View source for Biology:Timeline of human evolution

{{short description|Chronological outline of major events in the development of the human species}}
[[File:Age-of-Man-wiki.jpg|upright=1.2|thumb|[[Biography:Ernst Haeckel|Haeckel]]'s Paleontological Tree of Vertebrates (c.&nbsp;1879). The evolutionary history of species has been described as a "tree" with many branches arising from a single trunk. While Haeckel's tree is outdated, it illustrates clearly the principles that more complex and accurate modern reconstructions can obscure.]]
The '''timeline of human evolution''' outlines the major events in the evolutionary lineage of the modern human species, ''Homo sapiens'', 
throughout the [[Biology:History of life|history of life]], beginning some 4 billion years ago down to [[Biology:Recent human evolution|recent evolution]] within ''H. sapiens'' during and since the [[Earth:Last Glacial Period|Last Glacial Period]].

It includes brief explanations of the various [[Biology:Taxonomic rank|taxonomic rank]]s in the human lineage. The timeline reflects the  mainstream views in [[Biology:Taxonomy#Modern system of classification|modern taxonomy]], based on the principle of [[Biology:Phylogenetic nomenclature|phylogenetic nomenclature]];
in cases of open questions with no  clear consensus, the main competing possibilities are briefly outlined.

==Overview of taxonomic ranks==
A tabular overview of the [[Biology:Taxonomic rank|taxonomic rank]]ing   of ''Homo sapiens'' (with age estimates for each rank) is shown below.

{| class="wikitable"
! scope="col" | Rank
! scope="col" | Name
! scope="col" | Common name
! scope="col" | Millions of years ago (commencement)
|-
|
|Life
|
|4,200
|-
|
|[[Biology:Archaea|Archaea]]
|
|3,700
|-
|Domain||[[Biology:Eukaryote|Eukaryota]]||Eukaryotes||2,100
|-
|
|[[Biology:Podiata|Podiata]]
|Excludes Plants and their relatives
|1,540
|-
|
|[[Biology:Amorphea|Amorphea]]
|
|
|-
|
|[[Biology:Obazoa|Obazoa]]
|Excludes [[Biology:Amoebozoa|Amoebozoa]] (Amoebas)
|
|-
|
|[[Biology:Opisthokont|Opisthokont]]s
|Holozoa + [[Biology:Holomycota|Holomycota]] (Cristidicoidea and [[Biology:Fungi|Fungi]])
|1,300
|-
|
|[[Biology:Holozoa|Holozoa]]
|Excludes Holomycota
|1,100
|-
|
|[[Biology:Filozoa|Filozoa]]
|Choanozoa + Filasterea
|
|-
|
|[[Biology:Choanozoa|Choanozoa]]
|Choanoflagellates + Animals
|900
|-
|[[Biology:Kingdom|Kingdom]]||[[Biology:Animal|Animal]]ia||Animals||610
|-
|Subkingdom
|[[Biology:Eumetazoa|Eumetazoa]]
|Excludes [[Biology:Porifera|Porifera]] (Sponges)
|
|-
|
|[[Biology:ParaHoxozoa|Parahoxozoa]]
|Excludes [[Biology:Ctenophora|Ctenophora]] (Comb Jellies)
|
|-
|
|[[Biology:Bilateria|Bilateria]]
|Triploblasts / Worms||560
|-
|
|[[Biology:Nephrozoa|Nephrozoa]]
|
|
|-
|
|[[Biology:Deuterostome|Deuterostomes]]
|Division from [[Biology:Protostome|Protostome]]s
|
|-
|[[Biology:Phylum|Phylum]]||[[Biology:Chordate|Chordata]]||Chordates (Vertebrates and closely related invertebrates)||530
|-
|
|[[Biology:Olfactores|Olfactores]]
|Excludes [[Biology:Cephalochordate|cephalochordate]]s (Lancelets)
|
|-
|[[Biology:Subphylum|Subphylum]]||[[Biology:Vertebrate|Vertebrata]]||Fish / Vertebrates||505
|-
|Infraphylum||[[Biology:Gnathostomata|Gnathostomata]]||Jawed fish||460
|-
|
|[[Biology:Euteleostomi|Teleostomi]]
|Bony fish||420
|-
|
|[[Biology:Sarcopterygii|Sarcopterygii]]
|Lobe finned fish
|
|-
|Superclass||[[Biology:Tetrapod|Tetrapod]]a||Tetrapods (animals with four limbs)||395
|-
| ||[[Biology:Amniote|Amniota]]||Amniotes (fully terrestrial tetrapods whose eggs are  [[Biology:Amniote|"equipped with an amnion"]])     ||340
|-
|
|[[Biology:Synapsid|Synapsid]]a
|Proto-Mammals
|308
|-
|
|[[Biology:Therapsid|Therapsid]]
|Limbs beneath the body and other mammalian traits
|280
|-
|[[Biology:Class|Class]]||[[Biology:Mammal|Mammal]]ia||Mammals||220
|-
|[[Biology:Class|Subclass]]||[[Biology:Theria|Theria]]||Mammals that give birth to live young (i.e., non-egg-laying) ||160
|-
|Infraclass||[[Biology:Eutheria|Eutheria]]||Placental mammals  (i.e., non-marsupials) ||125
|-
|[[Biology:Magnorder|Magnorder]]||[[Biology:Boreoeutheria|Boreoeutheria]]||Supraprimates, (most) hoofed mammals, (most) carnivorous mammals, cetaceans, and bats ||124–101
|-
|Superorder||[[Biology:Euarchontoglires|Euarchontoglires]]||Supraprimates: primates, colugos, tree shrews, rodents, and rabbits ||100
|-
|[[Biology:Grandorder|Grandorder]]||[[Biology:Euarchonta|Euarchonta]]||[[Biology:Primates|Primates]], colugos, and tree shrews ||99–80
|-
|Mirorder||[[Biology:Primatomorpha|Primatomorpha]]||Primates and colugos||79.6
|-
|[[Biology:Order|Order]]||[[Biology:Primate|Primate]]s||Primates / [[Biology:Plesiadapiformes|Plesiadapiformes]]||66
|-
|Suborder||Haplorrhini||"Dry-nosed" (literally, "simple-nosed") primates:  tarsiers and monkeys (incl. apes) ||63
|-
|Infraorder||[[Biology:Simian|Simiiformes]]||monkeys (incl. apes) ||40
|-
|[[Biology:Parvorder|Parvorder]]||[[Biology:Catarrhini|Catarrhini]]||"Downward-nosed" primates: apes and old-world monkeys||30
|-
|[[Biology:Taxonomic rank|Superfamily]]||[[Biology:Hominoidea|Hominoidea]]||[[Biology:Ape|Ape]]s: great apes and lesser apes ([[Biology:Gibbon|gibbon]]s)            ||22-20
|-
|Family||[[Biology:Hominidae|Hominidae]]||Great apes: [[Biology:Human|human]]s, [[Biology:Chimpanzee|chimpanzee]]s, [[Biology:Gorilla|gorilla]]s and [[Biology:Orangutan|orangutan]]s—the hominids||20–15
|-
|Subfamily||[[Biology:Homininae|Homininae]]||Humans, chimpanzees, and gorillas (the African apes)<ref>Finarelli, J.A.; Clyde, W.C. (2004). "Reassessing hominoid phylogeny: Evaluating congruence in the morphological and temporal data". Paleobiology. 30 (4): 614.</ref> ||14–12
|-
|[[Biology:Tribe|Tribe]]||[[Biology:Hominini|Hominini]]||Includes both ''Homo'', ''Pan'' (chimpanzees), but [[Biology:Gorilla–human last common ancestor|not ''Gorilla'']].|| 10–8
|-
|[[Biology:Subtribe|Subtribe]]||[[Biology:Hominina|Hominina]]||Genus ''Homo'' and close human relatives and ancestors after [[Biology:Chimpanzee–human last common ancestor|splitting]] from ''Pan''—the hominins||8–4<ref>{{cite journal | vauthors = Patterson N, Richter DJ, Gnerre S, Lander ES, Reich D | s2cid = 2325560 | year = 2006 | title = Genetic evidence for complex speciation of humans and chimpanzees | journal = Nature | volume = 441 | issue = 7097| pages = 1103–08 | doi = 10.1038/nature04789 | pmid = 16710306 | bibcode = 2006Natur.441.1103P }}</ref>
|-
|(Genus)
|''[[Biology:Ardipithecus|Ardipithecus]] [[Biology:Sensu#Common qualifiers|s.l.]]''
|
|6-4
|-
|(Genus)
|''[[Biology:Australopithecus|Australopithecus]]''
|
|3
|-
|[[Biology:Genus|Genus]]||''[[Biology:Homo|Homo]] [[Biology:Homo habilis|(H. habilis)]]''||Humans||2.5
|-
|(Species)
|''[[Biology:Homo erectus|H. erectus]] s.l.''
|
|
|-
|(Species)
|''[[Biology:Homo heidelbergensis|H. heidelbergensis]] s.l.''
|
|
|-
|[[Biology:Species|Species]]||''Homo sapiens'' [[Biology:Sensu#Common qualifiers|s.s.]] ||Anatomically modern humans||0.8–0.3<ref>depending on the classification of the ''[[Biology:Homo heidelbergensis|Homo heidelbergensis]]'' lineage; 0.8 if Neanderthals are classed as ''H. sapiens neanderthalensis'', or if ''H. sapiens'' is defined cladistically from the divergence from ''H. neanderthalensis'', 0.3 based on the available fossil evidence.</ref>
|-
|}

==Timeline==
{{Human timeline}}
{{Life timeline}}
===Unicellular life===
{| class="wikitable"
! scope="col" | Date
! scope="col" | Event
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 4.3-4.1&nbsp;[[Year#SI prefix multipliers|Ga]]<br />
| style="vertical-align: top;" | [[Biology:Earliest known life forms|The earliest life]] appears, possibly as [[Biology:Protocell|protocell]]s. Their genetic material was probably composed of [[Biology:RNA|RNA]], capable of both self replication and enzymatic activity; their [[Biology:Cell membrane|membranes]] were composed of [[Chemistry:Lipid|lipid]]s. The [[Biology:Gene|gene]]s were separate strands, translated into [[Biology:Protein|protein]]s and often exchanged between the protocells. 
{{Further|Biology:Abiogenesis|Biology:RNA world|Biology:Earliest known life forms}}
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 4.0-3.8 Ga
|Prokaryotic [[Biology:Cell|cells]] appear; their genetic materials are composed of the more stable [[Biology:DNA|DNA]] and they use proteins for various reasons, primarily for aiding DNA to replicate itself by proteinaceous [[Biology:Enzyme|enzyme]]s (RNA now acts as an intermediary in this [[Biology:Central dogma of molecular biology|central dogma of genetic information flow]] of cellular life); [[Biology:Genotype|genes are now linked in sequences]] so all information passes to offsprings. They had [[Biology:Cell wall|cell wall]]s & [[Biology:Bacterial outer membrane|outer membrane]]s and were probably initially [[Earth:Thermophile|thermophile]]s.

|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 3.5 Ga
| This marks the first appearance of [[Biology:Cyanobacteria|cyanobacteria]] and their method of oxygenic photosynthesis and therefore the first occurrence of atmospheric [[Chemistry:Oxygen|oxygen]] on Earth.

For another billion years, prokaryotes would continue to diversify undisturbed.

|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 2.5-2.2 Ga
| First organisms to use oxygen. By 2400 Ma, in what is referred to as the [[Earth:Great Oxidation Event|Great Oxidation Event]], (GOE), most of the pre-oxygen anaerobic forms of life were wiped out by the oxygen producers.

{{Further|Earth:Geological history of oxygen}}
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 2.2-1.8 Ga
|Origin of the [[Biology:Eukaryote|eukaryote]]s: organisms with [[Biology:Cell nucleus|nuclei]], [[Biology:Endomembrane system|endomembrane system]]s (including [[Biology:Mitochondria|mitochondria]]) and complex [[Biology:Cytoskeleton|cytoskeleton]]s; they [[Biology:RNA splicing|spliced]] [[Biology:MRNA|mRNA]] between [[Biology:Transcription|transcription]] and [[Biology:Translation|translation]] (splicing also occurs in prokaryotes, but it is only of [[Biology:Non-coding RNA|non-coding RNA]]s). The evolution of eukaryotes, and possibly sex, is thought to be related to the GOE, as it probably pressured two or three lineages of prokaryotes (including an [[Biology:Aerobic organism|aerobe]] one, which later became mitochondria) to depend on each other, leading to endosymbiosis. Early eukaryotes lost their cell walls and outer membranes.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 1.2 Ga
| [[Biology:Evolution of sexual reproduction|Sexual reproduction evolves]] ([[Biology:Mitosis|mitosis]] and [[Biology:Meiosis|meiosis]]) by this time at least, leading to faster evolution<ref>"'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.'<br />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.<cite></cite>" [https://web.archive.org/web/20050404004325/http://news.nationalgeographic.com/news/2005/03/0330_050330_sexevolution.html Sex Speeds Up Evolution, Study Finds] (URL accessed on January 9, 2005)</ref> where genes are mixed in every generation enabling greater variation for subsequent selection.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 1.2-0.8 Ga
| [[File:Cronoflagelado2.svg|thumb|150px|[[Biology:Choanoflagellate|Choanoflagellate]]]]
The [[Biology:Holozoa|Holozoa]] lineage of eukaryotes evolves many features for making [[Biology:Colony|cell colonies]], and finally leads to the ancestor of animals (metazoans) and choanoflagellates.<ref>Dawkins, R. (2005), The Ancestor's Tale: A Pilgrimage to the Dawn of Evolution, Houghton Mifflin Harcourt, {{ISBN|978-0-618-61916-0}}</ref><ref>"Proterospongia is a rare freshwater protist, a colonial member of the Choanoflagellata." "Proterospongia itself is not the ancestor of sponges. However, it serves as a useful model for what the ancestor of sponges and other metazoans may have been like."  http://www.ucmp.berkeley.edu/protista/proterospongia.html Berkeley University</ref>

[[Biology:Proterospongia|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 [[Biology:Cell|cellular]] specialization for different tasks.
|-
|}

===Animalia===
{| class="wikitable"
! scope="col" | Date
! scope="col" | Event
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 800–650 [[Astronomy:Myr|Ma]]
|[[File:DickinsoniaCostata.jpg|thumb|200px|''[[Biology:Dickinsonia|Dickinsonia costata]]'' from the Ediacaran biota, 635–542 Ma, a possible early member of ''Animalia''.]]
[[Biology:Urmetazoan|Urmetazoan]]:
The first fossils that might represent [[Biology:Animal|animal]]s appear in the 665-million-year-old rocks of the Trezona Formation of South Australia. These fossils are interpreted as being early sponges.<ref>{{cite journal |title=Possible animal-body fossils in pre-Marinoan limestones from South Australia |journal=Nature Geoscience |volume=3 |pages=653–59 |date=17 August 2010 |doi=10.1038/ngeo934 |issue=9 |bibcode=2010NatGe...3..653M |last1=Maloof |first1=Adam C. |last2=Rose |first2=Catherine V. |last3=Beach |first3=Robert |last4=Samuels |first4=Bradley M. |last5=Calmet |first5=Claire C. |last6=Erwin |first6=Douglas H. |last7=Poirier |first7=Gerald R. |last8=Yao |first8=Nan |last9=Simons |first9=Frederik J.|s2cid=13171894 }}</ref> Multicellular animals may have existed from 800 Ma. Separation from the [[Biology:Porifera|Porifera]] ([[Biology:Sponge|sponge]]s) lineage.
[[Biology:Eumetazoa|Eumetazoa]]/Diploblast: separation from the [[Biology:Ctenophora|Ctenophora]] ("comb jellies") lineage.
Planulozoa/[[Biology:ParaHoxozoa|ParaHoxozoa]]: separation from the [[Biology:Placozoa|Placozoa]] and [[Biology:Cnidaria|Cnidaria]]  lineages.
All diploblasts possess epithelia, nerves, [[Biology:Muscle|muscle]]s and connective tissue and [[Biology:Mouth|mouth]]s, and except for placozoans, have some form of symmetry, with their ancestors probably having radial symmetry like that of cnidarians. Diploblasts separated their early embryonic cells into two germ layers ([[Biology:Ectoderm|ecto-]] and [[Biology:Endoderm|endoderm]]). [[Biology:Photoreceptor cell|Photoreceptive]] eye-spots evolve.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 650-600 Ma
| [[File:Proporus sp.png|thumb|200px|''[[Biology:Proporidae|Proporus]]'' sp., a xenacoelomorph.]]
[[Biology:Urbilaterian|Urbilaterian]]:
the last common ancestor of xenacoelomorphs, protostomes (including the [[Biology:Arthropod|arthropod]] [insect, crustacean, spider], [[Biology:Mollusc|mollusc]] [squid, snail, clam] and [[Biology:Annelid|annelid]] [earthworm] lineages) and the deuterostomes (including the vertebrate [human] lineage) (the last two are more related to each other and called [[Biology:Nephrozoa|Nephrozoa]]). Xenacoelomorphs all have a [[Biology:Gonopore|gonopore]] to expel [[Biology:Gamete|gamete]]s but nephrozoans merged it with their [[Biology:Anus|anus]]. Earliest development of bilateral symmetry, [[Biology:Mesoderm|mesoderm]], head (anterior [[Biology:Cephalization|cephalization]]) and various gut muscles (and thus [[Biology:Peristalsis|peristalsis]]) and, in the Nephrozoa, nephridia (kidney precursors), [[Biology:Coelom|coelom]] (or maybe pseudocoelom), distinct mouth and anus (evolution of through-gut), and possibly even [[Biology:Ventral nerve cord|nerve cords]] and [[Biology:Blood vessel|blood vessel]]s.<ref>Monahan-Earley, R., Dvorak, A. M., & Aird, W. C. (2013). Evolutionary origins of the blood vascular system and endothelium. ''Journal of Thrombosis and Haemostasis'', '''11''' (Suppl 1), 46–66.</ref> Reproductive tissue probably concentrates into a pair of gonads connecting just before the posterior orifice. "Cup-eyes" and [[Biology:Statocyst|balance organs]] evolve (the function of hearing added later as the more complex inner ear evolves in vertebrates). The nephrozoan through-gut had a wider portion in the front, called the [[Biology:Pharynx|pharynx]]. The [[Biology:Integument|integument]] or skin consists of an epithelial layer ([[Biology:Epidermis|epidermis]]) and a connective layer.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 600-540 Ma
|[[File:Actinopyga echinites1.jpg|thumb|200px|A [[Biology:Sea cucumber|sea cucumber]] (''Actinopyga echinites''), displaying its feeding tentacles and tube feet.]]
Most known animal phyla appeared in the fossil record as marine species during the '''[[Earth:Ediacaran|Ediacaran]]-[[Earth:Cambrian explosion|Cambrian explosion]]''', probably caused by long scale oxygenation since around 585 Ma (sometimes called the Neoproterozoic Oxygenation Event or NOE) and also an influx of oceanic minerals. Deuterostomes, the last common ancestor of the [[Biology:Chordata|Chordata]] [human] lineage, [[Biology:Hemichordata|Hemichordata]] ([[Biology:Acorn worm|acorn worm]]s and [[Biology:Graptolite|graptolite]]s) and [[Biology:Echinodermata|Echinodermata]] ([[Biology:Starfish|starfish]], [[Biology:Sea urchin|sea urchin]]s, [[Biology:Holothuroidea|sea cucumbers]], etc.), probably had both ventral and [[Biology:Dorsal nerve cord|dorsal nerve cord]]s like modern acorn worms.

An archaic survivor from this stage is the [[Biology:Acorn worm|acorn worm]], sporting an  open circulatory system (with less branched blood vessels) with a heart that also functions as a kidney. Acorn worms have a  [[Biology:Plexus|plexus]] concentrated into both dorsal and ventral nerve cords. The dorsal cord reaches into the proboscis, and is partially separated from the epidermis in that region. This part of the dorsal nerve cord is often hollow, and may well be homologous with the brain of vertebrates.<ref>{{cite book |author= Barnes, Robert D. |year=1982 |title= Invertebrate Zoology |publisher= Holt-Saunders International |location= Philadelphia|pages= 1018–26|isbn= 978-0-03-056747-6}}</ref> Deuterostomes also evolved [[Biology:Pharyngeal slit|pharyngeal slit]]s, which were probably used for filter feeding like in hemi- and proto-chordates.
|}

===Chordata===
{| class="wikitable"
! scope="col" | Date
! scope="col" | Event
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 540-520 Ma
|[[File:Pikaia BW.jpg|thumb|''[[Biology:Pikaia|Pikaia]]'']] 
The increased amount of oxygen causes many eukaryotes, including most animals, to become [[Biology:Obligate aerobe|obligate aerobe]]s.
 
The Chordata ancestor gave rise to the lancelets (Amphioxii) and Olfactores. Ancestral chordates evolved a post-anal [[Biology:Tail|tail]], notochord, and endostyle (precursor of thyroid). The pharyngeal slits (or gills) are now supported by connective tissue and used for filter feeding and possibly [[Physics:Breathing|breathing]].<ref>"Obviously vertebrates must have had ancestors living in the Cambrian, but they were assumed to be invertebrate forerunners of the true vertebrates — proto-chordates. ''Pikaia'' has been heavily promoted as the oldest fossil protochordate." [[Biography:Richard Dawkins|Richard Dawkins]] 2004 ''[[Biology:The Ancestor's Tale|The Ancestor's Tale]]'' p. 289, {{ISBN|0-618-00583-8}}</ref> Other, earlier chordate predecessors include ''[[Biology:Myllokunmingia|Myllokunmingia]] fengjiaoa'',<ref>{{Cite journal | last1 = Shu | first1 = D.G. | title = Lower Cambrian vertebrates from south China | last2 = Luo | first2 = H.L. | last3 = Conway Morris | first3 = S. | last4 = Zhang | first4 = X. L. | last5 = Hu | first5 = S.X. | last6 = Chen | first6 = L. | last7 = Han | first7 = J. | last8 = Zhu | first8 = M. | last9 = Li | first9 = Y. | last10 = Chen | first10 = L.Z. | s2cid = 4402854 | journal = Nature | volume = 402 | issue = 6757 | pages = 42–46 | year = 1999 | doi = 10.1038/46965|bibcode = 1999Natur.402...42S }}</ref> ''Haikouella lanceolata'',<ref>{{Cite journal | last1 = Chen | first1 = J.Y. | title = An early Cambrian craniate-like chordate | last2 = Huang | first2 = D.Y. | last3 = Li | first3 = C.W. | s2cid = 24895681 | journal = Nature | volume = 402 | pages = 518–22 | year = 1999 | doi = 10.1038/990080|bibcode = 1999Natur.402..518C | issue=6761}}</ref> and ''[[Biology:Haikouichthys|Haikouichthys]] ercaicunensis''.<ref>{{cite journal |last1=Shu |first1=D.-G. |last2=Morris |first2=S. Conway |last3=Han |first3=J. |last4=Zhang |first4=Z.-F. |last5=Yasui |first5=K. |last6=Janvier |first6=P. |last7=Chen |first7=L. |last8=Zhang |first8=X.-L. |last9=Liu |first9=J.-N. |last10=Li |first10=Y. |last11=Liu |first11=H.-Q. |s2cid=4401274 |title=Head and backbone of the Early Cambrian vertebrate Haikouichthys |journal=Nature |date=January 2003 |volume=421 |issue=6922 |pages=526–529 |doi=10.1038/nature01264 |pmid=12556891 |bibcode=2003Natur.421..526S }}</ref> They probably lost their ventral nerve cord and evolved a special region of the [[Biology:Central nervous system|dorsal one]], called the [[Biology:Brain|brain]], with glia becoming permanently associated with neurons. They probably evolved the first [[Biology:Blood cell|blood cell]]s (probably early leukocytes, indicating advanced innate immunity), which they made around the pharynx and gut.<ref>Udroiu, I., &amp; Sgura, A. (2017). The phylogeny of the spleen. The Quarterly Review of Biology, 92(4), 411–443. https://doi.org/10.1086/695327</ref> All chordates except tunicates sport an intricate, closed circulatory system, with highly branched blood vessels.

[[Biology:Olfactores|Olfactores]], last common ancestor of [[Biology:Tunicate|tunicate]]s and vertebrates in which [[Biology:Sense of smell|olfaction (smell)]] evolved. Since lancelets lack a [[Biology:Heart|heart]], it possibly emerged in this ancestor (previously the blood vessels themselves were contractile) though it could have been lost in lancelets after evolving in early deuterostomes (hemichordates and echinoderms have hearts).

|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 520-480 Ma
|[[File:Agnata.png|thumb|upright=0.9|Agnatha]]

The first [[Biology:Vertebrate|vertebrate]]s ("[[Biology:Fish|fish]]") appear: the [[Biology:Ostracoderm|ostracoderm]]s. ''[[Biology:Haikouichthys|Haikouichthys]]'' and ''[[Biology:Myllokunmingia|Myllokunmingia]]'' are examples of these jawless fish, or [[Biology:Agnatha|Agnatha]]; the jawless [[Biology:Cyclostomata|Cyclostomata]] diverge at this stage. They were jawless, had seven pairs of [[Biology:Pharyngeal arch|pharyngeal arch]]es like their descendants today, and their [[Biology:Endoskeleton|endoskeleton]]s were cartilaginous (then only consisting of the [[Biology:Chondrocranium|chondro]]cranium/braincase and vertebrae). The connective tissue below the epidermis differentiates into the [[Biology:Dermis|dermis]] and hypodermis.<ref>Elliot D.G. (2011) Functional Morphology of the Integumentary System in Fishes. In: Farrell A.P., (ed.), Encyclopedia of Fish Physiology: From Genome to Environment, volume 1, pp. 476–488. San Diego: Academic Press</ref> They depended on [[Biology:Fish gill|gills]] for respiration and evolved the unique sense of [[Biology:Taste|taste]] (the remaining sense of the skin now called "touch"), endothelia, camera [[Biology:Eye|eye]]s and inner ears (capable of hearing and balancing; each consists of a lagena, an [[Biology:Otolith|otolith]]ic organ and two [[Biology:Semicircular canals|semicircular canals]]) as well as [[Biology:Liver|liver]]s, [[Biology:Thyroid|thyroid]]s, kidneys and two-chambered hearts (one atrium and one [[Medicine:Ventricle (heart)|ventricle]]). They had a tail [[Biology:Fish fin|fin]] but lacked the paired (pectoral and pelvic) fins of more advanced fish. Brain divided into three parts (further division created distinct regions based on function). The [[Biology:Pineal gland|pineal gland]] of the brain penetrates to the level of the skin on the head, making it seem like a third eye. They evolved the first erythrocytes and thrombocytes.<ref>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</ref>
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 460-430 Ma
|[[File:Coccosteus BW.jpg|thumb|upright=0.9|A placoderm]]
The [[Biology:Placodermi|Placodermi]] were the first jawed fishes ([[Biology:Gnathostomata|Gnathostomata]]); their [[Medicine:Jaw|jaw]]s evolved from the first [[Biology:Branchial arch|gill]]/pharyngeal arch and they largely replaced their endoskeletal [[Biology:Cartilage|cartilage]] with [[Biology:Bone|bone]] and evolved pectoral and pelvic fins. Bones of the first gill arch became the [[Biology:Palatoquadrate|upper]] and [[Biology:Meckel's cartilage|lower]] jaw, while those from the second arch became the [[Biology:Hyomandibula|hyomandibula]], ceratohyal and basihyal; this closed two of the seven pairs of gills. The gap between the first and second arches just below the braincase (fused with upper jaw) created a pair of spiracles, which opened in the skin and led to the pharynx (water passed through them and left through gills). 
Placoderms had competition with the previous dominant animals, the cephalopods and [[Biology:Eurypterid|sea scorpions]], and rose to dominance themselves. A lineage of them probably evolved into the bony and cartilaginous fish, after evolving [[Biology:Scale (anatomy)#Fish scales|scales]], teeth (which allowed the transition to full carnivory), [[Biology:Stomach|stomach]]s, [[Biology:Spleen|spleen]]s, [[Biology:Thymus|thymus]]es, myelin sheaths, [[Biology:Hemoglobin|hemoglobin]] and advanced, adaptive immunity (the latter two occurred independently in the lampreys and hagfish). Jawed fish also have a third, lateral semicircular canal and their otoliths are divided between a saccule and utricle.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 430-410 Ma
|[[File:Latimeria Chalumnae - Coelacanth - NHMW.jpg|thumb|upright=0.9|[[Biology:Coelacanth|Coelacanth]] caught in 1974]]Bony fish split their jaws into several bones and evolve lungs, fin bones, two pairs of rib bones, and opercular bones, and diverge into the actinopterygii (with ray fins) and the sarcopterygii (with fleshy, lower fins);<ref>A fossil coelacanth jaw found in a stratum datable 410 [[Physics:Mya (unit)|mya]] that was collected near Buchan in Victoria, {{wipe|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. [http://www.int.iol.co.za/index.php?set_id=1&click_id=31&art_id=qw1156833901231B223]</ref> the latter transitioned from marine to freshwater habitats. Jawed fish also possess dorsal and anal fins.
|}

===Tetrapoda===
{{Further|Biology:Evolution of tetrapods}}
{| class="wikitable"
! scope="col" | Date
! scope="col" | Event
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 390 Ma
|[[File:Panderichthys BW.jpg|thumb|upright=0.9|''Panderichthys'']]
Some freshwater lobe-finned fish (sarcopterygii) develop limbs and give rise to the [[Biology:Tetrapodomorpha|Tetrapodomorpha]]. These fish evolved in shallow and [[Earth:Swamp|swamp]]y freshwater habitats, where they evolved large eyes and spiracles.

Primitive tetrapods ("fishapods") developed from tetrapodomorphs with a two-lobed [[Biology:Brain|brain]] in a flattened skull, a wide mouth and a medium snout, whose upward-facing eyes show that it was a bottom-dweller, and which had already developed adaptations of fins with fleshy bases and [[Biology:Bone|bone]]s. (The "living fossil" [[Biology:Coelacanth|coelacanth]] is a related [[Biology:Lobe-finned fish|lobe-finned fish]] without these shallow-water adaptations.) Tetrapod fishes used their fins as paddles in shallow-water habitats choked with plants and [[Earth:Detritus|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.<ref name="ucmp.berkeley.edu">"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</ref>

''[[Biology:Panderichthys|Panderichthys]]'' is a 90–130&nbsp;cm (35–50&nbsp;in) long fish from the Late Devonian period (380 [[Physics:Mya (unit)#Symbols y and yr|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 ''[[Biology:Ichthyostega|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''.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 375-350 Ma
|[[File:Tiktaalik BW.jpg|thumb|upright=0.9|''Tiktaalik'']]

''[[Biology:Tiktaalik|Tiktaalik]]'' is a genus of [[Biology:Sarcopterygii|sarcopterygian]] (lobe-finned) fishes from the late Devonian with many tetrapod-like features.  It shows a clear link between ''Panderichthys'' and ''Acanthostega''.

[[File:Acanthostega BW.jpg|thumb|upright=0.9|''Acanthostega'']]
[[File:Ichthyostega BW.jpg|thumb|upright=0.9|''Ichthyostega'']]
''[[Biology:Acanthostega|Acanthostega]]'' is an extinct [[Biology:Tetrapod|tetrapod]], 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. The dorsal pair of ribs form a rib cage to support the lungs, while the ventral pair disappears.

''[[Biology:Ichthyostega|Ichthyostega]]'' is another extinct tetrapod. Being one of the first animals with only two pairs of limbs (also unique since they end in digits and have bones), ''Ichthyostega'' is seen as an intermediate between a fish and an amphibian. ''[[Biology:Ichthyostega|Ichthyostega]]'' had limbs but these probably were not used for walking. They may have spent very brief periods out of water and would have used their limbs to paw their way through the [[Earth:Mud|mud]].<ref>"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''" [[Biography:Richard Dawkins|Richard Dawkins]] 2004 ''[[Biology:The Ancestor's Tale|The Ancestor's Tale]]'' p. 250, {{ISBN|0-618-00583-8}}</ref> They both had more than five digits (eight or seven) at the end of each of their limbs, and their bodies were scaleless (except their bellies, where they remained as [[Biology:Gastralia|gastralia]]). Many evolutionary changes occurred at this stage: eyelids and [[Biology:Tears|tear]] glands evolved to keep the eyes wet out of water and the eyes became connected to the pharynx for draining the liquid; the hyomandibula (now called [[Biology:Columella (auditory system)|columella]]) shrank into the spiracle, which now also connected to the inner ear at one side and the pharynx at another, becoming the [[Medicine:Eustachian tube|Eustachian tube]] (columella assisted in hearing); an early [[Medicine:Eardrum|eardrum]] (a patch of connective tissue) evolved on the end of each tube (called the [[Medicine:Otic notch|otic notch]]); and the ceratohyal and basihyal merged into the hyoid. These "fishapods" had more [[Biology:Ossification|ossified and stronger]] bones to support themselves on land (especially skull and [[Biology:Appendicular skeleton|limb bones]]). Jaw bones fuse together while gill and opercular bones disappear.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 350-330 Ma
|[[File:Pederpes22small.jpg|thumb|upright=0.9|''Pederpes'']]
''Pederpes'' from around 350 Ma indicates that the standard number of 5 digits evolved at the Early Carboniferous, when modern tetrapods (or "amphibians") split in two directions (one leading to the extant amphibians and the other to amniotes). At this stage, our ancestors evolved [[Biology:Vomeronasal organ|vomeronasal organ]]s, salivary glands, [[Biology:Tongue|tongue]]s, parathyroid glands, three-chambered hearts (with two atria and one ventricle) and [[Biology:Urinary bladder|bladders]], and completely removed their gills by adulthood. The [[Medicine:Glottis|glottis]] evolves to prevent food going into the [[Medicine:Respiratory tract|respiratory tract]]. Lungs and thin, moist skin allowed them to breathe; water was also needed to give birth to shell-less [[Biology:Egg|eggs]] and for early development. Dorsal, anal and tail fins all disappeared.

[[Biology:Lissamphibia|Lissamphibia]] (extant amphibians) retain many features of early amphibians but they have only four digits (caecilians have none).
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 330-300 Ma
|[[File:Hylonomus BW.jpg|thumb|upright=0.9|''Hylonomus'']]
From amphibians came the first reptiles: ''[[Biology:Hylonomus|Hylonomus]]'' is the earliest known [[Biology:Reptile|reptile]]. It was 20&nbsp;cm (8&nbsp;in) long (including the tail) and probably would have looked rather similar to modern [[Biology:Lizard|lizard]]s. It had small sharp teeth and probably ate small [[Biology:Millipede|millipede]]s and [[Biology:Insect|insect]]s. It is a precursor of later [[Biology:Amniote|amniote]]s (broadest sense of "reptile"). [[Biology:Keratin|Alpha keratin]] first evolves here; it is used in the claws of modern amniotes, and hair in mammals, indicating claws and a different type of [[Biology:Scale (anatomy)#Reptilian scales|scales]] evolved in amniotes (complete loss of gills as well).<ref>{{cite journal |last1=Eckhart |first1=L. |last2=Valle |first2=L. D. |last3=Jaeger |first3=K. |last4=Ballaun |first4=C. |last5=Szabo |first5=S. |last6=Nardi |first6=A. |last7=Buchberger |first7=M. |last8=Hermann |first8=M. |last9=Alibardi |first9=L. |last10=Tschachler |first10=E. |title=Identification of reptilian genes encoding hair keratin-like proteins suggests a new scenario for the evolutionary origin of hair |journal=Proceedings of the National Academy of Sciences |date=10 November 2008 |volume=105 |issue=47 |pages=18419–18423 |doi=10.1073/pnas.0805154105 |pmid=19001262 |pmc=2587626 |doi-access=free }}</ref>

Evolution of the amniotic egg gives rise to the amniotes, tetrapods that can reproduce on land and lay [[Biology:Eggshell|shelled eggs]] on dry land. They did not need to return to water for reproduction nor breathing. This adaptation and the desiccation-resistant scales gave them the capability to inhabit the uplands for the first time, albeit making them drink water through their mouths. At this stage, adrenal tissue may have concentrated into discrete glands.

Amniotes have advanced nervous systems, with twelve pairs of cranial nerves, unlike lower vertebrates. They also evolved true [[Biology:Sternum|sternum]]s but lost their eardrums and otic notches (hearing only by columella bone conduction).
|}

===Mammals===
{{Further|Biology:Evolution of mammals}}
{| class="wikitable"
! scope="col" | Date
! scope="col" | Event
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 300-260 Ma
| Shortly after the appearance of the first reptiles, two branches split off. One branch is the [[Biology:Sauropsida|Sauropsida]], from which come the modern reptiles and birds. The other branch is [[Biology:Synapsid|Synapsid]]a from which come modern mammals.  Both had [[Biology:Temporal fenestra|temporal fenestra]]e, 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 [[Biology:Diapsid|diapsid]]s (a branch of Sauropsida) had two. An early, inefficient version of diaphragm may have evolved in synapsids.

The earliest "mammal-like reptiles" are the [[Biology:Pelycosaur|pelycosaur]]s. The pelycosaurs were the first animals to have temporal fenestrae. Pelycosaurs were not [[Biology:Therapsid|therapsid]]s but their ancestors. The therapsids were, in turn, the ancestors of mammals.

The therapsids had temporal fenestrae larger and more mammal-like than pelycosaurs, their teeth showed more serial differentiation, their gait was semi-erect 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.<ref>"In many respects, the pelycosaurs are intermediate between the reptiles and mammals" http://www.ucmp.berkeley.edu/synapsids/pelycosaurs.html Berkeley University</ref> They had lost gastralia and, possibly, scales.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 260-230 Ma
|[[File:Cynognathus BW.jpg|right|thumb|upright=0.9|''[[Biology:Cynognathus|Cynognathus]]'']]
One subgroup of therapsids, the cynodonts, lose pineal eye & lumbar ribs and very likely became [[Biology:Warm-blooded|warm-blooded]]. The lower respiratory tract forms intricate branches in the lung [[Biology:Parenchyma|parenchyma]], ending in highly vascularized alveoli. Erythrocytes and thrombocytes lose their nuclei while [[Biology:Lymphatic system|lymphatic system]]s and advanced immunity emerge. They may have also had thicker [[Biology:Dermis|dermis]] like mammals today.

The jaws of cynodonts resembled modern mammal jaws; the anterior portion, the dentary, held differentiated teeth. This group of animals likely contains a species which is the ancestor of all modern mammals. Their temporal fenestrae merged with their [[Biology:Orbit (anatomy)|orbits]]. Their hindlimbs became erect and their posterior bones of the jaw progressively shrunk to the region of the [[Biology:Columella (auditory system)|columella]].<ref>"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." [[Biography:Richard Dawkins|Richard Dawkins]] 2004 ''[[Biology:The Ancestor's Tale|The Ancestor's Tale]]'' p. 211, {{ISBN|0-618-00583-8}}</ref>
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 230-170 Ma
|[[File:Repenomamus BW.jpg|right|thumb|upright=0.9|''[[Biology:Repenomamus|Repenomamus]]'']]
From [[Biology:Eucynodontia|Eucynodontia]] came the first [[Biology:Mammal|mammal]]s. Most early mammals were small shrew-like animals that fed on insects and had transitioned to [[Biology:Nocturnality|nocturnality]] to avoid competition with the dominant archosaurs — this led to the loss of the vision of [[Red|red]] and [[Physics:Ultraviolet|ultraviolet]] light (ancestral tetrachromacy of vertebrates reduced to [[Medicine:Dichromacy|dichromacy]]). Although there is no evidence in the fossil record, it is likely that these animals had a [[Physics:Thermoregulation|constant body temperature]], hair and milk glands for their young (the glands stemmed from the milk line). The [[Biology:Neocortex|neocortex]] (part of the cerebrum) region of the brain evolves in Mammalia, at the reduction of the tectum (non-smell senses which were processed here became integrated into neocortex but smell became primary sense). Origin of the [[Biology:Prostate|prostate]] gland and a pair of holes opening to the columella and nearby shrinking jaw bones; new [[Medicine:Eardrum|eardrum]]s stand in front of the columella and Eustachian tube. The skin becomes hairy, glandular (glands secreting sebum and sweat) and thermoregulatory. Teeth fully differentiate into incisors, [[Biology:Canine tooth|canines]], premolars and molars; mammals become [[Biology:Diphyodont|diphyodont]] and possess developed diaphragms and males have internal penises. All mammals have four chambered [[Biology:Heart|heart]]s (with two atria and two ventricles) and lack cervical ribs (now mammals only have thoracic ribs).

[[Biology:Monotreme|Monotreme]]s are an egg-laying group of mammals represented today by the [[Biology:Platypus|platypus]] and [[Biology:Echidna|echidna]].  Recent genome sequencing of the platypus indicates that its sex genes are closer to those of birds than to those of the [[Biology:Theria|theria]]n (live birthing) mammals. Comparing this to other mammals, it can be inferred that the first mammals to gain [[Biology:Sexual differentiation|sexual differentiation]] through the existence or lack of SRY gene (found in the y-Chromosome) evolved only in the therians. Early mammals and possibly their eucynodontian ancestors had epipubic bones, which serve to hold the pouch in modern marsupials (in both sexes).
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 170-120 Ma
|[[File:Juramaia NT.jpg|right|thumb|upright=0.9|''[[Biology:Juramaia|Juramaia sinensis]]'']]
Evolution of live birth (viviparity), with early therians probably having pouches for keeping their undeveloped young like in modern marsupials. [[Biology:Nipple|Nipple]]s stemmed out of the therian milk lines. The [[Biology:Cloaca|posterior orifice]] separates into anal and urogenital openings; males possess an external penis.

Monotremes and therians independently detach the malleus and incus from the dentary (lower jaw) and combine them to the shrunken columella (now called [[Biology:Stapes|stapes]]) in the [[Medicine:Tympanic cavity|tympanic cavity]] behind the eardrum (which is connected to the malleus and held by another bone detached from the dentary, the tympanic plus ectotympanic), and coil their lagena ([[Medicine:Cochlea|cochlea]]) to advance their hearing, with therians further evolving an external pinna and erect forelimbs. Female [[Biology:Placentalia|placentalia]]n mammals do not have pouches and epipubic bones but instead have a developed [[Biology:Placenta|placenta]] which penetrates the [[Biology:Uterus|uterus]] walls (unlike marsupials), allowing a longer gestation; they also have separated urinary and genital openings.<ref>Werneburg, Ingmar; Spiekman, Stephan N F (2018). 4. Mammalian embryology and organogenesis. In:
Zachos, Frank; Asher, Robert. Mammalian Evolution, Diversity and Systematics. Berlin: Walter de
Gruyter, 59-116. DOI: https://doi.org/10.1515/9783110341553-004</ref>
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 100-90 Ma
| Last common ancestor of rodents, rabbits, [[Biology:Ungulates|ungulates]], carnivorans, bats, shrews and humans (base of the clade [[Biology:Boreoeutheria|Boreoeutheria]]; males now have external testicles).
|}

===Primates===
{| class="wikitable"
! scope="col" | Date
! scope="col" | Event
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 90–66 Ma
|[[File:Plesiadapis2.jpg|right|thumb|upright=0.9|''Plesiadapis'']]
[[File:CarpolestesCL.png|right|thumb|upright=0.9|''Carpolestes simpsoni'']]
A group of small, nocturnal, arboreal, insect-eating mammals called [[Biology:Euarchonta|Euarchonta]] begins a speciation that will lead to the [[Biology:Order|orders]] of [[Biology:Primate|primate]]s, treeshrews and flying lemurs. They reduced the number of mammaries to only two pairs (on the chest). [[Biology:Primatomorpha|Primatomorpha]] is a subdivision of Euarchonta including primates and their ancestral stem-primates [[Biology:Plesiadapiformes|Plesiadapiformes]]. An early stem-primate, ''[[Biology:Plesiadapis|Plesiadapis]]'', still had claws and eyes on the side of the head, making it faster on the ground than in the trees, but it began to spend long times on lower branches, feeding on fruits and leaves.

The Plesiadapiformes very likely contain the ancestor species of all primates.<ref>"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" [[Biography:Richard Dawkins|Richard Dawkins]] 2004 ''[[Biology:The Ancestor's Tale|The Ancestor's Tale]]'' p. 136, {{ISBN|0-618-00583-8}}</ref>  They first appeared in the fossil record around 66 million years ago, soon after the [[Earth:Cretaceous–Paleogene extinction event|Cretaceous–Paleogene extinction event]] that eliminated about three-quarters of plant and animal species on Earth, including most dinosaurs.<ref name="Renne2013">{{cite journal |last1=Renne |first1=Paul R. |last2=Deino |first2=Alan L. |last3=Hilgen |first3=Frederik J. |last4=Kuiper |first4=Klaudia F. |last5=Mark |first5=Darren F. |last6=Mitchell |first6=William S. |last7=Morgan |first7=Leah E. |last8=Mundil |first8=Roland |last9=Smit |first9=Jan |s2cid=6112274 |title=Time Scales of Critical Events Around the Cretaceous-Paleogene Boundary |journal=Science |date=7 February 2013 |volume=339 |issue=6120 |pages=684–87 |doi=10.1126/science.1230492 |pmid=23393261 |bibcode=2013Sci...339..684R|url=https://semanticscholar.org/paper/411afc826d678575392ec4d2dc7a6f25c389c7d0 }}</ref><ref>[http://www.physorg.com/news88774682.html "Paleontologists discover most primitive primate skeleton"], Phys.org (January 23, 2007).</ref>

One of the last Plesiadapiformes is ''[[Biology:Carpolestes simpsoni|Carpolestes simpsoni]]'', having grasping digits but not forward-facing eyes.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 66-56 Ma
|[[Biology:Primate|Primate]]s diverge into suborders [[Biology:Strepsirrhini|Strepsirrhini]] (wet-nosed primates) and Haplorrhini (dry-nosed primates). Brain expands and cerebrum divides into 4 pairs of lobes. The [[Biology:Postorbital bar|postorbital bar]] evolves to separate the orbit from the temporal fossae as sight regains its position as the primary sense;  eyes became forward-facing. Strepsirrhini contain most [[Biology:Prosimian|prosimian]]s; modern examples include lemurs and [[Biology:Loris|loris]]es. The haplorrhines include the two living groups: prosimian [[Biology:Tarsier|tarsier]]s, and simian [[Biology:Monkey|monkey]]s, including [[Biology:Ape|ape]]s. The Haplorrhini metabolism lost the ability to produce [[Biology:Vitamin C|vitamin C]], forcing all descendants to include vitamin C-containing fruit in their diet. Early primates only had claws in their second digits; the rest were turned into nails.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 50-35 Ma
|[[File:Aegyptopithecus NT.jpg|right|thumb|upright=0.9|''Aegyptopithecus'']]
Simians split into infraorders Platyrrhini and [[Biology:Catarrhini|Catarrhini]]. They fully transitioned to [[Biology:Diurnality|diurnality]] and lacked any claw and [[Physics:Tapetum lucidum|tapetum lucidum]] (which evolved many times in various vertebrates). They possibly evolved at least some of the [[Medicine:Paranasal sinuses|paranasal sinuses]], and transitioned from estrous cycle to [[Medicine:Menstrual cycle|menstrual cycle]]. The number of mammaries is now reduced to only one thoracic pair. Platyrrhines, New World monkeys, have prehensile tails and males are color blind. The individuals whose descendants would become Platyrrhini are conjectured to have migrated to South America either on a [[Biology:Oceanic dispersal|raft of vegetation]] or via a [[Earth:Land bridge|land bridge]] (the hypothesis now favored<ref>Alan de Queiroz, ''The Monkey's Voyage'', Basic Books, 2014.</ref>). Catarrhines mostly stayed in [[Earth:Africa|Africa]] as the two continents drifted apart. Possible early ancestors of catarrhines include ''[[Biology:Aegyptopithecus|Aegyptopithecus]]'' and ''[[Biology:Saadanius|Saadanius]]''.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 35-20 Ma
|[[File:Proconsul NT.jpg|right|thumb|upright=0.9|''Proconsul'']]
[[Biology:Catarrhini|Catarrhini]] splits into 2 superfamilies, Old World monkeys (Cercopithecoidea) and [[Biology:Ape|ape]]s ([[Biology:Hominoidea|Hominoidea]]). Human trichromatic color vision had its genetic origins in this period. Catarrhines lost the vomeronasal organ (or possibly reduced it to vestigial status).

''[[Biology:Proconsul (mammal)|Proconsul]]'' was an early genus of catarrhine primates. They had a mixture of Old World monkey and [[Biology:Ape|ape]] characteristics. ''Proconsul'''s [[Biology:Monkey|monkey]]-like features include thin [[Biology:Tooth|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.

''[[Biology:Proconsul africanus|Proconsul africanus]]'' is a possible ancestor of both great and lesser apes, including humans.
|}

===Hominidae===
{| class="wikitable"
! scope="col" | Date
! scope="col" | Event
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 20-15 Ma
|[[Biology:Hominidae|Hominidae]] (great ape ancestors) speciate from the ancestors of the [[Biology:Hylobatidae|gibbon]] (lesser apes) between c. 20 to 16 Ma. They largely reduced their ancestral [[Biology:Snout|snout]] and lost the uricase enzyme (present in most organisms).<ref>{{Cite web|title=A new primate species at the root of the tree of extant hominoids|url=https://phys.org/news/2015-10-primate-species-root-tree-extant.html|website=phys.org|language=en|access-date=2020-05-29}}</ref>
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 16-12 Ma
|[[Biology:Homininae|Homininae]] ancestors speciate from the ancestors of the [[Biology:Orangutan|orangutan]] between c. 18 to 14 Ma.<ref>{{cite journal | last1 = Raauma | first1 = Ryan | last2 = Sternera | first2 = K | year = 2005 | title = Catarrhine primate divergence dates estimated from complete mitochondrial genomes | url = http://www.nyu.edu/gsas/dept/anthro/programs/csho/Content/Facultycvandinfo/Disotell/RAUMM&DISS.pdf | journal = Journal of Human Evolution | volume = 48 | issue = 3| pages = 237–57 | doi=10.1016/j.jhevol.2004.11.007 | pmid=15737392}}</ref>

''[[Biology:Pierolapithecus|Pierolapithecus catalaunicus]]'' is thought to be a common ancestor of [[Biology:Human|human]]s 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 [[Biology:Vertebral column|spine]], flexible wrists, and [[Biology:Scapula|shoulder blades]] that lie along its back.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 12 Ma
|''[[Biology:Danuvius guggenmosi|Danuvius guggenmosi]]'' is the first-discovered Late Miocene great ape with preserved [[Biology:Long bone|long bone]]s, and greatly elucidates the anatomical structure and locomotion of contemporary apes.<ref>{{Cite journal|last1=Böhme|first1=Madelaine|last2=Spassov|first2=Nikolai|last3=Fuss|first3=Jochen|last4=Tröscher|first4=Adrian|last5=Deane|first5=Andrew S.|last6=Prieto|first6=Jérôme|last7=Kirscher|first7=Uwe|last8=Lechner|first8=Thomas|last9=Begun|first9=David R.|s2cid=207888156|date=November 2019|title=A new Miocene ape and locomotion in the ancestor of great apes and humans|url=https://www.nature.com/articles/s41586-019-1731-0|journal=Nature|language=en|volume=575|issue=7783|pages=489–493|doi=10.1038/s41586-019-1731-0|pmid=31695194|bibcode=2019Natur.575..489B|issn=1476-4687}}</ref> It had adaptations for both hanging in trees (suspensory behavior) and walking on two legs (bipedalism)—whereas, among present-day hominids, humans are better adapted for the latter and the others for the former. ''Danuvius'' thus had a method of locomotion unlike any previously known ape called "extended limb clambering", walking directly along tree branches as well as using arms for suspending itself. The last common ancestor between humans and other apes possibly had a similar method of locomotion.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 12-8 Ma
|The clade currently represented by humans and the genus ''Pan'' ([[Biology:Chimpanzee|chimpanzee]]s and [[Biology:Bonobo|bonobo]]s) splits from the ancestors of the [[Biology:Gorilla|gorilla]]s between c. 12 to 8 Ma.<ref>{{Cite journal|last1=Popadin|first1=Konstantin|last2=Gunbin|first2=Konstantin|last3=Peshkin|first3=Leonid|last4=Annis|first4=Sofia|last5=Fleischmann|first5=Zoe|last6=Kraytsberg|first6=Genya|last7=Markuzon|first7=Natalya|last8=Ackermann|first8=Rebecca R.|last9=Khrapko|first9=Konstantin|date=2017-10-19|title=Mitochondrial pseudogenes suggest repeated inter-species hybridization in hominid evolution.|url=https://www.biorxiv.org/content/early/2017/10/19/134502|journal=bioRxiv|language=en|pages=134502|doi=10.1101/134502|doi-access=free|hdl=11427/36660|hdl-access=free}}</ref>
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 8-6 Ma
|[[File:SahelanthropusCL.png|right|thumb|upright=0.9|''Sahelanthropus tchadensis'']]

[[Biology:Hominini|Hominini]]: The [[Biology:Chimpanzee–human last common ancestor|latest common ancestor]] of [[Biology:Homo|humans]] and [[Biology:Pan (genus)|chimpanzees]]
is estimated to have lived between roughly 10 to 5 million years ago. Both chimpanzees and humans have a [[Medicine:Larynx|larynx]] that repositions during the first two years of life to a spot between the [[Biology:Pharynx|pharynx]] and the lungs, indicating that the common ancestors have this feature, a precondition for vocalized speech in humans.
Speciation may have begun shortly after 10 Ma, but late admixture between the lineages may have taken place until after 5 Ma. Candidates of [[Biology:Hominina|Hominina]] or [[Biology:Homininae|Homininae]] species which lived in this time period include
''[[Biology:Ouranopithecus|Ouranopithecus]]'' (c. 8 Ma),
''[[Biology:Graecopithecus|Graecopithecus]]'' (c. 7 Ma),
''Sahelanthropus tchadensis'' (c. 7 Ma),
''Orrorin tugenensis'' (c. 6 Ma).
[[File:Ardi.jpg|thumb|upright=0.9|''Ardipithecus'']]''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<ref name="NatGeo-News">{{cite web|url= http://news.nationalgeographic.com/news/2001/07/0712_ethiopianbones.html|archive-url= https://web.archive.org/web/20010715123225/http://news.nationalgeographic.com/news/2001/07/0712_ethiopianbones.html|url-status= dead|archive-date= July 15, 2001|title=Fossils From Ethiopia May Be Earliest Human Ancestor|first=David|last=Perlman|publisher=National Geographic News|date=July 12, 2001|quote=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.}}</ref> during the early Pliocene, and ''A. kadabba'', dated to approximately 5.6 million years ago<ref name="AndThePaleobiology">{{cite journal|last=White|first=Tim D.|author2=Asfaw, Berhane |author3=Beyene, Yonas |author4=Haile-Selassie, Yohannes |author5=Lovejoy, C. Owen |author6=Suwa, Gen |author7= WoldeGabriel, Giday |s2cid=20189444|year=2009 |title=''Ardipithecus ramidus'' and the Paleobiology of Early Hominids.|journal=Science |volume=326|issue=5949|pages=75–86|doi=10.1126/science.1175802|pmid=19810190|bibcode = 2009Sci...326...75W }}</ref> (late Miocene). ''A. ramidus'' had a small brain, measuring between 300 and 350&nbsp;cm<sup>3</sup>. This is about the same size as the modern bonobo and female chimpanzee brain; it is somewhat smaller than the brain of australopithecines like [[Biology:Lucy (Australopithecus)|Lucy]] (400 to 550&nbsp;cm<sup>3</sup>) 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.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 4-3.5 Ma
|[[File:Bienvenida al Museo de Lucy.jpg|thumb|upright|Reconstruction of "Lucy"]]
A member of the ''[[Biology:Australopithecus afarensis|Australopithecus afarensis]]'' left human-like footprints on volcanic ash in Laetoli, 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 ''[[Biology:Australopithecus|Australopithecus]]'' and the genus ''[[Biology:Homo|Homo]]''. Compared to the modern and extinct great [[Biology:Ape|ape]]s, ''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&nbsp;cm³) and a prognathic (anterior-projecting) face.

Australopithecines have been found in savannah environments; they probably developed their diet to include scavenged meat.  Analyses of ''[[Biology:Australopithecus africanus|Australopithecus africanus]]'' lower vertebrae suggests that these bones changed in females to support bipedalism even during pregnancy.
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 3.5–3.0 Ma
| ''Kenyanthropus platyops'', a possible ancestor of ''Homo'', emerges from the ''Australopithecus''. Stone tools are deliberately constructed.<ref>{{Cite journal | doi=10.1038/nature14464|pmid = 25993961| title=3.3-million-year-old stone tools from Lomekwi 3, West Turkana, Kenya| journal=Nature| volume=521| issue=7552| pages=310–15| year=2015| last1=Harmand| first1=Sonia| last2=Lewis| first2=Jason E.| last3=Feibel| first3=Craig S.| last4=Lepre| first4=Christopher J.| last5=Prat| first5=Sandrine| last6=Lenoble| first6=Arnaud| last7=Boës| first7=Xavier| last8=Quinn| first8=Rhonda L.| last9=Brenet| first9=Michel| last10=Arroyo| first10=Adrian| last11=Taylor| first11=Nicholas| last12=Clément| first12=Sophie| last13=Daver| first13=Guillaume| last14=Brugal| first14=Jean-Philip| last15=Leakey| first15=Louise| last16=Mortlock| first16=Richard A.| last17=Wright| first17=James D.| last18=Lokorodi| first18=Sammy| last19=Kirwa| first19=Christopher| last20=Kent| first20=Dennis V.| last21=Roche| first21=Hélène|s2cid = 1207285|bibcode = 2015Natur.521..310H}}</ref>
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 3 Ma
| The bipedal [[Biology:Australopithecus|australopithecines]] (a genus of the subtribe [[Biology:Hominina|Hominina]]) evolve in the savannas of [[Earth:Africa|Africa]] being hunted by ''[[Biology:Megantereon|Megantereon]]''. Loss of body hair occurs from 3 to 2 Ma, in parallel with the development of full [[Bipedalism|bipedalism]] and slight enlargement of the brain.<ref>{{Cite journal|last1=Ruxton|first1=Graeme D.|last2=Wilkinson|first2=David M.|date=2011-12-27|title=Avoidance of overheating and selection for both hair loss and bipedality in hominins|journal=Proceedings of the National Academy of Sciences|language=en|volume=108|issue=52|pages=20965–20969|doi=10.1073/pnas.1113915108|issn=0027-8424|pmid=22160694|pmc=3248486|bibcode=2011PNAS..10820965R|doi-access=free}}</ref>
|}

===Homo===
{| class="wikitable"
! scope="col" | Date
! scope="col" | Event
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 2.5–2.0 Ma
|
Early ''[[Biology:Homo|Homo]]'' appears in East Africa,   speciating from [[Biology:Australopithecine|australopithecine]] ancestors.
The Lower Paleolithic is defined by the beginning of use of [[Engineering:Stone tool|stone tool]]s.
''[[Biology:Australopithecus garhi|Australopithecus garhi]]'' was using stone tools at about 2.5 Ma.
''[[Biology:Homo habilis|Homo habilis]]'' is the oldest species given the designation ''Homo'', by Leakey et al in 1964.
''H. habilis'' is  intermediate between ''[[Biology:Australopithecus afarensis|Australopithecus afarensis]]'' and ''[[Biology:Homo erectus|H. erectus]]'', and there have been suggestions to re-classify it within genus ''Australopithecus'', as ''Australopithecus habilis''.

Stone tools found at the Shangchen site in China and dated to 2.12 million years ago are considered the earliest known evidence of hominins outside Africa, surpassing Dmanisi in Georgia by 300,000 years.<ref>{{Cite journal | doi=10.1038/s41586-018-0299-4|pmid = 29995848| title=Hominin occupation of the Chinese Loess Plateau since about 2.1 million years ago| journal=Nature| volume=559| issue=7715| pages=608–12| year=2018| last1=Zhu| first1=Zhaoyu| last2=Dennell| first2=Robin| last3=Huang| first3=Weiwen| last4=Wu| first4=Yi| last5=Qiu| first5=Shifan| last6=Yang| first6=Shixia| last7=Rao| first7=Zhiguo| last8=Hou| first8=Yamei| last9=Xie| first9=Jiubing| last10=Han| first10=Jiangwei| last11=Ouyang| first11=Tingping|s2cid = 49670311|bibcode = 2018Natur.559..608Z}}</ref>
{{Further|Biology:Homo naledi|Biology:Homo rudolfensis}}
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 1.9–0.8 Ma
|[[File:Homo.erectus.adult.female.smithsonian.timevanson.flickr.jpg|thumb|upright|Reconstruction of a female ''H. erectus'']]
''[[Biology:Homo erectus|Homo erectus]]''  derives from early ''Homo'' or late ''Australopithecus''.

''[[Biology:Homo habilis|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 ''H. erectus'' for almost half a million years (until about 1.5 Ma).
From its earliest appearance at about 1.9 Ma, ''H. erectus'' is distributed in East Africa and Southwest Asia (''Homo georgicus''). 
''H. erectus'' is  the first known species to develop [[Biology:Control of fire by early humans|control of fire]], by about 1.5 Ma.

''H. erectus'' later migrates throughout Eurasia, reaching Southeast Asia by 0.7 Ma.
It is described in a number of subspecies.<ref>NOVA: Becoming Human Part 2 http://video.pbs.org/video/1319997127/</ref> Early humans were social and initially scavenged, before becoming active hunters. The need to communicate and hunt prey efficiently in a new, fluctuating environment (where the locations of resources need to be memorized and told) may have driven the expansion of the brain from 2 to 0.8 Ma.

Evolution of dark skin at about 1.2 Ma.<ref name=jabl04>{{cite journal |last1=Jablonski |first1=Nina G. |s2cid=53481281 |title=The Evolution of Human Skin and Skin Color |journal=Annual Review of Anthropology |date=October 2004 |volume=33 |issue=1 |pages=585–623 |doi=10.1146/annurev.anthro.33.070203.143955 }}</ref>

''[[Biology:Homo antecessor|Homo antecessor]]'' may be a common ancestor of humans and Neanderthals.<ref>{{cite journal |last1=Bermudez de Castro |first1=J. M. |title=A Hominid from the Lower Pleistocene of Atapuerca, Spain: Possible Ancestor to Neandertals and Modern Humans |journal=Science |date=30 May 1997 |volume=276 |issue=5317 |pages=1392–1395 |doi=10.1126/science.276.5317.1392 |pmid=9162001 }}</ref><ref>{{cite journal |last1=Green |first1=Richard E. |last2=Krause |first2=Johannes |last3=Ptak |first3=Susan E. |last4=Briggs |first4=Adrian W. |last5=Ronan |first5=Michael T. |last6=Simons |first6=Jan F. |last7=Du |first7=Lei |last8=Egholm |first8=Michael |last9=Rothberg |first9=Jonathan M. |last10=Paunovic |first10=Maja |last11=Pääbo |first11=Svante |s2cid=4320907 |title=Analysis of one million base pairs of Neanderthal DNA |journal=Nature |date=November 2006 |volume=444 |issue=7117 |pages=330–336 |doi=10.1038/nature05336 |pmid=17108958 |bibcode=2006Natur.444..330G |doi-access=free }}</ref> At present estimate, humans have approximately 20,000–25,000 [[Biology:Gene|gene]]s and share 99% of their [[Biology:DNA|DNA]] with the now extinct [[Biology:Neanderthal|Neanderthal]]<ref>"<cite>Rubin also said analysis so far suggests human and Neanderthal DNA are some 99.5 percent to nearly 99.9 percent identical.</cite>" [http://www.cnn.com/2006/TECH/science/11/15/neanderthal.ap/index.html Neanderthal bone gives DNA clues] (URL accessed on November 16, 2006)</ref> and 95–99% of their [[Biology:DNA|DNA]] with their closest living evolutionary relative, the [[Biology:Chimpanzee|chimpanzee]]s.<ref>"<cite>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.</cite>" {{cite journal|doi=10.1073/pnas.172510699|title=Divergence between samples of chimpanzee and human DNA sequences is 5%, counting indels|author=Britten, R.J. |journal = PNAS |year=2002|volume=99|pages=13633–35|pmid=12368483|issue=21|pmc=129726|bibcode = 2002PNAS...9913633B |doi-access=free}}</ref><ref>"<cite>...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.</cite>" {{Cite journal|title=What makes us human?|author=Pollard, K.S.|s2cid=38866839|journal = Scientific American|year=2009|volume=300–5|pages=44–49|doi=10.1038/scientificamerican0509-44|pmid=19438048|issue=5|bibcode=2009SciAm.300e..44P}}</ref>  The human variant of the [[Biology:FOXP2|FOXP2]] gene (linked to the control of speech) has been found to be identical in Neanderthals.<ref name="pmid17949978">{{cite journal | title = The derived FOXP2 variant of modern humans was shared with Neandertals | journal = Curr. Biol. | volume = 17 | issue = 21 | pages = 1908–12 |date=November 2007 | pmid = 17949978 | doi = 10.1016/j.cub.2007.10.008 |vauthors=Krause J, Lalueza-Fox C, Orlando L, Enard W, Green RE, Burbano HA, Hublin JJ, Hänni C, Fortea J, de la Rasilla M, Bertranpetit J, Rosas A, Pääbo S | s2cid = 9518208 |  hdl = 11858/00-001M-0000-000F-FED3-1 | hdl-access = free }}
*{{cite news |author=Nicholas Wade |date=October 19, 2007 |title=Neanderthals Had Important Speech Gene, DNA Evidence Shows |newspaper=The New York Times |url=https://www.nytimes.com/2007/10/19/science/19speech-web.html?ref=world |url-access=subscription}}</ref>
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 0.8–0.3 Ma
|[[File:Homo heidelbergensis - forensic facial reconstruction-crop.png|thumb|upright|Reconstruction of ''Homo heidelbergensis'']]
Divergence of [[Biology:Neanderthal|Neanderthal]] and [[History:Denisovan|Denisovan]] lineages from a common ancestor.<ref>{{cite journal |last1=Stein |first1=Richard A. |title=Copy Number Analysis Starts to Add Up |journal=Genetic Engineering & Biotechnology News |date=October 2015 |volume=35 |issue=17 |pages=20, 22–23 |doi=10.1089/gen.35.17.09 }}</ref>
''[[Biology:Homo heidelbergensis|Homo heidelbergensis]]'' (in Africa also known as ''[[Biology:Homo rhodesiensis|Homo rhodesiensis]]'') had long been thought to be a likely candidate for the last common ancestor of the Neanderthal and modern human lineages.
However, genetic evidence from the [[Biology:Sima de los Huesos|Sima de los Huesos]] fossils published in 2016 seems to suggest that ''H. heidelbergensis'' in its entirety should be included in the Neanderthal lineage, as "pre-Neanderthal" or "early Neanderthal", while the divergence time between the Neanderthal and modern lineages has been pushed back to before the emergence of ''H. heidelbergensis'', to about 600,000 to 800,000 years ago, the approximate age of ''[[Biology:Homo antecessor|Homo antecessor]]''.<ref>{{Cite journal | doi=10.1038/nature17405| pmid=26976447| title=Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins| journal=Nature| volume=531| issue=7595| pages=504–07| date=March 2016| last1=Meyer| first1=Matthias| last2=Arsuaga| first2=Juan-Luis| last3=De Filippo| first3=Cesare| last4=Nagel| first4=Sarah| last5=Aximu-Petri| first5=Ayinuer| last6=Nickel| first6=Birgit| last7=Martínez| first7=Ignacio| last8=Gracia| first8=Ana| last9=De Castro| first9=José María Bermúdez| last10=Carbonell| first10=Eudald| last11=Viola| first11=Bence| last12=Kelso| first12=Janet| last13=Prüfer| first13=Kay| last14=Pääbo| first14=Svante| s2cid=4467094| bibcode=2016Natur.531..504M}}</ref><ref>{{Cite journal | doi=10.1038/531286a| pmid=26983523| title=Oldest ancient-human DNA details dawn of Neanderthals| journal=Nature| volume=531| issue=7594| pages=296–86| year=2016| last1=Callaway| first1=Ewen| s2cid=4459329| bibcode=2016Natur.531..296C| doi-access=free}}</ref> Brain expansion (enlargement) between 0.8 and 0.2 Ma may have occurred due to the extinction of most African [[Biology:Megafauna|megafauna]] (which made humans feed from smaller prey and plants, which required greater intelligence due to greater speed of the former and uncertainty about whether the latter were poisonous or not), extreme climate variability after [[Earth:Mid-Pleistocene Transition|Mid-Pleistocene Transition]] (which intensified the situation, and resulted in frequent migrations), and in general selection for more social life (and intelligence) for greater chance of survival, reproductivity, and care for mothers. Solidified footprints dated to about 350 ka and associated with ''H. heidelbergensis'' were found in  southern Italy in 2003.<ref>{{cite journal|last1=Mietto|first1=Paolo|last2=Avanzini|first2=Marco|last3=Rolandi|first3=Giuseppe|s2cid=2396763|title=Palaeontology: Human footprints in Pleistocene volcanic ash|journal=Nature|volume=422|issue=6928|year=2003|page=133|doi=10.1038/422133a|pmid=12634773|bibcode=2003Natur.422..133M|doi-access=free}}</ref> 

''H. sapiens'' lost the brow ridges from their hominid ancestors as well as the snout completely, though their noses evolve to be protruding (possibly from the time of ''H. erectus''). By 200 ka, humans had stopped their brain expansion.
|-
|}

===Homo sapiens===
{{see|Homo sapiens|Neanderthal|Interbreeding between archaic and modern humans|Recent human evolution|Human genetic variation}}
{| class="wikitable"
! scope="col" | Date
! scope="col" | Event
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 300–130 ka
|[[File:Homo sapiens sapiens (Fundort Jebel Irhoud Marokko).jpg|thumb|upright|Reconstruction of early ''Homo sapiens'' from Jebel Irhoud, Morocco {{c.|315 000 years BP}}]]
Neanderthals and Denisovans emerge from the northern ''[[Biology:Homo heidelbergensis|Homo heidelbergensis]]'' lineage around 500-450 ka while [[Biology:Human|sapients]] emerge  from the southern lineage around 350-300 ka.<ref>Timmermann, A., Yun, KS., Raia, P. et al. Climate effects on archaic human habitats and species successions. Nature '''604''', 495–501 (2022). https://doi.org/10.1038/s41586-022-04600-9</ref>

Fossils attributed to ''H. sapiens,'' along with stone tools, dated to approximately 300,000 years ago, found at [[Biology:Jebel Irhoud|Jebel Irhoud]], Morocco<ref name="NAT-20170607a">{{cite journal |last1=Callaway |first1=Ewen |title=Oldest Homo sapiens fossil claim rewrites our species' history |journal=Nature |date=7 June 2017 |doi=10.1038/nature.2017.22114 }}</ref> yield the earliest fossil evidence for anatomically modern ''Homo sapiens''.
Modern human presence in [[Place:East Africa|East Africa]] (Gademotta), at 276 kya.<ref name="Tryon Faith 2013">{{cite journal | last1 = Tryon | first1 = Christopher A. | last2 = Faith | first2 = Tyler | year = 2013 | title = Variability in the Middle Stone Age of Eastern Africa | doi = 10.1086/673752 | journal = Current Anthropology | volume = 54 | issue = 8| pages = 234–54 | s2cid = 14124486 | url = http://espace.library.uq.edu.au/view/UQ:321153/UQ321153_OA.pdf }}</ref> In July 2019, anthropologists reported the discovery of 210,000 year old remains of a ''H. sapiens'' in Apidima Cave, Peloponnese, Greece.<ref name="NYT-20190710">{{cite news |last=Zimmer |first=Carl |title=A Skull Bone Discovered in Greece May Alter the Story of Human Prehistory - The bone, found in a cave, is the oldest modern human fossil ever discovered in Europe. It hints that humans began leaving Africa far earlier than once thought. |url=https://www.nytimes.com/2019/07/10/science/skull-neanderthal-human-europe-greece.html |date=10 July 2019 |work={{wipe|The New York Times}} |access-date=11 July 2019 }}</ref><ref name="PHYS-20190710">{{cite news |author=Staff |title='Oldest remains' outside Africa reset human migration clock |url=https://phys.org/news/2019-07-oldest-africa-reset-human-migration.html |date=10 July 2019 |work=Phys.org |access-date=10 July 2019 }}</ref><ref name="NAT-20190710">{{cite journal |last=Harvati |first=Katerina |s2cid=195873640 |display-authors=et al. |title=Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia |date=10 July 2019 |journal=Nature |volume=571 |issue=7766 |pages=500–504 |doi=10.1038/s41586-019-1376-z |pmid=31292546 |url=https://zenodo.org/record/6646855 }}</ref>

[[Biology:Y-chromosomal Adam|Patrilineal]] and [[Biology:Mitochondrial Eve|matrilineal]] most recent common ancestors (MRCAs) of living humans roughly between 200 and 100 kya<ref>{{cite journal |last1=Heinz |first1=Tanja |last2=Pala |first2=Maria |last3=Gómez-Carballa |first3=Alberto |last4=Richards |first4=Martin B. |last5=Salas |first5=Antonio |title=Updating the African human mitochondrial DNA tree: Relevance to forensic and population genetics |journal=Forensic Science International: Genetics |date=March 2017 |volume=27 |pages=156–159 |doi=10.1016/j.fsigen.2016.12.016 |pmid=28086175 }}</ref><ref>{{cite journal |last1=Poznik |first1=G. D. |last2=Henn |first2=B. M. |last3=Yee |first3=M.-C. |last4=Sliwerska |first4=E. |last5=Euskirchen |first5=G. M. |last6=Lin |first6=A. A. |last7=Snyder |first7=M. |last8=Quintana-Murci |first8=L. |last9=Kidd |first9=J. M. |last10=Underhill |first10=P. A. |last11=Bustamante |first11=C. D. |title=Sequencing Y Chromosomes Resolves Discrepancy in Time to Common Ancestor of Males Versus Females |journal=Science |date=1 August 2013 |volume=341 |issue=6145 |pages=562–565 |doi=10.1126/science.1237619 |pmid=23908239 |pmc=4032117 |bibcode=2013Sci...341..562P }}</ref>
with some estimates on the patrilineal MRCA somewhat higher, ranging up to 250 to 500 kya.<ref>{{cite journal |last1=Karmin |first1=Monika |last2=Saag |first2=Lauri |last3=Vicente |first3=Mário |last4=Sayres |first4=Melissa A. Wilson |last5=Järve |first5=Mari |last6=Talas |first6=Ulvi Gerst |last7=Rootsi |first7=Siiri |last8=Ilumäe |first8=Anne-Mai |last9=Mägi |first9=Reedik |last10=Mitt |first10=Mario |last11=Pagani |first11=Luca |last12=Puurand |first12=Tarmo |last13=Faltyskova |first13=Zuzana |last14=Clemente |first14=Florian |last15=Cardona |first15=Alexia |last16=Metspalu |first16=Ene |last17=Sahakyan |first17=Hovhannes |last18=Yunusbayev |first18=Bayazit |last19=Hudjashov |first19=Georgi |last20=DeGiorgio |first20=Michael |last21=Loogväli |first21=Eva-Liis |last22=Eichstaedt |first22=Christina |last23=Eelmets |first23=Mikk |last24=Chaubey |first24=Gyaneshwer |last25=Tambets |first25=Kristiina |last26=Litvinov |first26=Sergei |last27=Mormina |first27=Maru |last28=Xue |first28=Yali |last29=Ayub |first29=Qasim |last30=Zoraqi |first30=Grigor |last31=Korneliussen |first31=Thorfinn Sand |last32=Akhatova |first32=Farida |last33=Lachance |first33=Joseph |last34=Tishkoff |first34=Sarah |last35=Momynaliev |first35=Kuvat |last36=Ricaut |first36=François-Xavier |last37=Kusuma |first37=Pradiptajati |last38=Razafindrazaka |first38=Harilanto |last39=Pierron |first39=Denis |last40=Cox |first40=Murray P. |last41=Sultana |first41=Gazi Nurun Nahar |last42=Willerslev |first42=Rane |last43=Muller |first43=Craig |last44=Westaway |first44=Michael |last45=Lambert |first45=David |last46=Skaro |first46=Vedrana |last47=Kovačevic´ |first47=Lejla |last48=Turdikulova |first48=Shahlo |last49=Dalimova |first49=Dilbar |last50=Khusainova |first50=Rita |last51=Trofimova |first51=Natalya |last52=Akhmetova |first52=Vita |last53=Khidiyatova |first53=Irina |last54=Lichman |first54=Daria V. |last55=Isakova |first55=Jainagul |last56=Pocheshkhova |first56=Elvira |last57=Sabitov |first57=Zhaxylyk |last58=Barashkov |first58=Nikolay A. |last59=Nymadawa |first59=Pagbajabyn |last60=Mihailov |first60=Evelin |last61=Seng |first61=Joseph Wee Tien |last62=Evseeva |first62=Irina |last63=Migliano |first63=Andrea Bamberg |last64=Abdullah |first64=Syafiq |last65=Andriadze |first65=George |last66=Primorac |first66=Dragan |last67=Atramentova |first67=Lubov |last68=Utevska |first68=Olga |last69=Yepiskoposyan |first69=Levon |last70=Marjanovic´ |first70=Damir |last71=Kushniarevich |first71=Alena |last72=Behar |first72=Doron M. |last73=Gilissen |first73=Christian |last74=Vissers |first74=Lisenka |last75=Veltman |first75=Joris A. |last76=Balanovska |first76=Elena |last77=Derenko |first77=Miroslava |last78=Malyarchuk |first78=Boris |last79=Metspalu |first79=Andres |last80=Fedorova |first80=Sardana |last81=Eriksson |first81=Anders |last82=Manica |first82=Andrea |last83=Mendez |first83=Fernando L. |last84=Karafet |first84=Tatiana M. |last85=Veeramah |first85=Krishna R. |last86=Bradman |first86=Neil |last87=Hammer |first87=Michael F. |last88=Osipova |first88=Ludmila P. |last89=Balanovsky |first89=Oleg |last90=Khusnutdinova |first90=Elza K. |last91=Johnsen |first91=Knut |last92=Remm |first92=Maido |last93=Thomas |first93=Mark G. |last94=Tyler-Smith |first94=Chris |last95=Underhill |first95=Peter A. |last96=Willerslev |first96=Eske |last97=Nielsen |first97=Rasmus |last98=Metspalu |first98=Mait |last99=Villems |first99=Richard |display-authors=29 |last100=Kivisild |first100=Toomas |title=A recent bottleneck of Y chromosome diversity coincides with a global change in culture |journal=Genome Research |date=April 2015 |volume=25 |issue=4 |pages=459–466 |doi=10.1101/gr.186684.114 |pmid=25770088 |pmc=4381518 }}</ref>

160,000 years ago, ''[[Biology:Homo sapiens idaltu|Homo sapiens idaltu]]'' in the Awash River Valley (near present-day [[Earth:Bouri Formation#Herto|Herto village]], Ethiopia) practiced excarnation.<ref>{{cite journal |last1=Clark |first1=J. Desmond |last2=Beyene |first2=Yonas |last3=WoldeGabriel |first3=Giday |last4=Hart |first4=William K. |last5=Renne |first5=Paul R. |last6=Gilbert |first6=Henry |last7=Defleur |first7=Alban |last8=Suwa |first8=Gen |last9=Katoh |first9=Shigehiro |last10=Ludwig |first10=Kenneth R. |last11=Boisserie |first11=Jean-Renaud |last12=Asfaw |first12=Berhane |last13=White |first13=Tim D. |s2cid=4312418 |title=Stratigraphic, chronological and behavioural contexts of Pleistocene Homo sapiens from Middle Awash, Ethiopia |journal=Nature |date=June 2003 |volume=423 |issue=6941 |pages=747–752 |doi=10.1038/nature01670 |pmid=12802333 |bibcode=2003Natur.423..747C }}</ref>

|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 130–80 ka
| [[Earth:Marine Isotope Stage 5|Marine Isotope Stage 5]] ([[Earth:Eemian|Eemian]]).
Modern human presence in [[Place:Southern Africa|Southern Africa]] and [[Place:West Africa|West Africa]].<ref>{{cite encyclopedia |doi=10.1093/acrefore/9780190277734.013.137 |chapter=The Stone Age Archaeology of West Africa |title=Oxford Research Encyclopedia of African History |year=2017 |last1=Scerri |first1=Eleanor |isbn=9780190277734 |chapter-url=https://ora.ox.ac.uk/objects/uuid:1442d1c1-7f76-454d-97db-8fd52a4c6c1e }}</ref>
Appearance of mitochondrial haplogroup (mt-haplogroup) [[Biology:Haplogroup L2 (mtDNA)|L2]].
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 80–50 ka
| [[Physics:Marine isotope stage|MIS 4]], beginning of the [[Earth:Upper Paleolithic|Upper Paleolithic]].
Early evidence for [[Philosophy:Behavioral modernity|behavioral modernity]].<ref>Henshilwood, C.S. and B. Dubreuil 2009. Reading the artifacts: gleaning language skills from the Middle Stone Age in southern Africa. In R. Botha and C. Knight (eds), The Cradle of Language. Oxford: Oxford University Press, pp. 41-61.</ref>
Appearance of mt-haplogroups [[Biology:Haplogroup M (mtDNA)|M]] and [[Biology:Haplogroup N (mtDNA)|N]]. [[Earth:Southern Dispersal|Southern Dispersal]]  migration out of Africa, Proto-Australoid [[Earth:Southern Dispersal|peopling of Oceania]].<ref name = "pmid12594511">{{cite journal|vauthors=Bowler JM, Johnston H, Olley JM, Prescott JR, Roberts RG, Shawcross W, Spooner NA |s2cid = 4365526 |title = New ages for human occupation and climatic change at Lake Mungo, Australia | journal = Nature  |volume = 421 |issue = 6925 | year = 2003 | pages = 837–40 | pmid = 12594511 | doi = 10.1038/nature01383  |bibcode = 2003Natur.421..837B}}</ref> Archaic admixture from Neanderthals in Eurasia,<ref name = "green">{{Cite journal | author = Richard E. Green 
 | title = A Draft Sequence of the Neandertal Genome | journal = Science
 | volume = 328 | issue = 5979 | pages = 710–22 | doi = 10.1126/science.1188021 | pmid = 20448178 | year = 2010|bibcode = 2010Sci...328..710G | last2 = Krause | first2 = J. | last3 = Briggs | first3 = A.W. | last4 = Maricic | first4 = T. | last5 = Stenzel | first5 = U. | last6 = Kircher | first6 = M. | last7 = Patterson | first7 = N. | last8 = Li | first8 = H. | last9 = Zhai | first9 = W. | last10 = Fritz | first10 = M.H.Y. | last11 = Hansen | first11 = N.F. | last12 = Durand | first12 = E.Y. | last13 = Malaspinas | first13 = A.S. | last14 = Jensen | first14 = J.D. | last15 = Marques-Bonet | first15 = T. | last16 = Alkan | first16 = C. | last17 = Prufer | first17 = K. | last18 = Meyer | first18 = M. | last19 = Burbano | first19 = H.A. | last20 = Good | first20 = J.M. | last21 = Schultz | first21 = R. | last22 = Aximu-Petri | first22 = A. | last23 = Butthof | first23 = A. | last24 = Hober | first24 = B. | last25 = Hoffner | first25 = B. | last26 = Siegemund | first26 = M. | last27 = Weihmann | first27 = A. | last28 = Nusbaum | first28 = C. | last29 = Lander | first29 = E.S. | last30 = Russ | first30 = C. | display-authors = 8 | pmc=5100745}}</ref><ref name = "BBC">{{cite news
  | last = Rincon | first = Paul | title = Neanderthal genes 'survive in us'
  | work = BBC News | publisher = BBC
  | date = 2010-05-06 | url = http://news.bbc.co.uk/2/hi/science/nature/8660940.stm
  | access-date = 2010-05-07}}</ref> from  Denisovans in Oceania with trace amounts in Eastern Eurasia,<ref name=sank16-comb>{{cite journal |last1=Sankararaman |first1=Sriram |last2=Mallick |first2=Swapan |last3=Patterson |first3=Nick |last4=Reich |first4=David |title=The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans |journal=Current Biology |date=2016 |volume=26 |issue=9 |pages=1241–1247 |doi=10.1016/j.cub.2016.03.037|pmid=27032491 |pmc=4864120 }}</ref> and from an unspecified African lineage of archaic humans in Sub-Saharan Africa as well as an interbred species of Neanderthals and Denisovans in Asia and Oceania.<ref>{{cite journal |last1=Callaway |first1=Ewen |s2cid=87081207 |title=Hunter-gatherer genomes a trove of genetic diversity |journal=Nature |date=26 July 2012 |doi=10.1038/nature.2012.11076 }}</ref><ref>{{cite journal |last1=Lachance |first1=Joseph |last2=Vernot |first2=Benjamin |last3=Elbers |first3=Clara C. |last4=Ferwerda |first4=Bart |last5=Froment |first5=Alain |last6=Bodo |first6=Jean-Marie |last7=Lema |first7=Godfrey |last8=Fu |first8=Wenqing |last9=Nyambo |first9=Thomas B. |last10=Rebbeck |first10=Timothy R. |last11=Zhang |first11=Kun |last12=Akey |first12=Joshua M. |last13=Tishkoff |first13=Sarah A. |title=Evolutionary History and Adaptation from High-Coverage Whole-Genome Sequences of Diverse African Hunter-Gatherers |journal=Cell |date=August 2012 |volume=150 |issue=3 |pages=457–469 |doi=10.1016/j.cell.2012.07.009 |pmid=22840920 |pmc=3426505}}</ref><ref>{{cite journal |last1=Xu |first1=Duo |last2=Pavlidis |first2=Pavlos |last3=Taskent |first3=Recep Ozgur |last4=Alachiotis |first4=Nikolaos |last5=Flanagan |first5=Colin |last6=DeGiorgio |first6=Michael |last7=Blekhman |first7=Ran |last8=Ruhl |first8=Stefan |last9=Gokcumen |first9=Omer |title=Archaic Hominin Introgression in Africa Contributes to Functional Salivary MUC7 Genetic Variation |journal=Molecular Biology and Evolution |date=October 2017 |volume=34 |issue=10 |pages=2704–2715 |doi=10.1093/molbev/msx206 |pmid=28957509 |pmc=5850612 }}</ref><ref>{{cite journal |last1=Mondal |first1=Mayukh |last2=Bertranpetit |first2=Jaume |last3=Lao |first3=Oscar |title=Approximate Bayesian computation with deep learning supports a third archaic introgression in Asia and Oceania |journal=Nature Communications |date=16 January 2019 |volume=10 |issue=1 |pages=246 |doi=10.1038/s41467-018-08089-7 |pmid=30651539 |pmc=6335398 |bibcode=2019NatCo..10..246M }}</ref>
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | 50–25 ka
|
[[File:Homo sapiens sapiens, Oase, Rumänien (Daniela Hitzemann).jpg|thumb|upright|Reconstruction of Oase 2 (c. 40 ka)]]
[[Philosophy:Behavioral modernity#Great leap forward|Behavioral modernity]] develops by this time or earlier, according to the "great leap forward" theory.<ref name="Klein 1995">{{cite journal|last1=Klein|first1=Richard|s2cid=10402296|title=Anatomy, behavior, and modern human origins|journal=Journal of World Prehistory|date=1995|volume=9|issue=2|pages=167–98|doi=10.1007/bf02221838}}</ref>
Extinction of ''[[Biology:Homo floresiensis|Homo floresiensis]]''.<ref>{{cite journal|last1=Sutikna|first1=Thomas|last2=Tocheri|first2=Matthew W.|last3=Morwood|first3=Michael J.|last4=Saptomo|first4=E. Wahyu|last5=Jatmiko|last6=Awe|first6=Rokus Due|last7=Wasisto|first7=Sri|last8=Westaway|first8=Kira E.|last9=Aubert|first9=Maxime|last10=Li|first10=Bo|last11=Zhao|first11=Jian-xin|last12=Storey|first12=Michael|last13=Alloway|first13=Brent V.|last14=Morley|first14=Mike W.|last15=Meijer|first15=Hanneke J.M.|last16=van den Bergh|first16=Gerrit D.|last17=Grün|first17=Rainer|last18=Dosseto|first18=Anthony|author19-link=Adam Brumm|last19=Brumm|first19=Adam|last20=Jungers|first20=William L.|last21=Roberts|first21=Richard G.|s2cid=4469009|title=Revised stratigraphy and chronology for Homo floresiensis at Liang Bua in Indonesia|journal=Nature|date=30 March 2016|volume=532|issue=7599|pages=366–69|doi=10.1038/nature17179|pmid=27027286|bibcode=2016Natur.532..366S|url=https://ro.uow.edu.au/smhpapers/3778}}</ref>
M168 mutation (carried by all non-African males).
Appearance of mt-haplogroups [[Biology:Haplogroup U (mtDNA)|U]] and [[Biology:Haplogroup K (mtDNA)|K]].
Peopling of Europe, peopling of the North Asian [[Earth:Mammoth steppe|Mammoth steppe]]. Paleolithic art.
[[Biology:Neanderthal extinction|Extinction of Neanderthals]] and other archaic human variants  (with possible survival of hybrid populations in Asia and Africa.)
Appearance of Y-Haplogroup R2; mt-haplogroups [[Biology:Haplogroup J (mtDNA)|J]] and [[Biology:Haplogroup X (mtDNA)|X]].
|-
| style="vertical-align: top; text-align: right; white-space: nowrap;" | after 25 ka
|[[File:Homo sapiens - Neolithic - reconstruction - MUSE.jpg|thumb|upright|Reconstruction of a Neolithic farmer from Europe, Science Museum in Trento]]
[[Earth:Last Glacial Maximum|Last Glacial Maximum]]; Epipaleolithic / Mesolithic / [[Earth:Holocene|Holocene]].
Peopling of the Americas.
Appearance of: Y-Haplogroup R1a; mt-haplogroups [[Biology:Haplogroup V (mtDNA)|V]] and [[Biology:Haplogroup T (mtDNA)|T]].
Various [[Biology:Recent human evolution|recent divergence]] associated with environmental pressures,
e.g.  light skin in Europeans and East Asians (KITLG, [[Biology:Agouti signalling peptide|ASIP]]), after 30 ka;<ref name=Belezal2012>{{cite journal |url= |title=The timing of pigmentation lightening in Europeans |journal=Molecular Biology and Evolution |year=2012 |doi=10.1093/molbev/mss207 |pmid=22923467 |pmc=3525146 |volume=30 |issue=1 |pages=24–35|last2=Santos |first2=A.M. |last3=McEvoy |first3=B. |last4=Alves |first4=I. |last5=Martinho |first5=C. |last6=Cameron |first6=E. |last7=Shriver |first7=M.D. |last8=Parra |first8=E.J. |last9=Rocha |first9=J.|last1=Belezal|first1=Sandra}}</ref>
Inuit adaptation to high-fat diet and cold climate, 20 ka.<ref>{{cite journal |last1=Fumagalli |first1=M. |last2=Moltke |first2=I. |last3=Grarup |first3=N. |last4=Racimo |first4=F. |last5=Bjerregaard |first5=P. |last6=Jorgensen |first6=M. E. |last7=Korneliussen |first7=T. S. |last8=Gerbault |first8=P. |last9=Skotte |first9=L. |last10=Linneberg |first10=A. |last11=Christensen |first11=C. |last12=Brandslund |first12=I. |last13=Jorgensen |first13=T. |last14=Huerta-Sanchez |first14=E. |last15=Schmidt |first15=E. B. |last16=Pedersen |first16=O. |last17=Hansen |first17=T. |last18=Albrechtsen |first18=A. |last19=Nielsen |first19=R. |s2cid=546365 |title=Greenlandic Inuit show genetic signatures of diet and climate adaptation |journal=Science |date=17 September 2015 |volume=349 |issue=6254 |pages=1343–1347 |doi=10.1126/science.aab2319 |pmid=26383953 |bibcode=2015Sci...349.1343F |hdl=10044/1/43212 |hdl-access=free }}</ref>

Extinction of late surviving [[Biology:Archaic humans|archaic humans]] at the beginning of the [[Earth:Holocene|Holocene]] (12 ka).
Accelerated divergence due to selection pressures in populations participating in the [[History:Neolithic Revolution|Neolithic Revolution]] after 12 ka, e.g.
East Asian types of ADH1B associated with [[Biology:Oryza sativa#Continental East Asia|rice domestication]],<ref>{{cite journal |last1=Peng |first1=Yi |last2=Shi |first2=Hong |last3=Qi |first3=Xue-bin |last4=Xiao |first4=Chun-jie |last5=Zhong |first5=Hua |last6=Ma |first6=Run-lin Z |last7=Su |first7=Bing |title=The ADH1B Arg47His polymorphism in East Asian populations and expansion of rice domestication in history |journal=BMC Evolutionary Biology |date=2010 |volume=10 |issue=1 |pages=15 |doi=10.1186/1471-2148-10-15 |pmid=20089146 |pmc=2823730 |doi-access=free }}</ref> or  [[Biology:Lactase persistence|lactase persistence]].<ref>{{cite journal |last1=Ségurel |first1=Laure |last2=Bon |first2=Céline |title=On the Evolution of Lactase Persistence in Humans |journal=Annual Review of Genomics and Human Genetics |date=31 August 2017 |volume=18 |issue=1 |pages=297–319 |doi=10.1146/annurev-genom-091416-035340 |pmid=28426286 }}</ref><ref>{{cite journal |last1=Ingram |first1=Catherine J. E. |last2=Mulcare |first2=Charlotte A. |last3=Itan |first3=Yuval |last4=Thomas |first4=Mark G. |last5=Swallow |first5=Dallas M. |s2cid=3329285 |title=Lactose digestion and the evolutionary genetics of lactase persistence |journal=Human Genetics |date=26 November 2008 |volume=124 |issue=6 |pages=579–591 |doi=10.1007/s00439-008-0593-6 |pmid=19034520 }}</ref> A slight decrease in brain size occurred a few thousand years ago.
|-
|}

==See also==
{{div col |colwidth=30em}}
* [[Philosophy:Evolution of human intelligence|Evolution of human intelligence]]
* [[Astronomy:Graphical timeline of the universe|Graphical timeline of the universe]]
* [[Biology:Human evolution|Human evolution]]
* [[Biology:Recent human evolution|Recent human evolution]]
* [[Biology:List of human evolution fossils|List of human evolution fossils]]
* [[Earth:Natural history|Natural history]]
* [[History:Human history|Human history]]
* [[Earth:History of Earth|History of Earth]]
* [[History:Timeline of prehistory|Timeline of prehistory]]
* [[Biology:Timeline of the evolutionary history of life|Timeline of the evolutionary history of life]]
* [[History:List of timelines|List of timelines]]
{{div col end}}

==References==
{{Reflist}}

==External links==
* [http://www.palaeos.com Palaeos]
* [https://web.archive.org/web/20110707140754/http://andabien.com/html/hominid-timeline.htm Hominid Timeline]
* [http://www.ucmp.berkeley.edu Berkeley Evolution]
* [http://sci.waikato.ac.nz/evolution/AnimalEvolution.shtml History of Animal Evolution]
* [http://tolweb.org/Life_on_Earth/1 Tree of Life Web Project] – explore complete phylogenetic tree interactively
* [http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive Human Timeline (Interactive)] – Smithsonian, National Museum of Natural History (August 2016).

{{Human Evolution}}
{{Evolution}}

{{DEFAULTSORT:Timeline Of Human Evolution}}
[[Category:Biology timelines|Human evolution]]
[[Category:Human evolution|*]]

{{Sourceattribution|Timeline of human evolution}}