
7 million years ago,
Orrorin tugenensis dating from
5.7 million years ago, and
Ardipithecus kadabba dating to
5.6 million years ago. Each of these have been argued to be a
bipedal
ancestor of later hominins but, in each case, the claims have been
contested. It is also possible that one or more of these species are
ancestors of another branch of African apes, or that they represent a
shared ancestor between hominins and other apes.
The question then of the relationship between these early fossil
species and the hominin lineage is still to be resolved. From these
early species, the australopithecines arose around
4 million years ago and diverged into
robust (also called
Paranthropus) and
gracile branches, one of which (possibly
A. garhi) probably went on to become ancestors of the genus
Homo. The australopithecine species that is best represented in the fossil record is
Australopithecus afarensis
with more than one hundred fossil individuals represented, found from
Northern Ethiopia (such as the famous "Lucy"), to Kenya, and
South Africa. Fossils of robust australopithecines such as
Au. robustus (or alternatively
Paranthropus robustus) and
Au./P. boisei are particularly abundant in South Africa at sites such as
Kromdraai and
Swartkrans, and around
Lake Turkana in Kenya.
The earliest member of the genus Homo is Homo habilis which evolved around 2.8 million years ago. Homo habilis is the first species for which we have positive evidence of the use of stone tools. They developed the
Oldowan lithic technology, named after the Olduvai Gorge in which the first specimens were found. Some scientists consider
Homo rudolfensis, a larger bodied group of fossils with similar morphology to the original
H. habilis fossils, to be a separate species while others consider them to be part of
H. habilis—simply representing intraspecies variation, or perhaps even
sexual dimorphism.
The brains of these early hominins were about the same size as that of a
chimpanzee, and their main adaptation was bipedalism as an adaptation
to terrestrial living.
During the next million years, a process of encephalization began and, by the arrival (about
1.9 million years ago) of
Homo erectus in the fossil record, cranial capacity had doubled.
Homo erectus were the first of the hominins to emigrate from Africa, and, from
1.8 to 1.3 million years ago, this species spread through Africa, Asia, and Europe. One population of
H. erectus, also sometimes classified as a separate species
Homo ergaster, remained in Africa and evolved into
Homo sapiens. It is believed that these species,
H. erectus and
H. ergaster, were the first to use fire and complex tools.
The earliest transitional fossils between
H. ergaster/erectus and archaic
H. sapiens are from Africa, such as
Homo rhodesiensis, but seemingly transitional forms were also found at
Dmanisi,
Georgia. These descendants of African
H. erectus spread through Eurasia from ca. 500,000 years ago evolving into
H. antecessor,
H. heidelbergensis and
H. neanderthalensis. The earliest fossils of
anatomically modern humans are from the Middle Paleolithic, about 200,000 years ago such as the
Omo remains of Ethiopia; later fossils from
Es Skhul cave in
Israel and Southern Europe begin around 90,000 years ago (
0.09 million years ago).
As modern humans spread out from Africa, they encountered other hominins such as
Homo neanderthalensis and the so-called Denisovans, who may have evolved from populations of
Homo erectus that had left Africa around
2 million years ago.
The nature of interaction between early humans and these sister species
has been a long-standing source of controversy, the question being
whether humans replaced these earlier species or whether they were in
fact similar enough to interbreed, in which case these earlier
populations may have contributed genetic material to modern humans.
[95][96]
This migration out of Africa is estimated to have begun about 85,000 years
[97] BP dated by a finger bone unearthed in
Saudi Arabia’s
Nefud
Desert and modern humans subsequently spread globally, replacing
earlier hominins either through competition or hybridization. They
inhabited Eurasia and Oceania by 40,000 years BP, and the Americas by at
least 14,500 years BP.
[98]
Before Homo
Early evolution of primates
Evolutionary history of the primates can be traced back 65 million years.
[99] One of the oldest known primate-like mammal species, the
Plesiadapis, came from North America;
[100] another,
Archicebus, came from China.
[101] Other similar basal primates were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and
Eocene.
David R. Begun
[102] 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.
[103] Its ancestry is thought to be species related to
Aegyptopithecus,
Propliopithecus, and
Parapithecus from the Faiyum, at around 35 million years ago.
[104] In 2010,
Saadanius was described as a close relative of the last common ancestor of the
crown catarrhines, and tentatively dated to 29–28 million years ago, helping to fill an 11-million-year gap in the fossil record.
[105]
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 (family
Hylobatidae) diverged from the line of great apes some 18–12 million
years ago, and that of orangutans (subfamily Ponginae) 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 South East Asian hominoid population, but fossil
proto-orangutans may be represented by
Sivapithecus from
India and
Griphopithecus from
Turkey, dated to around 10 million years ago.
[16]
Divergence of the human clade from other great apes
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.
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.
[106] 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 chimpanzee, may have evolved via the process of
self-domestication. Consequently, arguing against the so-called "chimpanzee referential model"
[107] the authors suggest it is no longer tenable to use
common 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.[106]: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
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, the
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.[108] 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.
[109]
Given the opposable big toe found on Little Foot, it seems that he was a
good climber, and it is thought given the night predators of the
region, he probably, like gorillas and chimpanzees, built a nesting
platform at night, in the trees.
Evolution of genus Homo
The earliest documented representative of the genus
Homo is
Homo habilis, which evolved around
2.8 million years ago,
[94]
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 cm
3.
[110] (Such an increase in human brain size is equivalent to each generation having 125,000 more
neurons than their parents.) It is believed that
Homo erectus and
Homo 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.
One current view of the temporal and geographical distribution of genus
Homo populations.
[111] Other interpretations differ mainly in the taxonomy and geographical distribution of hominin species.
According to the recent African origin of modern humans theory, modern humans evolved in Africa possibly from
Homo heidelbergensis,
Homo rhodesiensis or
Homo antecessor and migrated out of the continent some 50,000 to 100,000 years ago, gradually replacing local populations of
Homo erectus,
Denisova hominins,
Homo floresiensis and
Homo neanderthalensis.
[112][113][114][115][116] Archaic Homo sapiens, the forerunner of
anatomically modern humans, evolved in the
Middle Paleolithic between 400,000 and 250,000 years ago.
[117][118][119] 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 inter-breeding between these species.
[78][120][80] 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
[121] although others point to evidence that suggests that a gradual change in behavior took place over a longer time span.
[122]
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.
[123][124]
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.
[124] 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
[30] of various
Homo species and to study the role of diet in physical and behavioral evolution within
Homo.
[27][125][126][127][128]
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,
[129] killing the majority of humans and creating a
population bottleneck that affected the genetic inheritance of all humans today.
[130]
H. habilis and H. gautengensis
Homo habilis lived from about 2.8
[94] 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.
Homo habilis
had smaller molars and larger brains than the australopithecines, and
made tools from stone and perhaps animal bones. One of the first known
hominins, it was nicknamed 'handy man' by discoverer
Louis Leakey due to its association with
stone tools. Some scientists have proposed moving this species out of
Homo and into
Australopithecus due to the morphology of its skeleton being more adapted to
living on trees rather than to
moving on two legs like
Homo sapiens.
[131]
In May 2010, a new species,
Homo gautengensis, was discovered in South Africa.
[132]
H. rudolfensis and H. georgicus
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 another Homo habilis, but this has not been confirmed.[133]
- Homo georgicus, from Georgia, may be an intermediate form between Homo habilis and Homo erectus,[134] or a sub-species of Homo erectus.[135]
H. ergaster and H. erectus
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 a 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).
[136] 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
Homo erectus.
[citation needed] 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
Homo floresiensis survived it. The early phase of
Homo erectus, from 1.8 to 1.25 Ma, is considered by some to be a separate species,
Homo ergaster, or as
Homo erectus ergaster, a subspecies of
Homo erectus.
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.
[137]
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
suggests that the fact that Homo seems to have been ground dwelling,
with reduced intestinal length, smaller dentition, "and swelled our
brains to their current, horrendously fuel-inefficient size",
[138]
suggest that control of fire and releasing increased nutritional value
through cooking was the key adaptation that separated Homo from
tree-sleeping Australopitheicines.
[139]
famous example of
Homo erectus is
Peking Man; others were found in Asia (notably in Indonesia), Africa, and Europe. Many paleoanthropologists now use the term
Homo ergaster for the non-Asian forms of this group, and reserve
Homo erectus only for those fossils that are found in Asia and meet certain skeletal and dental requirements which differ slightly from
H. ergaster.
H. cepranensis and H. antecessor
These are proposed as species that may be 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.[140][141]
- H. cepranensis refers to a single skull cap from Italy, estimated to be about 800,000 years old.[142]
H. 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.
[143]
H. rhodesiensis, and the Gawis cranium
- 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.[144]
Neanderthal and Denisovan
Homo neanderthalensis, alternatively designated as
Homo sapiens neanderthalensis,
[145] lived in Europe and Asia from 400,000
[146] to about 28,000 years ago.
[147] There are a number of clear anatomical differences between
anatomically modern humans
(AMH) and Neanderthal populations. Many of these relate to the superior
adaptation to cold environments possessed by the Neanderthal
populations. Their
surface to volume ratio is an extreme version of that found amongst
Inuit
populations, indicating that they were less inclined to lose body heat
than were AMH. From brain Endocasts, Neanderthals also had significantly
larger brains. This would seem to indicate that the intellectual
superiority of AMH populations may be questionable. More recent research
by Eiluned Pearce, Chris Stringer, R. I. M. Dunbar, however, have shown
important differences in Brain architecture. For example, in both the
orbital chamber size and in the size of the
occipital lobe,
the larger size suggests that the Neanderthal had a better visual
acuity than modern humans. This would give a superior vision in the
inferior light conditions found in Glacial Europe. It also seems that
the higher body mass of Neanderthals had a correspondingly larger brain
mass required for body care and control. The Neanderthal populations
seem to have been physically superior to AMH populations. These
differences may have been sufficient to give Neanderthal populations an
environmental superiority to AMH populations from 75,000 to 45,000 years
BP. With these differences, Neanderthal brains show a smaller area was
available for social functioning. Plotting group size possible from
endocrainial volume, suggests that AMH populations (minus occipital lobe
size), had a
Dunbars number
of 144 possible relationships. Neanderthal populations seem to have
been limited to about 120 individuals. This would show up in a larger
number of possible mates for AMH humans, with increased risks of
inbreeding amongst Neanderthal populations. It also suggests that humans
had larger trade catchment areas than Neanderthals (confirmed in the
distribution of stone tools). With larger populations, social and
technological innovations were easier to fix in human populations, which
may have all contributed to the fact that modern Homo sapiens replaced
the Neanderthal populations by 28,000 BP.
[148]
Dermoplastic reconstruction of a Neanderthal
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.
[149][150][151]
However, a sequencing of the Neanderthal genome in 2010 indicated that
Neanderthals did indeed interbreed with anatomically modern humans
circa
45,000 to 80,000 years ago (at the approximate time that modern humans
migrated out from Africa, but before they dispersed into Europe, Asia
and elsewhere).
[152]
The genetic sequencing of a human from Romania dated 40,000 years ago
showed that 11% of their genome was Neanderthal. This would indicate
that this individual had a Neanderthal great grandparent, 4 generations
previously. It seems that this individual has left no living
descendants.
Nearly all modern non-African humans have 1% to 4% of their DNA derived from Neanderthal DNA,
[152]
and this finding is consistent with recent studies indicating that the
divergence of some human alleles dates to one Ma, although the
interpretation of these studies has been questioned.
[153][154] Neanderthals and
Homo sapiens
could have co-existed in Europe for as long as 10,000 years, during
which human populations exploded vastly outnumbering Neanderthals,
possibly outcompeting them by sheer numerical strength.
[155]
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 Denisovans.
[156] 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.
[78][157]
While the divergence point of the mtDNA was unexpectedly deep in time,
[158]
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.
[78] Modern humans are known to have overlapped with Neanderthals in Europe and the Near East for possibly more than 40,000 years,
[159]
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.
[157][160] 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.
[79][161]
Alleles thought to have originated in Neanderthals and Denisovans
have been identified at several genetic loci in the genomes of modern
humans outside of Africa. HLA haplotypes from Denisovans and Neanderthal
represent more than half the HLA alleles of modern Eurasians,
[80] 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 likelihood to suffer depression more frequently.
[162]
The flow of genes from Neanderthal populations to modern human was
not all one way. Sergi Castellano of the Max Planck Institute for
Evolutionary Anthropology
in Leipzig, Germany, has in 2016 reported that while Denisovan and
Neanderthal genomes are more related to each other than they are to us,
Siberian Neanderthal genomes show similarity to the modern human gene
pool, more so than to European Neanderthal populations. The evidence
suggests that the Neanderthal populations interbred with modern humans
possibly 100,000 years ago, probably somewhere in the Near East.
[163]
Studies of a Neanderthal child at Gibraltar show from brain
development and teeth eruption that Neanderthal children may have
matured more rapidly than is the case for Homo sapiens.
[164]
H. floresiensis
Models representing Homo sapiens, Homo neanderthalensis and Homo floresiensis
H. floresiensis, which lived from approximately 190,000 to 50,000 years before present, has been nicknamed
hobbit for its small size, possibly a result of
insular dwarfism.
[165] 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 cm
3 (considered small for a chimpanzee and less than a third of the
H. sapiens average of 1400 cm
3).
However, there is an ongoing debate over whether
H. floresiensis is indeed a separate species.
[166] Some scientists hold that
H. floresiensis was a modern
H. sapiens with pathological dwarfism.
[167] 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.
[167]
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.
[citation needed]
H. sapiens
H. sapiens (the adjective
sapiens is Latin for "wise" or "intelligent") emerged around 300,000 years ago, likely derived from
Homo heidelbergensis.
[168] 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.
[169]
The
Toba catastrophe theory, which postulates a
population bottleneck for
H. sapiens about 70,000 years ago,
[170] was controversial from its first proposal in the 1990s and by the 2010s had very little support.
[171]
Distinctive
human genetic variability has arisen as the result of the
founder effect, by
archaic admixture and by
recent evolutionary pressures.
Use of tools
"A sharp rock", an
Oldowan pebble tool, the most basic of human stone tools.
The harnessing of fire was a pivotal milestone in human history.
Acheulean hand-axes from
Kent.
Homo erectus flint work. The types shown are (clockwise from top) cordate, ficron and ovate.
The use of tools has been interpreted as a sign of intelligence, and
it has been theorized that tool use may have stimulated certain aspects
of human evolution, especially the continued expansion of the human
brain.
[172]
Paleontology has yet to explain the expansion of this organ over
millions of years despite being extremely demanding in terms of energy
consumption. The brain of a modern human consumes about 13
watts (260 kilocalories per day), a fifth of the body's resting power consumption.
[173]
Increased tool use would allow hunting for energy-rich meat products,
and would enable processing more energy-rich plant products. Researchers
have suggested that early hominins were thus under evolutionary
pressure to increase their capacity to create and use tools.
[174]
Precisely when early humans started to use tools is difficult to
determine, because the more primitive these tools are (for example,
sharp-edged stones) the more difficult it is to decide whether they are
natural objects or human artifacts.
[172] There is some evidence that the australopithecines (4 Ma)
may have used broken bones as tools, but this is debated.
[175]
It should be noted that
many species make and use tools,
but it is the human genus that dominates the areas of making and using
more complex tools. The oldest known tools are the Oldowan stone tools
from Ethiopia, 2.5–2.6 million years old. A
Homo fossil was found near some Oldowan tools, and its age was noted at 2.3 million years old, suggesting that maybe the
Homo species did indeed create and use these tools. It is a possibility but does not yet represent solid evidence.
[176] The
third metacarpal styloid process
enables the hand bone to lock into the wrist bones, allowing for
greater amounts of pressure to be applied to the wrist and hand from a
grasping thumb and fingers. It allows humans the dexterity and strength
to make and use complex tools. This unique anatomical feature separates
humans from apes and other nonhuman primates, and is not seen in human
fossils older than 1.8 million years.
[177]
Bernard Wood noted that
Paranthropus co-existed with the early
Homo
species in the area of the "Oldowan Industrial Complex" over roughly
the same span of time. Although there is no direct evidence which
identifies
Paranthropus as the tool makers, their anatomy lends
to indirect evidence of their capabilities in this area. Most
paleoanthropologists agree that the early
Homo species were
indeed responsible for most of the Oldowan tools found. They argue that
when most of the Oldowan tools were found in association with human
fossils,
Homo was always present, but
Paranthropus was not.
[176]
In 1994, Randall Susman used the anatomy of opposable thumbs as the basis for his argument that both the
Homo and
Paranthropus
species were toolmakers. He compared bones and muscles of human and
chimpanzee thumbs, finding that humans have 3 muscles which are lacking
in chimpanzees. Humans also have thicker metacarpals with broader heads,
allowing more precise grasping than the chimpanzee hand can perform.
Susman posited that modern anatomy of the human opposable thumb is an
evolutionary response to the requirements associated with making and
handling tools and that both species were indeed toolmakers.
[176]
Stone tools
Stone tools are first attested around 2.6 Million years ago, when
H. habilis in Eastern Africa used so-called
pebble tools,
choppers made out of round pebbles that had been split by simple strikes.
[178] This marks the beginning of the
Paleolithic, or Old
Stone Age; its end is taken to be the end of the last
Ice Age, around 10,000 years ago. The Paleolithic is subdivided into the
Lower Paleolithic (Early Stone Age), ending around 350,000–300,000 years ago, the
Middle Paleolithic (Middle Stone Age), until 50,000–30,000 years ago, and the
Upper Paleolithic, (Late Stone Age), 50,000-10,000 years ago.
Archaeologists working in the Great Rift Valley in Kenya claim to
have discovered the oldest known stone tools in the world. Dated to
around 3.3 million years ago, the implements are some 700,000 years
older than stone tools from Ethiopia that previously held this
distinction.
[179][180][181]
The period from 700,000–300,000 years ago is also known as the
Acheulean, when
H. ergaster (or
erectus) made large stone
hand axes out of
flint and
quartzite, at first quite rough (Early Acheulian), later "
retouched" by additional, more-subtle strikes at the sides of the
flakes. After 350,000 BP the more refined so-called
Levallois technique
was developed, a series of consecutive strikes, by which scrapers,
slicers ("racloirs"), needles, and flattened needles were made.
[178] Finally, after about 50,000 BP, ever more refined and specialized flint tools were made by the Neanderthals and the immigrant
Cro-Magnons (knives, blades, skimmers). In this period they also started to make tools out of bone.
Transition to behavioral modernity
Until about 50,000–40,000 years ago, the use of stone tools seems to have progressed stepwise. Each phase (
H. habilis,
H. ergaster,
H. neanderthalensis)
started at a higher level than the previous one, but after each phase
started, further development was slow. Currently paleoanthropologists
are debating whether these
Homo species possessed some or many of
the cultural and behavioral traits associated with modern humans such
as language, complex symbolic thinking, technological creativity etc. It
seems that they were culturally conservative maintaining simple
technologies and foraging patterns over very long periods.
Around 50,000 BP, modern human culture started to evolve more
rapidly. The transition to behavioral modernity has been characterized
by most as a Eurasian "Great Leap Forward",
[182] or as the "Upper Palaeolithic Revolution",
[183] due to the sudden appearance of distinctive signs of modern behavior and
big game hunting[84]
in the archaeological record. Some other scholars consider the
transition to have been more gradual, noting that some features had
already appeared among archaic African
Homo sapiens since 200,000 years ago.
[184][185]
Recent evidence suggests that the Australian Aboriginal population
separated from the African population 75,000 years ago, and that they
made a sea journey of up to 160 km 60,000 years ago, which may diminish
the evidence of the Upper Paleolithic Revolution.
[186]
Modern humans started burying their dead, using animal hides to make
clothing, hunting with more sophisticated techniques (such as using
trapping pits or driving animals off cliffs), and engaging in
cave painting.
[187]
As human culture advanced, different populations of humans introduced
novelty to existing technologies: artifacts such as fish hooks, buttons,
and bone needles show signs of variation among different populations of
humans, something that had not been seen in human cultures prior to
50,000 BP. Typically,
H. neanderthalensis populations do not vary in their technologies, although the
Chatelperronian assemblages have been found to be Neanderthal innovations produced as a result of exposure to the Homo sapiens
Aurignacian technologies.
[188]
Among concrete examples of
modern human behavior,
anthropologists include specialization of tools, use of jewellery and
images (such as cave drawings), organization of living space, rituals
(for example, burials with grave gifts), specialized hunting techniques,
exploration of less hospitable geographical areas, and
barter
trade networks. Debate continues as to whether a "revolution" led to
modern humans ("the big bang of human consciousness"), or whether the
evolution was more "gradual".
[122]
Recent and current human evolution
Evolution has continued in
anatomically modern human populations, which are affected by both natural selection and
genetic drift. Although
selection pressure
on some traits, such as resistance to smallpox, has decreased in modern
human life, humans are still undergoing natural selection for many
other traits. Some of these are due to specific environmental pressures,
while others are related to lifestyle changes since the development of
agriculture (10,000 years ago), urban civilization (5,000), and
industrialization
(250 years ago). It has been argued that human evolution has
accelerated since the development of agriculture 10,000 years ago and
civilization some 5,000 years ago, resulting, it is claimed, in
substantial genetic differences between different current human
populations.
[189]
Particularly conspicuous is variation in superficial characteristics, such as
Afro-textured hair, or the recent evolution of
light skin and
blond
hair in some populations, which are attributed to differences in
climate. Particularly strong selective pressures have resulted in
high-altitude adaptation in humans, with different ones in different isolated populations. Studies of the
genetic basis show that some developed very recently, with Tibetans evolving over 3,000 years to have high proportions of an allele of
EPAS1 that is adaptive to high altitudes.
Other evolution is related to
endemic diseases: the presence of
malaria selected for
sickle cell trait (the
heterozygote form of sickle cell gene), while the absence of malaria and the health effects of
sickle-cell anemia select against this trait. For example, the population at risk of the severe debilitating disease
kuru has significant over-representation of an immune variant of the
prion protein gene G127V versus non-immune alleles. The frequency of this
genetic variant is due to the survival of immune persons.
[190][191]
Recent human evolution related to agriculture includes genetic
resistance to infectious disease that has appeared in human populations
by crossing the species barrier from domesticated animals,
[192] as well as changes in metabolism due to changes in diet, such as
lactase persistence.
In contemporary times, since industrialization, some trends have been
observed: for instance, menopause is evolving to occur later.
[193]
Other reported trends appear to include lengthening of the human
reproductive period and reduction in cholesterol levels, blood glucose
and blood pressure in some populations.
[193]
Species list
This list is in
chronological order across the table by
genus. Some species/subspecies names are well-established, and some are less established – especially in genus
Homo. Please see articles for more information.
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