Timeline of human evolution
The timeline of human evolution outlines the major events in the development of the human species, Homo sapiens, and the evolution of human ancestors. It includes brief explanations of some of the species, genera, and the higher ranks of taxa that are seen today as possible ancestors of modern humans.
This timeline is based on studies from anthropology, paleontology, developmental biology, morphology, and from anatomical and genetic data. It does not address the origin of life, which discussion is provided by abiogenesis, but presents one possible line of evolutionary descent of species that eventually led to humans.
A caution: Other than Mr Haeckel's historic and emblematic "tree", this article provides no phylogenetics analysis to help portray the complex, nonlinear facts of human evolution. It is important that the "possible line of evolutionary descent" should not be interpreted as providing a linear progression from very early taxa to a (presupposed) objective of Homo sapiens; no such directed progression is implied here.
Taxonomy of Homo sapiens
|Taxonomic rank||Name||Common name||Millions of years ago|
|Domain||Eukaryota||Cells with a nucleus||2,100|
|Phylum||Chordata||Vertebrates and closely related invertebrates||530|
|Superclass||Tetrapoda||Tetrapods (animals with four limbs)||395|
|Unranked||Amniota||Amniotes (fully terrestrial tetrapods whose eggs are "equipped with an amnios")||340|
|Subclass||Theria||Mammals that give birth to live young (i.e., non-egg-laying)|
|Infraclass||Eutheria||Placental mammals (i.e., non-marsupials)||125|
|Magnorder||Boreoeutheria||Supraprimates, (most) hoofed mammals, (most) carnivorous mammals, whales, and bats|
|Superorder||Euarchontoglires||Supraprimates: primates, colugos, tree shrews, rodents, and rabbits||100|
|Grandorder||Euarchonta||Primates, colugos, and tree shrews|
|Mirorder||Primatomorpha||Primates and colugos||79.6|
|Suborder||Haplorrhini||"Dry-nosed" (literally, "simple-nosed") primates: apes, monkeys, and tarsiers||40|
|Infraorder||Simiiformes||"Higher" primates (Simians): apes and monkeys|
|Parvorder||Catarrhini||"Downward-nosed" primates: apes and old-world monkeys||30|
|Superfamily||Hominoidea||Apes: great apes and lesser apes (gibbons)||28|
|Family||Hominidae||Great apes: humans, chimpanzees, gorillas, and orangutans—the hominids||15|
|Subfamily||Homininae||Humans, chimpanzees, and gorillas (the African apes)||8|
|Tribe||Hominini||Genera Homo, Pan, and the Australopithecines||5.8|
|Subtribe||Hominina||Genus Homo and close human relatives and ancestors after splitting from Pan—the hominins||2.5|
|Species||(Archaic) Homo sapiens||Neanderthals||0.5|
|Subspecies||Homo sapiens sapiens||Anatomically modern humans||0.2|
First living beings
|The earliest life appears.
Further information: Abiogenesis
|3900 Ma||Cells resembling prokaryotes appear. This marks the first appearance of photosynthesis and therefore the first occurrence of large quantities of atmospheric oxygen on the earth.
Further information: Cell (biology) § Origins
|2500 Ma||First organisms to utilize oxygen. By 2400 Ma, in what is referred to as the Great Oxygenation Event, the pre-oxygen anaerobic forms of life were wiped out by the oxygen consumers.|
|2100 Ma||More complex cells appear: the eukaryotes.
Further information: Eukaryote § Origin of eukaryotes
|1200 Ma||Sexual reproduction evolves, leading to faster evolution where genes are mixed in every generation enabling greater variation for subsequent selection.|
Proterospongia (members of the Choanoflagellata) are the best living examples of what the ancestor of all animals may have looked like. They live in colonies, and show a primitive level of cellular specialization for different tasks.
|600 Ma||It is thought that the earliest multicellular animal was a sponge-like creature.
Sponges are among the simplest of animals, with partially differentiated tissues.
Sponges (Porifera) are the phylogenetically oldest animal phylum extant today.
|580 Ma||Multicellular animal movement may have started with cnidarians. Almost all cnidarians possess nerves and muscles. Because they are the simplest animals to possess them, their direct ancestors were very probably the first animals to use nerves and muscles together. Cnidarians are also the first animals with an actual body of definite form and shape. They have radial symmetry. The first eyes evolved at this time.|
Flatworms are the earliest animals to have a brain, and the simplest animals alive to have bilateral symmetry. They are also the simplest animals with organs that form from three germ layers.
Most known animal phyla appeared in the fossil record as marine species during the Cambrian explosion.
|540 Ma||Acorn worms are considered more highly specialised and advanced than other similarly shaped worm-like creatures. They have a circulatory system with a heart that also functions as a kidney. Acorn worms have a gill-like structure used for breathing, a structure similar to that of primitive fish. Acorn worms are thus sometimes said to be a link between vertebrates and invertebrates.|
|530 Ma||Pikaia is an iconic ancestor of modern chordates and vertebrates. Other, earlier chordate predecessors include Myllokunmingia fengjiaoa, Haikouella lanceolata, and Haikouichthys ercaicunensis. Conodonts are a famous type of early (495 Mya and later) chordate fossil; they have the peculiar teeth of an eel-shaped animal characterized by large eyes, fins with fin rays, chevron-shaped muscles and a notochord. The animal is sometimes called a conodont, and sometimes a conodontophore (conodont-bearer) to avoid confusion.|
The first vertebrates appear: the ostracoderms, jawless fish related to present-day lampreys and hagfishes. Haikouichthys and Myllokunmingia are examples of these jawless fish, or Agnatha. (See also prehistoric fish). They were jawless and their internal skeletons were cartilaginous. They lacked the paired (pectoral and pelvic) fins of more advanced fish. They were precursors to the Osteichthyes (bony fish).
The Placodermi were prehistoric fishes. Placoderms were some of the first jawed fishes (Gnathostomata), their jaws evolving from the first gill arch. A placoderm's head and thorax were covered by articulated armoured plates and the rest of the body was scaled or naked. However, the fossil record indicates that they left no descendents after the end of the Devonian and are less closely related to living bony fishes than sharks are.
|410 Ma||The first coelacanth appears; this order of animals had been thought to have no extant members until living specimens were discovered in 1938. It is often referred to as a living fossil.|
Primitive tetrapods developed from a lobe-finned fish (an "osteolepid Sarcopterygian"), with a two-lobed brain in a flattened skull, a wide mouth and a short snout, whose upward-facing eyes show that it was a bottom-dweller, and which had already developed adaptations of fins with fleshy bases and bones. (The "living fossil" coelacanth is a related lobe-finned fish without these shallow-water adaptations.) Tetrapod fishes used their fins as paddles in shallow-water habitats choked with plants and detritus. The universal tetrapod characteristics of front limbs that bend backward at the elbow and hind limbs that bend forward at the knee can plausibly be traced to early tetrapods living in shallow water. 
Panderichthys is a 90–130 cm (35–50 in) long fish from the Late Devonian period (380 Mya). It has a large tetrapod-like head. Panderichthys exhibits features transitional between lobe-finned fishes and early tetrapods.
Trackway impressions made by something that resembles Ichthyostega's limbs were formed 390 Ma in Polish marine tidal sediments. This suggests tetrapod evolution is older than the dated fossils of Panderichthys through to Ichthyostega.
Acanthostega is an extinct amphibian, among the first animals to have recognizable limbs. It is a candidate for being one of the first vertebrates to be capable of coming onto land. It lacked wrists, and was generally poorly adapted for life on land. The limbs could not support the animal's weight. Acanthostega had both lungs and gills, also indicating it was a link between lobe-finned fish and terrestrial vertebrates.
Ichthyostega is an early tetrapod. Being one of the first animals with legs, arms, and finger bones, Ichthyostega is seen as a hybrid between a fish and an amphibian. Ichthyostega had legs but its limbs probably were not used for walking. They may have spent very brief periods out of water and would have used their legs to paw their way through the mud.
From amphibians came the first reptiles: Hylonomus is the earliest known reptile. It was 20 cm (8 in) long (including the tail) and probably would have looked rather similar to modern lizards. It had small sharp teeth and probably ate millipedes and early insects. It is a precursor of later Amniotes and mammal-like reptiles. Αlpha keratin first evolves here which is used in claws in modern lizards and birds, and hair in mammals.
Evolution of the amniotic egg gives rise to the Amniota, reptiles that can reproduce on land and lay eggs on dry land. They did not need to return to water for reproduction. This adaptation gave them the capability to colonize the uplands for the first time.
Reptiles have advanced nervous systems, compared to amphibians. They have twelve pairs of cranial nerves.
Sauropsids, from which come the modern reptiles and birds. The other branch is Synapsida, from which come modern mammals. Both had temporal fenestrae, a pair of holes in their skulls behind the eyes, which were used to increase the space for jaw muscles. Synapsids had one opening on each side, while diapsids had two.
The earliest mammal-like reptiles are the pelycosaurs. The pelycosaurs were the first animals to have temporal fenestrae. Pelycosaurs are not therapsids but soon they gave rise to them. The Therapsida were the direct ancestor of mammals.
The therapsids have temporal fenestrae larger and more mammal-like than pelycosaurs, their teeth show more serial differentiation, and later forms had evolved a secondary palate. A secondary palate enables the animal to eat and breathe at the same time and is a sign of a more active, perhaps warm-blooded, way of life.
One sub-group of therapsids, the cynodonts, evolved more mammal-like characteristics.
The jaws of cynodonts resemble modern mammal jaws. It is very likely that this group of animals contains a species which is the direct ancestor of all modern mammals.
From Eucynodontia (cynodonts) came the first mammals. Most early mammals were small shrew-like animals that fed on insects. Although there is no evidence in the fossil record, it is likely that these animals had a constant body temperature and milk glands for their young. The neocortex region of the brain first evolved in mammals and thus is unique to them.
Monotremes are an egg-laying group of mammals represented amongst modern animals by the platypus and echidna. Recent genome sequencing of the platypus indicates that its sex genes are closer to those of birds than to those of the therian (live birthing) mammals. Comparing this to other mammals, it can be inferred that the first mammals to gain gender differentiation through the existence or lack of SRY gene (found in the y-Chromosome) evolved after the monotreme lineage split off.
|100 Ma||Last common ancestor of mice and humans (base of the clade Euarchontoglires).|
A group of small, nocturnal and arboreal, insect-eating mammals called the Euarchonta begins a speciation that will lead to the primate, treeshrew and flying lemur orders. The Primatomorpha is a subdivision of Euarchonta that includes the primates and the stem-primates Plesiadapiformes. One of the early stem-primates is Plesiadapis. Plesiadapis still had claws and the eyes located on each side of the head. Because of this they were faster on the ground than on the top of the trees, but they began to spend long times on lower branches of trees, feeding on fruits and leaves. The Plesiadapiformes very likely contain the species which is the ancestor of all primates.
One of the last Plesiadapiformes is Carpolestes simpsoni. It had grasping digits but no forward-facing eyes.
|63 Ma||Primates diverge into suborders Strepsirrhini (wet-nosed primates) and Haplorrhini (dry-nosed primates). Strepsirrhini contain most of the prosimians; modern examples include the lemurs and lorises. The haplorrhines include the three living groups: prosimian tarsiers, simian monkeys, and apes. One of the earliest haplorrhines is Teilhardina asiatica, a mouse-sized, diurnal creature with small eyes. The Haplorrhini metabolism lost the ability to make its own Vitamin C. This means that it and all its descendants had to include fruit in its diet, where Vitamin C could be obtained externally.|
Haplorrhini splits into infraorders Platyrrhini and Catarrhini. Platyrrhines, New World monkeys, have prehensile tails and males are color blind. They may have migrated to South America on a raft of vegetation across the relatively narrow Atlantic ocean (approx. 700 km). Catarrhines mostly stayed in Africa as the two continents drifted apart. Possible early ancestors of catarrhines include Aegyptopithecus and Saadanius.
Proconsul was an early genus of catarrhine primates. They had a mixture of Old World monkey and ape characteristics. Proconsul's monkey-like features include thin tooth enamel, a light build with a narrow chest and short forelimbs, and an arboreal quadrupedal lifestyle. Its ape-like features are its lack of a tail, ape-like elbows, and a slightly larger brain relative to body size.
Proconsul africanus is a possible ancestor of both great and lesser apes, including humans.
|15 Ma||Hominidae (great apes) speciate from the ancestors of the gibbon (lesser apes).|
|13 Ma||Homininae ancestors speciate from the ancestors of the orangutan.
Pierolapithecus catalaunicus is believed to be a common ancestor of humans and the other great apes, or at least a species that brings us closer to a common ancestor than any previous fossil discovery. It had the special adaptations for tree climbing as do present-day humans and other great apes: a wide, flat rib cage, a stiff lower spine, flexible wrists, and shoulder blades that lie along its back.
|10 Ma||The lineage currently represented by humans and the Pan genus (common chimpanzees and bonobos) speciates from the ancestors of the gorillas.|
The Hominina, a subtribe of Hominini who are closely related to or ancestors to humans, speciate from the ancestors of the chimpanzees. Both chimpanzees and humans have a larynx that repositions during the first two years of life to a spot between the pharynx and the lungs, indicating that the common ancestors have this feature, a precondition for vocalized speech in humans.
The latest common ancestor of humans and chimpanzees lived around the time of Sahelanthropus tchadensis, c. 7 Ma ; and S. tchadensis is sometimes claimed to be that last common ancestor of humans and chimpanzees, but the claim has not been established. The earliest known representative from the ancestral human line post-dating the separation with the chimpanzee lines is Orrorin tugenensis (Millennium Man, Kenya; c. 6 Ma).
Ardipithecus is, or may be, a very early hominin genus (tribe Hominini and subtribe Hominina). Two species are described in the literature: A. ramidus, which lived about 4.4 million years ago during the early Pliocene, and A. kadabba, dated to approximately 5.6 million years ago (late Miocene). A. ramidus had a small brain, measuring between 300 and 350 cm3. This is about the same size as the modern bonobo and female common chimpanzee brain; it is much smaller than the brain of australopithecines like Lucy (400 to 550 cm3) and slightly over a fifth the size of the modern Homo sapiens brain.
Ardipithecus was arboreal, meaning it lived largely in the forest where it competed with other forest animals for food, no doubt including the contemporary ancestor of the chimpanzees. Ardipithecus was probably bipedal as evidenced by its bowl shaped pelvis, the angle of its foramen magnum and its thinner wrist bones, though its feet were still adapted for grasping rather than walking for long distances.
A member of the Australopithecus afarensis left human-like footprints on volcanic ash in Laetoli, Kenya (Northern Tanzania), providing strong evidence of full-time bipedalism. Australopithecus afarensis lived between 3.9 and 2.9 million years ago, and is considered one of the earliest hominins—those species that developed and comprised the lineage of Homo and Homo's closest relatives after the split from the line of the chimpanzees.
It is thought that A. afarensis was ancestral to both the genus Australopithecus and the genus Homo. Compared to the modern and extinct great apes, A. afarensis had reduced canines and molars, although they were still relatively larger than in modern humans. A. afarensis also has a relatively small brain size (380–430 cm³) and a prognathic (i.e. projecting anteriorly) face.
Australopithecines have been found in savannah environments; from scavenging opportunities, they probably developed their diet to include meat. Analyses of Australopithecus africanus lower vertebrae suggests that these bones changed in females to support bipedalism even during pregnancy.
|3.5–3.3 Ma||Kenyanthropus platyops, a possible ancestor of Homo, emerges from the Australopithecus genus. Stone tools are deliberately constructed.|
|3 Ma||The bipedal australopithecines (a genus of the Hominina subtribe) evolve in the savannas of Africa being hunted by Dinofelis. Loss of body hair occurs from 3 to 2 Ma, in parallel with the development of full bipedalism.|
Homo appears in East Africa; with most Australopithecines they are considered the first hominins—that is, they are designated (by some) as those earliest humans and human relatives or ancestors to rise after splitting from the lineage of Pan, the chimpanzees. Others consider the genus Pan as hominins also, and perhaps the first hominins.
Homo habilis, although significantly different of anatomy and physiology, is thought to be the ancestor of Homo ergaster, or African Homo erectus; but it is also known to have coexisted with Homo erectus for some one-half million years (until about 1.5 Ma).
Homo erectus evolves in Africa. Homo erectus would bear a striking resemblance to modern humans, but had a brain about 74 percent of the size of modern man. Its forehead is less sloping than that of Homo habilis and the teeth are smaller.
Homo ergaster, known as African Homo erectus, and other hominin species such as Homo georgicus, Homo pekinensis, Homo heidelbergensis are often put under the umbrella species name of Homo erectus. Starting with Homo georgicus—found in what is now the Republic of Georgia, dated at 1.8 Ma—the pelvis and backbone grew more human-like, which would enable H. georgicus to cover very long distances and to follow herds of prey animals; this is the oldest fossil of a hominin found outside of Africa.
Control of fire by early humans is achieved about 1.5 Ma by Homo ergaster. Homo ergaster reaches a height of around 1.9 metres (6.2 ft). Evolution of dark skin, which is linked to the loss of body hair in human ancestors, is complete by 1.2 Ma. Homo pekinensis first appears in Asia around 700 ka but, according to the theory of a recent African origin of modern humans, they could not be ancestors to modern humans, but rather, were an offshoot cousin species from Homo erectus. Homo heidelbergensis was a very large hominin that developed a more advanced complement of cutting tools and may have hunted big game such as horses.
|1.2 Ma||Homo antecessor may be a common ancestor of humans and Neanderthals. At present estimate, humans have approximately 20,000–25,000 genes and share 99% of their DNA with the now extinct Neanderthal  and 95–99% of their DNA with their closest living evolutionary relative, the chimpanzees. The human variant of the FOXP2 gene (linked to the control of speech) has been found to be identical in Neanderthals.|
Three 1.5 m (5 ft) tall Homo heidelbergensis left footprints in powdery volcanic ash solidified in Italy. Homo heidelbergensis may be a common ancestor of humans and Neanderthals. It is morphologically very similar to Homo erectus but Homo heidelbergensis had a larger brain-case, about 93% the size of that of Homo sapiens. The holotype of the species was tall, 1.8 m (6 ft) and more muscular than modern humans.
Beginning of the Middle Paleolithic.
|500 ka||Divergence of Neanderthal and Denisovan lineages from a common ancestor.|
|200 ka||Omo1 and Omo2 sites (Omo River, Ethiopia) yield the earliest fossil evidence for anatomically modern Homo sapiens.
By a 2015 study, the hypothetical man Y-chromosomal Adam is estimated to have lived in East Africa about 250 ka. He would be the most recent common ancestor from whom all male human Y chromosomes are descended.
|160 ka||Homo sapiens (Homo sapiens idaltu) in Ethiopia, Awash River (near present-day Herto village) practiced mortuary rituals. Potential earliest evidence of anatomical and aspects of behavioral modernity consistent with the continuity hypothesis including use of red ochre and fishing.|
|90 ka||Appearance of mitochondrial haplogroup (mt-haplogroup) L2.|
|60 ka||Appearance of mt-haplogroups M and N, which participated in a migration out of Africa. Homo sapiens who leave Africa in this wave may have interbred with the Neanderthals they encounter.|
|50 ka||Behavioral modernity develops, according to the "great leap forward" theory.|
|40 ka||Migration to Australia and Europe; Cro-Magnon develops in Europe.|
|25–40 ka||The independent Neanderthal lineage dies out.|
|10–20 ka||Beginning of the Mesolithic / Holocene.|
- Chimpanzee-human last common ancestor
- Dawn of Humanity (film)
- Human evolution
- Human taxonomy
- Most recent common ancestor
- List of human evolution fossils
- March of Progress – famous illustration of 25 million years of human evolution
- Prehistoric amphibian
- Prehistoric fish
- Prehistoric reptile
- The Ancestor's Tale by Richard Dawkins – timeline comprising 40 rendezvous points
- Timeline of evolution – explains the evolution of animals living today
- Timeline of prehistory
- Y-DNA haplogroups by ethnic groups
- Evolutionary history of life
- Graphical timeline of the universe
- History of Earth
- History of the world
- Natural history
- "'Experiments with sex have been very hard to conduct,' Goddard said. 'In an experiment, one needs to hold all else constant, apart from the aspect of interest. This means that no higher organisms can be used, since they have to have sex to reproduce and therefore provide no asexual control.'
Goddard and colleagues instead turned to a single-celled organism, yeast, to test the idea that sex allows populations to adapt to new conditions more rapidly than asexual populations." Sex Speeds Up Evolution, Study Finds (URL accessed on January 9, 2005)
- Dawkins, R. (2005), The Ancestor's Tale: A Pilgrimage to the Dawn of Evolution, Houghton Mifflin Harcourt, ISBN 978-0-618-61916-0
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- "Obviously vertebrates must have had ancestors living in the Cambrian, but they were assumed to be invertebrate forerunners of the true vertebrates — protochordates. Pikaia has been heavily promoted as the oldest fossil protochordate." Richard Dawkins 2004 The Ancestor's Tale Page 289, ISBN 0-618-00583-8
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- These first vertebrates lacked jaws, like the living hagfish and lampreys. Jawed vertebrates appeared 100 million years later, in the Silurian. http://www.ucmp.berkeley.edu/vertebrates/vertintro.html Berkeley University
- "Bones of first gill arch became upper and lower jaws." (Image)
- A fossil coelacanth jaw found in a stratum datable 410 mya that was collected near Buchan in Victoria, Australia's East Gippsland, currently holds the record for oldest coelacanth; it was given the name Eoactinistia foreyi when it was published in September 2006. 
- "Lungfish are believed to be the closest living relatives of the tetrapods, and share a number of important characteristics with them. Among these characters are tooth enamel, separation of pulmonary blood flow from body blood flow, arrangement of the skull bones, and the presence of four similarly sized limbs with the same position and structure as the four tetrapod legs." http://www.ucmp.berkeley.edu/vertebrates/sarco/dipnoi.html Berkeley University
- "the ancestor that amphibians share with reptiles and ourselves?" "These possibly transitional fossils have been much studied, among them Acanthostega, which seems to have been wholly aquatic, and Ichthyostega" Richard Dawkins 2004 The Ancestor's Tale page 250, ISBN 0-618-00583-8
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- "In many respects, the pelycosaurs are intermediate between the reptiles and mammals" http://www.ucmp.berkeley.edu/synapsids/pelycosaurs.html Berkeley University
- "Thrinaxodon, like any fossil, should be thought of as a cousin of our ancestor, not the ancestor itself. It was a member of a group of mammal-like reptiles called the cynodonts. The cynodonts were so mammal-like, it is tempting to call them mammals. But who cares what we call them? They are almost perfect intermediates." Richard Dawkins 2004 The Ancestor's Tale page 211, ISBN 0-618-00583-8
- "A Jurassic eutherian mammal and divergence of marsupials and placentals". Nature 476: 442–445. Aug 2011. doi:10.1038/nature10291. PMID 21866158.
- "Fossils that might help us reconstruct what Concestor 8 was like include the large group called plesiadapi-forms. They lived about the right time, and they have many of the qualities you would expect of the grand ancestor of all the primates" Richard Dawkins 2004 The Ancestor's Tale page 136, ISBN 0-618-00583-8
- Raauma, Ryan; Sternera, K (2005). "Catarrhine primate divergence dates estimated from complete mitochondrial genomes" (PDF). Journal of Human Evolution 48: 237–257. doi:10.1016/j.jhevol.2004.11.007. PMID 15737392.
- Perlman, David (July 12, 2001). "Fossils From Ethiopia May Be Earliest Human Ancestor". National Geographic News. Retrieved July 2009.
Another co-author is Tim D. White, a paleoanthropologist at UC-Berkeley who in 1994 discovered a pre-human fossil, named Ardipithecus ramidus, that was then the oldest known, at 4.4 million years.
- White, Tim D.; Asfaw, Berhane; Beyene, Yonas; Haile-Selassie, Yohannes; Lovejoy, C. Owen; Suwa, Gen; WoldeGabriel, Giday (2009). "Ardipithecus ramidus and the Paleobiology of Early Hominids.". Science 326 (5949): 75–86. Bibcode:2009Sci...326...75W. doi:10.1126/science.1175802. PMID 19810190.
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- NOVA: Becoming Human Part 2 http://video.pbs.org/video/1319997127/
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- "Rubin also said analysis so far suggests human and Neanderthal DNA are some 99.5 percent to nearly 99.9 percent identical." Neanderthal bone gives DNA clues (URL accessed on November 16, 2006)
- "The conclusion is the old saw that we share 98.5% of our DNA sequence with chimpanzee is probably in error. For this sample, a better estimate would be that 95% of the base pairs are exactly shared between chimpanzee and human DNA." Britten, R.J. (2002). "Divergence between samples of chimpanzee and human DNA sequences is 5%, counting indels". PNAS 99 (21): 13633–5. Bibcode:2002PNAS...9913633B. doi:10.1073/pnas.172510699. PMC 129726. PMID 12368483.
- "...of the three billion letters that make up the human genome, only 15 million—less than 1 percent—have changed in the six million years or so since the human and chimp lineages diverged." Pollard, K.S. (2009). "What makes us human?". Scientific American. 300-5 (5): 44–49. doi:10.1038/scientificamerican0509-44
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- Mounier,Aurélien; François Marchal and Silvana Condemi "Is Homo heidelbergensis a distinct species? New insight on the Mauer mandible" Journal of Human Evolution Volume 56, Issue 3, March 2009, Pages 219–246 
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