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Timeline of human evolution

From Wikipedia, the free encyclopedia

Haeckel's Paleontological Tree of Vertebrates (c. 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 While Haeckel's tree is outdated, it illustrates clearly the principles that more complex and accurate modern reconstructions can obscure.

Thetimeline of human evolution outlines the major events in the evolutionary lineage of themodern human species,Homo sapiens, throughout thehistory of life, beginning some 4 billion years ago down torecent evolution withinH. sapiens during and since theLast Glacial Period.

It includes brief explanations of the varioustaxonomic ranks in the human lineage. The timeline reflects the mainstream views inmodern taxonomy, based on the principle ofphylogenetic nomenclature;in cases of open questions with no clear consensus, the main competing possibilities are briefly outlined.

Overview of taxonomic ranks

[edit]

A tabular overview of thetaxonomic ranking ofHomo sapiens (with age estimates for each rank) is shown below.

RankNameCommon nameStarted
(millions
of years ago)
Life4,200
Archaea3,700
DomainEukaryotaEukaryotes2,100
OpimodaExcludes Plants and their relatives1,540
Amorphea
ObazoaExcludesAmoebozoa (Amoebas)
OpisthokontaHolozoa +Holomycota (Cristidicoidea andFungi)1,300
HolozoaExcludes Holomycota1,100
FilozoaChoanozoa +Filasterea
ChoanozoaChoanoflagellates + Animals900
KingdomAnimaliaAnimals610
SubkingdomEumetazoaExcludesPorifera (Sponges)
ParahoxozoaExcludesCtenophora (Comb Jellies)
BilateriaTriploblasts / Worms560
Nephrozoa
DeuterostomiaDivision fromProtostomes
PhylumChordataChordates (Vertebrates and closely related invertebrates)530
OlfactoresExcludescephalochordates (Lancelets)
SubphylumVertebrataFish / Vertebrates505
InfraphylumGnathostomataJawed fish460
TeleostomiBony fish420
SarcopterygiiLobe finned fish
SuperclassTetrapodaTetrapods (animals with four limbs)395
AmniotaAmniotes (fully terrestrial tetrapods whose eggs are"equipped with an amnion")340
SynapsidaProto-Mammals308
TherapsidaLimbs beneath the body and other mammalian traits280
ClassMammaliaMammals220
SubclassTheriaMammals that give birth to live young (i.e. non-egg-laying)160
InfraclassEutheriaPlacental mammals (i.e. non-marsupials)125
MagnorderBoreoeutheriaSupraprimates, (most) hoofed mammals, (most) carnivorous mammals, cetaceans, and bats124–101
SuperorderEuarchontogliresSupraprimates: primates, colugos, tree shrews, rodents, and rabbits100
GrandorderEuarchontaPrimates,colugos, andtree shrews99–80
MirorderPrimatomorphaPrimates and colugos79.6
OrderPrimatesPrimates /Plesiadapiformes66
SuborderHaplorrhini"Dry-nosed" (literally, "simple-nosed") primates:tarsiers andmonkeys (incl. apes)63
InfraorderSimiiformesmonkeys (incl. apes)40
ParvorderCatarrhini"Downward-nosed" primates: apes and old-world monkeys30
SuperfamilyHominoideaApes: great apes andlesser apes (gibbons)22–20
FamilyHominidaeGreat apes:humans,chimpanzees,gorillas andorangutans—thehominids20–15
SubfamilyHomininaeHumans, chimpanzees, and gorillas (the African apes)[1]14–12
TribeHomininiIncludes bothHomo andPan (chimpanzees), butnotGorilla.10–8
SubtribeHomininaGenusHomo and close human relatives and ancestors aftersplitting fromPan—thehominins8–4[2]
(Genus)Ardipithecuss.l.6-4
(Genus)Australopithecus3
GenusHomo(H. habilis)Humans2.5
(Species)H. erectus s.l.
(Species)H. heidelbergensis s.l.
SpeciesHomo sapienss.s.Anatomically modern humans0.8–0.3[3]

Timeline

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Unicellular life

[edit]
DateEvent
4.3-4.1 Ga
The earliest life appears, possibly asprotocells. Their genetic material was probably composed ofRNA, capable of both self replication and enzymatic activity; theirmembranes were composed oflipids. Thegenes were separate strands, translated intoproteins and often exchanged between the protocells.
4.0-3.8 GaProkaryoticcells appear; their genetic materials are composed of the more stableDNA and they use proteins for various reasons, primarily for aiding DNA to replicate itself by proteinaceousenzymes (RNA now acts as an intermediary in thiscentral dogma of genetic information flow of cellular life);genes are now linked in sequences so all information passes to offsprings. They hadcell walls &outer membranes and were probably initiallythermophiles.
Further information:Cell (biology) § Origins
3.5 GaThis marks the first appearance ofcyanobacteria and their method of oxygenic photosynthesis and therefore the first occurrence of atmosphericoxygen on Earth.

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

2.5-2.2 GaFirst organisms to use oxygen. By 2400 Ma, in what is referred to as theGreat Oxidation Event, (GOE), most of the pre-oxygen anaerobic forms of life were wiped out by the oxygen producers.
Further information:Geological history of oxygen
2.2-1.8 GaOrigin of theeukaryotes: organisms withnuclei,endomembrane systems (includingmitochondria) and complexcytoskeletons; theysplicedmRNA betweentranscription andtranslation (splicing also occurs in prokaryotes, but it is only ofnon-coding RNAs). 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 anaerobe one, which later became mitochondria) to depend on each other, leading toendosymbiosis. Early eukaryotes lost their cell walls and outer membranes.
1.2 GaSexual reproduction evolves (mitosis andmeiosis) by this time at least, leading to faster evolution[4] where genes are mixed in every generation enabling greater variation for subsequent selection.
1.2-0.8 Ga
Choanoflagellate

TheHolozoa lineage of eukaryotes evolves many features for makingcell colonies, and finally leads to the ancestor of animals (metazoans) andchoanoflagellates.[5][6]

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 ofcellular specialization for different tasks.

Animalia

[edit]
DateEvent
800–650Ma
Dickinsonia costata from theEdiacaran biota, 635–542 Ma, a possible early member ofAnimalia.

Urmetazoan:The first fossils that might representanimals appear in the 665-million-year-old rocks of theTrezona Formation ofSouth Australia. These fossils are interpreted as being early sponges.[7]Multicellular animals may have existed from 800 Ma. Separation from thePorifera (sponges) lineage.Eumetazoa/Diploblast: separation from theCtenophora ("comb jellies") lineage.Planulozoa/ParaHoxozoa: separation from thePlacozoa andCnidaria lineages.All diploblasts possessepithelia,nerves,muscles andconnective tissue andmouths, and except for placozoans, have some form of symmetry, with their ancestors probably havingradial symmetry like that of cnidarians. Diploblasts separated their early embryonic cells into twogerm layers (ecto- andendoderm).Photoreceptive eye-spots evolve.

650-600 Ma
Proporus sp., axenacoelomorph.

Urbilaterian:the last common ancestor of xenacoelomorphs,protostomes (including thearthropod [insect, crustacean, spider],mollusc [squid, snail, clam] andannelid [earthworm] lineages) and thedeuterostomes (including the vertebrate [human] lineage) (the last two are more related to each other and calledNephrozoa). Xenacoelomorphs all have agonopore to expelgametes but nephrozoans merged it with theiranus. Earliest development ofbilateral symmetry,mesoderm,head (anteriorcephalization) and various gut muscles (and thusperistalsis) and, in the Nephrozoa,nephridia (kidney precursors),coelom (or maybepseudocoelom), distinct mouth and anus (evolution ofthrough-gut), and possibly evennerve cords andblood vessels.[8] Reproductive tissue probably concentrates into a pair ofgonads connecting just before the posterior orifice. "Cup-eyes" andbalance organs evolve (the function of hearing added later as the more complexinner ear evolves in vertebrates). The nephrozoan through-gut had a wider portion in the front, called thepharynx. Theintegument or skin consists of an epithelial layer (epidermis) and a connective layer.

600-540 Ma
Asea cucumber (Actinopyga echinites), displaying its feeding tentacles andtube feet.

Most known animal phyla appeared in the fossil record as marine species during theEdiacaran-Cambrian explosion, probably caused by long scale oxygenation since around 585 Ma (sometimes called theNeoproterozoic Oxygenation Event or NOE) and also an influx of oceanic minerals.Deuterostomes, the last common ancestor of theChordata [human] lineage,Hemichordata (acorn worms andgraptolites) andEchinodermata (starfish,sea urchins,sea cucumbers, etc.), probably had both ventral anddorsal nerve cords like modern acorn worms.

An archaic survivor from this stage is theacorn worm, sporting anopen circulatory system (with less branched blood vessels) with a heart that also functions as a kidney. Acorn worms have aplexus 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.[9] Deuterostomes also evolvedpharyngeal slits, which were probably used forfilter feeding like in hemi- and proto-chordates.

Chordata

[edit]
DateEvent
540-520 Ma
Pikaia

The increased amount of oxygen causes many eukaryotes, including most animals, to becomeobligate aerobes.

The Chordata ancestor gave rise to thelancelets (Amphioxii) and Olfactores. Ancestral chordates evolved a post-analtail,notochord, andendostyle (precursor of thyroid). The pharyngeal slits (orgills) are now supported by connective tissue and used for filter feeding and possiblybreathing. The first of these basal chordates to be discovered by science wasPikaia gracilens.[10] Other, earlier chordate predecessors includeMyllokunmingia fengjiaoa,[11]Yunnanozoon lividum,[12] andHaikouichthys ercaicunensis.[13] They probably lost their ventral nerve cord and evolved a special region of thedorsal one, called thebrain, withglia becoming permanently associated withneurons. They probably evolved the firstblood cells (probably earlyleukocytes, indicating advancedinnate immunity), which they made around the pharynx and gut.[14] All chordates except tunicates sport an intricate,closed circulatory system, with highly branched blood vessels.

Olfactores, last common ancestor oftunicates and vertebrates in whicholfaction (smell) evolved. Since lancelets lack aheart, 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).

520-480 Ma
Agnatha

The firstvertebrates ("fish") appear: theAgnathans. They were jawless, had seven pairs ofpharyngeal arches like their descendants today, and theirendoskeletons were cartilaginous (then only consisting of thechondrocranium/braincase andvertebrae). The jawlessCyclostomata diverge at this stage. The connective tissue below the epidermis differentiates into thedermis andhypodermis.[15] They depended ongills for respiration and evolved the unique sense oftaste (the remaining sense of the skin now called "touch"),endothelia, cameraeyes and inner ears (capable of hearing and balancing; each consists of alagena, anotolithic organ and twosemicircular canals) as well aslivers,thyroids,kidneys and two-chambered hearts (oneatrium and oneventricle). They had a tailfin 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). Thepineal gland of the brain penetrates to the level of the skin on the head, making it seem like athird eye. They evolved the firsterythrocytes andthrombocytes.[16]

460-430 Ma
Aplacoderm

ThePlacodermi werethe first jawed fishes (Gnathostomata); theirjaws evolved from the firstgill/pharyngeal arch and they largely replaced their endoskeletalcartilage withbone and evolved pectoral and pelvic fins. Bones of the first gill arch became theupper andlower jaw, while those from the second arch became thehyomandibula, 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 ofspiracles, 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, thecephalopods andsea scorpions, and rose to dominance themselves. A lineage of them probably evolved into the bony andcartilaginous fish, after evolvingscales,teeth (which allowed the transition to fullcarnivory),stomachs,spleens,thymuses,myelin sheaths,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 asaccule andutricle.

430-410 Ma
Coelacanth caught in 1974
Bony fish split their jaws into several bones and evolvelungs, fin bones, two pairs ofrib bones, andopercular bones, and diverge into theactinopterygii (withray fins) and thesarcopterygii (with fleshy, lower fins);[17] the latter transitioned from marine to freshwater habitats. Jawed fish also possess dorsal and anal fins.

Tetrapoda

[edit]
Further information:Evolution of tetrapods
DateEvent
390 Ma
Panderichthys

Some freshwater lobe-finned fish (sarcopterygii) develop limbs and give rise to theTetrapodomorpha. These fish evolved in shallow andswampy freshwaterhabitats, where they evolved large eyes and spiracles.

Primitive tetrapods ("fishapods") developed from tetrapodomorphs with a two-lobedbrain 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 andbones. (The "living fossil"coelacanth is a relatedlobe-finned fish without these shallow-water adaptations.) Tetrapod fishes used their fins as paddles in shallow-water habitats choked with plants anddetritus. 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.[18]

Panderichthys is a 90–130 cm (35–50 in) long fish from the LateDevonian period (380Mya). It has a large tetrapod-like head.Panderichthys exhibits features transitional between lobe-finned fishes and early tetrapods.

Trackway impressions made by something that resemblesIchthyostega's limbs were formed 390 Ma in Polish marine tidal sediments. This suggests tetrapod evolution is older than the dated fossils ofPanderichthys through toIchthyostega.

375-350 Ma
Tiktaalik

Tiktaalik is a genus ofsarcopterygian (lobe-finned) fishes from the late Devonian with many tetrapod-like features. It shows a clear link betweenPanderichthys andAcanthostega.

Acanthostega
Ichthyostega

Acanthostega is an extincttetrapod, 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 arib cage to support the lungs, while the ventral pair disappears.

Ichthyostega is another extinct tetrapod. Being one of the first animals with only two pairs oflimbs (also unique since they end indigits and have bones),Ichthyostega is seen as an intermediate between a fish and an amphibian.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 themud.[19] 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 asgastralia). Many evolutionary changes occurred at this stage:eyelids andtearglands evolved to keep the eyes wet out of water and the eyes became connected to the pharynx for draining the liquid; the hyomandibula (now calledcolumella) shrank into the spiracle, which now also connected to the inner ear at one side and the pharynx at another, becoming theEustachian tube (columella assisted in hearing); an earlyeardrum (a patch of connective tissue) evolved on the end of each tube (called theotic notch); and the ceratohyal and basihyal merged into thehyoid. These "fishapods" had moreossified and stronger bones to support themselves on land (especially skull andlimb bones). Jaw bones fuse together while gill and opercular bones disappear.

350-330 Ma
Pederpes

Pederpes from around 350 Ma indicates that the standard number of 5 digits evolved at theEarly 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 evolvedvomeronasal organs,salivary glands,tongues,parathyroid glands, three-chambered hearts (with two atria and one ventricle) andbladders, and completely removed their gills by adulthood. Theglottis evolves to prevent food going into therespiratory tract. Lungs and thin, moist skin allowed them to breathe; water was also needed to give birth to shell-lesseggs and for early development. Dorsal, anal and tail fins all disappeared.

Lissamphibia (extant amphibians) retain many features of early amphibians but they have only four digits (caecilians have none).

330-300 Ma
Hylonomus

From amphibians came the firstamniotes:Hylonomus, a primitivereptile, is the earliest amniote known. It was 20 cm (8 in) long (including the tail) and probably would have looked rather similar to modernlizards. It had small sharp teeth and probably ate smallmillipedes andinsects. It is a precursor of later amniotes (including both the reptiles and the ancestors of mammals).Alpha keratin firstevolves here; it is used in the claws of modern amniotes, and hair in mammals, indicatingclaws and a different type ofscales evolved in amniotes (complete loss of gills as well).[20]

Evolution of the amniotic egg allows the amniotes to reproduce on land and layshelled 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 discreteglands.

Amniotes have advanced nervous systems, with twelve pairs ofcranial nerves, unlike lower vertebrates. They also evolved truesternums but lost their eardrums and otic notches (hearing only by columella bone conduction).

Mammalia

[edit]
Further information:Evolution of mammals
DateEvent
300-260 MaShortly after the appearance of the first amniotes, two branches split off. One branch is theSauropsida, from which come the reptiles, including birds. The other branch isSynapsida, from which come modern mammals. Both hadtemporal fenestrae, a pair of holes in their skulls behind the eyes, which were used to increase the space for jaw muscles. Synapsids had one opening on each side, whilediapsids (a branch of Sauropsida) had two. An early, inefficient version ofdiaphragm may have evolved in synapsids.

The earliest synapsids, or "proto-mammals," are thepelycosaurs. The pelycosaurs were the first animals to have temporal fenestrae. Pelycosaurs were nottherapsids but their ancestors. The therapsids were, in turn, the ancestors ofmammals.

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 asecondary 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.[21] They had lost gastralia and, possibly, scales.

260-230 Ma
Cynognathus

One subgroup of therapsids, the cynodonts, losepineal eye and lumbar ribs and very likely becamewarm-blooded. The lower respiratory tract forms intricate branches in the lungparenchyma, ending in highly vascularizedalveoli. Erythrocytes and thrombocytes lose their nuclei whilelymphatic systems and advanced immunity emerge. They may have also had thickerdermis 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 theirorbits. Their hindlimbs became erect and their posterior bones of the jaw progressively shrunk to the region of thecolumella.[22]

230-170 Ma
Repenomamus

FromEucynodontia came the firstmammals. Most early mammals were small shrew-like animals that fed on insects and had transitioned tonocturnality to avoid competition with the dominantarchosaurs — this led to the loss of the vision ofred andultraviolet light (ancestraltetrachromacy of vertebrates reduced todichromacy). Although there is no evidence in the fossil record, it is likely that these animals had aconstant body temperature,hair andmilkglands for their young (the glands stemmed from themilk line). Theneocortex (part of the cerebrum) region of the brain evolves in Mammalia, at the reduction of thetectum (non-smell senses which were processed here became integrated into neocortex but smell became primary sense). Origin of theprostate gland and a pair ofholes opening to the columella and nearby shrinking jaw bones; neweardrums stand in front of the columella and Eustachian tube. The skin becomes hairy, glandular (glands secretingsebum andsweat) and thermoregulatory. Teeth fully differentiate intoincisors,canines,premolars andmolars; mammals becomediphyodont and possess developed diaphragms and males have internalpenises. All mammals have four chamberedhearts (with two atria and two ventricles) and lack cervical ribs (now mammals only have thoracic ribs).

Monotremes are an egg-laying group of mammals represented today by theplatypus andechidna. Recent genome sequencing of the platypus indicates that its sex genes are closer to those of birds than to those of thetherian (live birthing) mammals. Comparing this to other mammals, it can be inferred that the first mammals to gainsexual differentiation through the existence or lack ofSRY gene (found in the y-Chromosome) evolved only in the therians. Early mammals and possibly their eucynodontian ancestors hadepipubic bones, which serve to hold the pouch in modern marsupials (in both sexes).

170-120 Ma
Juramaia sinensis

Evolution of live birth (viviparity), with early therians probably having pouches for keeping their undeveloped young like in modernmarsupials.Nipples stemmed out of the therian milk lines. Theposterior orifice separates into anal and urogenital openings; males possess an external penis.

Monotremes and therians independently detach themalleus andincus from thedentary (lower jaw) and combine them to the shrunken columella (now calledstapes) in thetympanic cavity behind the eardrum (which is connected to the malleus and held by another bone detached from the dentary, thetympanic plusectotympanic), and coil their lagena (cochlea) to advance their hearing, with therians further evolving an externalpinna and erect forelimbs. Femaleplacentalian mammals do not have pouches and epipubic bones but instead have a developedplacenta which penetrates theuterus walls (unlike marsupials), allowing a longergestation; they also have separated urinary and genital openings.[23]

100-90 MaLast common ancestor ofrodents,rabbits,ungulates,carnivorans,bats,shrews and humans (base of the cladeBoreoeutheria; males now haveexternal testicles).

Primates

[edit]
Further information:Evolution of primates
DateEvent
90–66 Ma
Plesiadapis
Carpolestes simpsoni

A group of small, nocturnal, arboreal, insect-eating mammals calledEuarchonta begins a speciation that will lead to theorders ofprimates,treeshrews andflying lemurs. They reduced the number of mammaries to only two pairs (on the chest).Primatomorpha is a subdivision of Euarchonta including primates and their ancestral stem-primatesPlesiadapiformes. An early stem-primate,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.[24] They first appeared in the fossil record around 66 million years ago, soon after theCretaceous–Paleogene extinction event that eliminated about three-quarters of plant and animal species on Earth, including most dinosaurs.[25][26]

One of the last Plesiadapiformes isCarpolestes simpsoni, having grasping digits but not forward-facing eyes.

66-56 MaPrimates diverge into subordersStrepsirrhini (wet-nosed primates) andHaplorrhini (dry-nosed primates). Brain expands and cerebrum divides into 4 pairs oflobes. Thepostorbital bar evolves to separate the orbit from thetemporal fossae as sight regains its position as the primary sense; eyes became forward-facing. Strepsirrhini contain mostprosimians; modern examples includelemurs andlorises. The haplorrhines include the two living groups: prosimiantarsiers, and simianmonkeys, includingapes. The Haplorrhini metabolism lost the ability to producevitamin 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 intonails.
50-35 Ma
Aegyptopithecus

Simians split into infraordersPlatyrrhini andCatarrhini. They fully transitioned todiurnality and lacked any claw andtapetum lucidum (which evolved many times in various vertebrates). They possibly evolved at least some of theparanasal sinuses, and transitioned fromestrous cycle tomenstrual 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 araft of vegetation or via aland bridge (the hypothesis now favored[27]). Catarrhines mostly stayed inAfrica as the two continents drifted apart. Possible early ancestors of catarrhines includeAegyptopithecus andSaadanius.

35-20 Ma
Proconsul

Catarrhini splits into 2 superfamilies,Old World monkeys (Cercopithecoidea) andapes (Hominoidea). Humantrichromatic color vision had its genetic origins in this period. Catarrhines lost the vomeronasal organ (or possibly reduced it to vestigial status).

Proconsul was an early genus of catarrhine primates. They had a mixture ofOld World monkey andape characteristics.Proconsul'smonkey-like features include thintooth enamel, a light build with a narrow chest and short forelimbs, and an arboreal quadrupedal lifestyle. Its ape-like features are its lack of a tail, ape-like elbows, and a slightly larger brain relative to body size.

Proconsul africanus is a possible ancestor of both great and lesser apes, including humans.

Hominidae

[edit]
DateEvent
20-15 MaHominidae (great ape ancestors) speciate from the ancestors of thegibbon (lesser apes) between c. 20 to 16 Ma. They largely reduced their ancestralsnout and lost theuricase enzyme (present in most organisms).[28]
16-12 MaHomininae ancestors speciate from the ancestors of theorangutan between c. 18 to 14 Ma.[29]

Pierolapithecus catalaunicus is thought to be acommon ancestor ofhumans 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, flatrib cage, a stiff lowerspine, flexible wrists, andshoulder blades that lie along its back.

12 MaDanuvius guggenmosi is the first-discovered Late Miocene great ape with preservedlong bones, and greatly elucidates the anatomical structure and locomotion of contemporary apes.[30] 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.
12-8 MaThe clade currently represented by humans and the genusPan (chimpanzees andbonobos) splits from the ancestors of thegorillas between c. 12 to 8 Ma.[31]
8-6 Ma
Sahelanthropus tchadensis

Hominini: Thelatest common ancestor ofhumans andchimpanzeesis estimated to have lived between roughly 10 to 5 million years ago. Both chimpanzees and humans have alarynx that repositions during the first two years of life to a spot between thepharynx 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 ofHominina orHomininae species which lived in this time period includeGraecopithecus (c. 7 Ma),Sahelanthropus tchadensis (c. 7 Ma),Orrorin tugenensis (c. 6 Ma).

Ardipithecus
Ardipithecus is, or may be, a very earlyhominingenus (tribeHominini andsubtribeHominina). Two species are described in the literature:A. ramidus, which lived about 4.4 million years ago[32] during the earlyPliocene, andA. kadabba, dated to approximately 5.6 million years ago[33] (lateMiocene).A. ramidus had a small brain, measuring between 300 and 350 cm3. This is about the same size as the modernbonobo and femalechimpanzee brain; it is somewhat smaller than the brain of australopithecines likeLucy (400 to 550 cm3) and slightly over a fifth the size of the modernHomo 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 probablybipedal as evidenced by its bowl shaped pelvis, the angle of itsforamen magnum and its thinner wrist bones, though its feet were still adapted for grasping rather than walking for long distances.

4-3.5 Ma
Reconstruction of "Lucy"

A member of theAustralopithecus afarensis left human-like footprints on volcanic ash inLaetoli, northernTanzania, 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 earliesthominins—those species that developed and comprised the lineage ofHomo andHomo's closest relatives after the split from the line of the chimpanzees.

It is thought thatA. afarensis was ancestral to both the genusAustralopithecus and the genusHomo. Compared to the modern and extinct greatapes,A. afarensis had reduced canines and molars, although they were still relatively larger than in modern humans.A. afarensis also has a relatively small brain size (380–430 cm3) and a prognathic (anterior-projecting) face.

Australopithecines have been found in savannah environments; they probably developed their diet to include scavenged meat. Analyses ofAustralopithecus africanus lowervertebrae suggests that these bones changed in females to support bipedalism even during pregnancy.

3.5–3.0 MaKenyanthropus platyops, a possible ancestor ofHomo, emerges from theAustralopithecus. Stone tools are deliberately constructed, possibly byKenyanthropus platyops orAustralopithecus afarensis.[34]
3 MaThe bipedalaustralopithecines (a genus of the subtribeHominina) evolve in the savannas ofAfrica being hunted byMegantereon. Loss ofbody hair occurs from 3 to 2 Ma, in parallel with the development of fullbipedalism and slight enlargement of the brain.[35]

Homo

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DateEvent
2.8–2.0 Ma

EarlyHomo appears in East Africa, speciating fromaustralopithecine ancestors.TheLower Paleolithic is defined by the beginning of use ofstone tools.Australopithecus garhi was using stone tools at about 2.5 Ma.Homo habilis is the oldest species given the designationHomo, byLeakey et al. in 1964.H. habilis is intermediate betweenAustralopithecus afarensis andH. erectus, and there have been suggestions to re-classify it within genusAustralopithecus, asAustralopithecus habilis.

LD 350-1 is now considered the earliest known specimen of thegenusHomo, dating to 2.75–2.8 Ma, found in theLedi-Geraru site in theAfar Region ofEthiopia. It is currently unassigned to a species, and it is unclear if it represents the ancestor toH. habilis andH. rudolfensis, which are estimated to have evolved around 2.4 Ma.[36]

Stone tools found at theShangchen site in China and dated to 2.12 million years ago are considered the earliest known evidence ofhominins outside Africa, surpassingDmanisi hominins found in Georgia by 300,000 years, although whether these hominins were an early species in the genusHomo or another hominin species is unknown.[37]

Further information:Homo naledi andHomo rudolfensis
1.9–0.8 Ma
Reconstruction of a femaleH. erectus

Homo erectus derives from earlyHomo or lateAustralopithecus.

Homo habilis, although significantly different of anatomy and physiology, is thought to be the ancestor ofHomo ergaster, or AfricanHomo erectus; but it is also known to have coexisted withH. 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 developcontrol of fire, by about 1.5 Ma.

H. erectus later migratesthroughout Eurasia, reaching Southeast Asia by 0.7 Ma.It is described in a number ofsubspecies.[38] 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.[39]

Homo antecessor may be a common ancestor ofHomo sapiens andNeanderthals.[40][41] At present estimate, humans have approximately 20,000–25,000genes and share 99% of theirDNA with the now extinctNeanderthal[42] and 95–99% of theirDNA with their closest living evolutionary relative, thechimpanzees.[43][44] The human variant of theFOXP2 gene (linked to the control of speech) has been found to be identical in Neanderthals.[45]

0.8–0.3 Ma
Reconstruction ofHomo heidelbergensis

Divergence ofNeanderthal andDenisovan lineages from a common ancestor.[46]Homo heidelbergensis (in Africa also known asHomo 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 theSima de los Huesos fossils published in 2016 seems to suggest thatH. 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 ofH. heidelbergensis, to about 600,000 to 800,000 years ago, the approximate age ofHomo antecessor.[47][48] Brain expansion (enlargement) between 0.8 and 0.2 Ma may have occurred due to the extinction of most Africanmegafauna (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 afterMid-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 withH. heidelbergensis were found in southern Italy in 2003.[49]

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 ofH. erectus). By 200 ka, humans had stopped their brain expansion.

Homo sapiens

[edit]
Further information:Homo sapiens,Neanderthal,Interbreeding between archaic and modern humans,Recent human evolution, andHuman genetic variation
DateEvent
300–130 ka
Reconstruction of earlyHomo sapiens fromJebel Irhoud, Moroccoc. 315 000 years BP

Neanderthals andDenisovans emerge from the northernHomo heidelbergensis lineage around 500-450 ka whilesapients emerge from the southern lineage around 350-300 ka.[50]

Fossils attributed toH. sapiens, along with stone tools, dated to approximately 300,000 years ago, found atJebel Irhoud, Morocco[51] yield the earliest fossil evidence foranatomically modernHomo sapiens.Modern human presence inEast Africa (Gademotta), at 276 kya.[52] In July 2019, anthropologists reported the discovery of 210,000 year old remains of what may possibly have been aH. sapiens inApidima Cave,Peloponnese,Greece.[53][54][55]

Patrilineal andmatrilineal most recent common ancestors (MRCAs) of living humans roughly between 200 and 100 kya[56][57]with some estimates on the patrilineal MRCA somewhat higher, ranging up to 250 to 500 kya.[58]

160,000 years ago,Homo sapiens idaltu in theAwash River Valley (near present-dayHerto village, Ethiopia) practicedexcarnation.[59]

130–80 kaMarine Isotope Stage 5 (Eemian).

Modern human presence inSouthern Africa andWest Africa.[60]Appearance of mitochondrial haplogroup (mt-haplogroup)L2.

80–50 kaMIS 4, beginning of theUpper Paleolithic.

Early evidence forbehavioral modernity.[61]Appearance of mt-haplogroupsM andN.Southern Dispersalmigration out of Africa,Proto-Australoidpeopling of Oceania.[62]Archaic admixture fromNeanderthals in Eurasia,[63][64] fromDenisovans in Oceania with trace amounts in Eastern Eurasia,[65] 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.[66][67][68][69]

50–25 ka
Reconstruction ofOase 2 (c. 40 ka)

Behavioral modernity develops by this time or earlier, according to the"great leap forward" theory.[70]Extinction ofHomo floresiensis.[71]M168 mutation (carried by all non-African males).Appearance of mt-haplogroupsU andK.Peopling of Europe, peopling of the North AsianMammoth steppe.Paleolithic art.Extinction of Neanderthals and other archaic human variants (with possible survival ofhybrid populations in Asia and Africa).Appearance of Y-HaplogroupR2; mt-haplogroupsJ andX.

after 25 ka
Reconstruction of aNeolithic farmer from Europe,Science Museum in Trento

Last Glacial Maximum;Epipaleolithic /Mesolithic /Holocene.Peopling of the Americas.Appearance of: Y-HaplogroupR1a; mt-haplogroupsV andT.Variousrecent divergence associated with environmental pressures,e.g.light skin in Europeans and East Asians (KITLG,ASIP), after 30 ka;[72]Inuit adaptation to high-fat diet and cold climate, 20 ka.[73]

Extinction of late survivingarchaic humans at the beginning of theHolocene (12 ka).Accelerated divergence due to selection pressures in populations participating in theNeolithic Revolution after 12 ka, e.g.East Asian types ofADH1B associated withrice domestication,[74] orlactase persistence.[75][76] The past 100,000 years have seen selective reductions in brain size for some human lineages during warmer interglacial periods.[77]

See also

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