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Timeline of human evolution
Timeline of human evolution
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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 While Haeckel's tree is outdated, it illustrates clearly the principles that more complex and accurate modern reconstructions can obscure.

The timeline of human evolution outlines the major events in the evolutionary lineage of the modern human species, Homo sapiens, throughout the history of life, beginning some 4 billion years ago down to recent evolution within H. sapiens during and since the Last Glacial Period.

It includes brief explanations of the various taxonomic ranks in the human lineage. The timeline reflects the mainstream views in modern taxonomy, based on the principle of phylogenetic 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 the taxonomic ranking of Homo sapiens (with age estimates for each rank) is shown below.

Rank Name Common name Started
(millions
of years ago)
Life 4,200
Archaea 3,700
Domain Eukaryota Eukaryotes 2,100
Opimoda Excludes Plants and their relatives 1,540
Amorphea
Obazoa Excludes Amoebozoa (Amoebas)
Opisthokonta Holozoa + Holomycota (Cristidicoidea and Fungi) 1,300
Holozoa Excludes Holomycota 1,100
Filozoa Choanozoa + Filasterea
Choanozoa Choanoflagellates + Animals 900
Kingdom Animalia Animals 610
Subkingdom Eumetazoa Excludes Porifera (Sponges)
Parahoxozoa Excludes Ctenophora (Comb Jellies)
Bilateria Triploblasts / Worms 560
Nephrozoa
Deuterostomia Division from Protostomes
Phylum Chordata Chordates (Vertebrates and closely related invertebrates) 530
Olfactores Excludes cephalochordates (Lancelets)
Subphylum Vertebrata Fish / Vertebrates 505
Infraphylum Gnathostomata Jawed fish 460
Teleostomi Bony fish 420
Sarcopterygii Lobe finned fish
Superclass Tetrapoda Tetrapods (animals with four limbs) 395
Amniota Amniotes (fully terrestrial tetrapods whose eggs are "equipped with an amnion") 340
Synapsida Proto-Mammals 308
Therapsida Limbs beneath the body and other mammalian traits 280
Class Mammalia Mammals 220
Subclass Theria Mammals that give birth to live young (i.e. non-egg-laying) 160
Infraclass Eutheria Placental mammals (i.e. non-marsupials) 125
Magnorder Boreoeutheria Supraprimates, (most) hoofed mammals, (most) carnivorous mammals, cetaceans, and bats 124–101
Superorder Euarchontoglires Supraprimates: primates, colugos, tree shrews, rodents, and rabbits 100
Grandorder Euarchonta Primates, colugos, and tree shrews 99–80
Mirorder Primatomorpha Primates and colugos 79.6
Order Primates Primates / Plesiadapiformes 66
Suborder Haplorrhini "Dry-nosed" (literally, "simple-nosed") primates: tarsiers and monkeys (incl. apes) 63
Infraorder Simiiformes monkeys (incl. apes) 40
Parvorder Catarrhini "Downward-nosed" primates: apes and old-world monkeys 30
Superfamily Hominoidea Apes: great apes and lesser apes (gibbons) 22–20
Family Hominidae Great apes: humans, chimpanzees, gorillas and orangutans—the hominids 20–15
Subfamily Homininae Humans, chimpanzees, and gorillas (the African apes)[1] 14–12
Tribe Hominini Includes both Homo and Pan (chimpanzees), but not Gorilla. 10–8
Subtribe Hominina Genus Homo and close human relatives and ancestors after splitting from Pan—the hominins 8–4[2]
(Genus) Ardipithecus s.l. 6-4
(Genus) Australopithecus 3
Genus Homo (H. habilis) Humans 2.5
(Species) H. erectus s.l.
(Species) H. heidelbergensis s.l.
Species Homo sapiens s.s. Anatomically modern humans 0.8–0.3[3]

Timeline

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−4500 —
−4000 —
−3500 —
−3000 —
−2500 —
−2000 —
−1500 —
−1000 —
−500 —
0 —
 
 
 
 
 
Plants
 
 
 
 

Unicellular life

[edit]
Date Event
4.3-4.1 Ga
The earliest life appears, possibly as protocells. Their genetic material was probably composed of RNA, capable of both self replication and enzymatic activity; their membranes were composed of lipids. The genes were separate strands, translated into proteins and often exchanged between the protocells.
4.0-3.8 Ga Prokaryotic cells appear; their genetic materials are composed of the more stable DNA and they use proteins for various reasons, primarily for aiding DNA to replicate itself by proteinaceous enzymes (RNA now acts as an intermediary in this central dogma of genetic information flow of cellular life); genes are now linked in sequences so all information passes to offsprings. They had cell walls & outer membranes and were probably initially thermophiles.
3.5 Ga This marks the first appearance of cyanobacteria and their method of oxygenic photosynthesis and therefore the first occurrence of atmospheric oxygen on Earth.

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

2.5-2.2 Ga First organisms to use oxygen. By 2400 Ma, in what is referred to as the Great Oxidation Event, (GOE), most of the pre-oxygen anaerobic forms of life were wiped out by the oxygen producers.
2.2-1.8 Ga Origin of the eukaryotes: organisms with nuclei, endomembrane systems (including mitochondria) and complex cytoskeletons; they spliced mRNA between transcription and translation (splicing also occurs in prokaryotes, but it is only of non-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 an aerobe one, which later became mitochondria) to depend on each other, leading to endosymbiosis. Early eukaryotes lost their cell walls and outer membranes.
1.2 Ga Sexual reproduction evolves (mitosis and meiosis) 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

The Holozoa lineage of eukaryotes evolves many features for making cell colonies, and finally leads to the ancestor of animals (metazoans) and choanoflagellates.[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 of cellular specialization for different tasks.

Animalia

[edit]
Date Event
800–650 Ma
Dickinsonia costata from the Ediacaran biota, 635–542 Ma, a possible early member of Animalia.

Urmetazoan: The first fossils that might represent animals appear in the 665-million-year-old rocks of the Trezona Formation of South Australia. These fossils are interpreted as being early sponges.[7] Multicellular animals may have existed from 800 Ma. Separation from the Porifera (sponges) lineage. Eumetazoa/Diploblast: separation from the Ctenophora ("comb jellies") lineage. Planulozoa/ParaHoxozoa: separation from the Placozoa and Cnidaria lineages. All diploblasts possess epithelia, nerves, muscles and connective tissue and mouths, 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 (ecto- and endoderm). Photoreceptive eye-spots evolve.

650-600 Ma
Proporus sp., a xenacoelomorph.

Urbilaterian: the last common ancestor of xenacoelomorphs, protostomes (including the arthropod [insect, crustacean, spider], mollusc [squid, snail, clam] and annelid [earthworm] lineages) and the deuterostomes (including the vertebrate [human] lineage) (the last two are more related to each other and called Nephrozoa). Xenacoelomorphs all have a gonopore to expel gametes but nephrozoans merged it with their anus. Earliest development of bilateral symmetry, mesoderm, head (anterior cephalization) and various gut muscles (and thus peristalsis) and, in the Nephrozoa, nephridia (kidney precursors), coelom (or maybe pseudocoelom), distinct mouth and anus (evolution of through-gut), and possibly even nerve cords and blood vessels.[8] Reproductive tissue probably concentrates into a pair of gonads connecting just before the posterior orifice. "Cup-eyes" and 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 pharynx. The integument or skin consists of an epithelial layer (epidermis) and a connective layer.

600-540 Ma
A 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 Ediacaran-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 Chordata [human] lineage, Hemichordata (acorn worms and graptolites) and Echinodermata (starfish, sea urchins, sea cucumbers, etc.), probably had both ventral and dorsal nerve cords like modern acorn worms.

An archaic survivor from this stage is the 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 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.[9] Deuterostomes also evolved pharyngeal slits, which were probably used for filter feeding like in hemi- and proto-chordates.

Chordata

[edit]
Date Event
540-520 Ma
Pikaia

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

The Chordata ancestor gave rise to the lancelets (Amphioxii) and Olfactores. Ancestral chordates evolved a post-anal 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 breathing. The first of these basal chordates to be discovered by science was Pikaia gracilens.[10] Other, earlier chordate predecessors include Myllokunmingia fengjiaoa,[11] Yunnanozoon lividum,[12] and Haikouichthys ercaicunensis.[13] They probably lost their ventral nerve cord and evolved a special region of the dorsal one, called the brain, with glia becoming permanently associated with neurons. They probably evolved the first blood cells (probably early leukocytes, indicating advanced innate 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 of tunicates and vertebrates in which olfaction (smell) evolved. Since lancelets lack a 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).

520-480 Ma
Agnatha

The first vertebrates ("fish") appear: the Agnathans. They were jawless, had seven pairs of pharyngeal arches like their descendants today, and their endoskeletons were cartilaginous (then only consisting of the chondrocranium/braincase and vertebrae). The jawless Cyclostomata diverge at this stage. The connective tissue below the epidermis differentiates into the dermis and hypodermis.[15] They depended on gills for respiration and evolved the unique sense of taste (the remaining sense of the skin now called "touch"), endothelia, camera eyes and inner ears (capable of hearing and balancing; each consists of a lagena, an otolithic organ and two semicircular canals) as well as livers, thyroids, kidneys and two-chambered hearts (one atrium and one ventricle). They had a tail 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 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.[16]

460-430 Ma
A placoderm

The Placodermi were the first jawed fishes (Gnathostomata); their jaws evolved from the first gill/pharyngeal arch and they largely replaced their endoskeletal cartilage with bone and evolved pectoral and pelvic fins. Bones of the first gill arch became the upper and lower jaw, while those from the second arch became the 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 sea scorpions, and rose to dominance themselves. A lineage of them probably evolved into the bony and cartilaginous fish, after evolving scales, teeth (which allowed the transition to full carnivory), 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 a saccule and utricle.

430-410 Ma
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);[17] the latter transitioned from marine to freshwater habitats. Jawed fish also possess dorsal and anal fins.

Tetrapoda

[edit]
Date Event
390 Ma
Panderichthys

Some freshwater lobe-finned fish (sarcopterygii) develop limbs and give rise to the Tetrapodomorpha. These fish evolved in shallow and swampy freshwater habitats, where they evolved large eyes and spiracles.

Primitive tetrapods ("fishapods") developed from tetrapodomorphs with a two-lobed 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 bones. (The "living fossil" coelacanth is a related lobe-finned fish without these shallow-water adaptations.) Tetrapod fishes used their fins as paddles in shallow-water habitats choked with plants and detritus. The universal tetrapod characteristics of front limbs that bend backward at the elbow and hind limbs that bend forward at the knee can plausibly be traced to early tetrapods living in shallow water.[18]

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

Trackway impressions made by something that resembles Ichthyostega's limbs were formed 390 Ma in Polish marine tidal sediments. This suggests tetrapod evolution is older than the dated fossils of Panderichthys through to Ichthyostega.

375-350 Ma
Tiktaalik

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

Acanthostega
Ichthyostega

Acanthostega is an extinct 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.

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. 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 mud.[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 as gastralia). Many evolutionary changes occurred at this stage: eyelids and 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 columella) shrank into the spiracle, which now also connected to the inner ear at one side and the pharynx at another, becoming the Eustachian tube (columella assisted in hearing); an early eardrum (a patch of connective tissue) evolved on the end of each tube (called the otic notch); and the ceratohyal and basihyal merged into the hyoid. These "fishapods" had more ossified and stronger bones to support themselves on land (especially skull and limb 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 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 vomeronasal organs, salivary glands, tongues, parathyroid glands, three-chambered hearts (with two atria and one ventricle) and bladders, and completely removed their gills by adulthood. The glottis evolves to prevent food going into the respiratory tract. Lungs and thin, moist skin allowed them to breathe; water was also needed to give birth to shell-less eggs 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 first amniotes: Hylonomus, a primitive reptile, is the earliest amniote known. It was 20 cm (8 in) long (including the tail) and probably would have looked rather similar to modern lizards. It had small sharp teeth and probably ate small millipedes and insects. It is a precursor of later amniotes (including both the reptiles and the ancestors of mammals). 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 scales evolved in amniotes (complete loss of gills as well).[20]

Evolution of the amniotic egg allows the amniotes to reproduce on land and lay 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 sternums but lost their eardrums and otic notches (hearing only by columella bone conduction).

Mammalia

[edit]
Date Event
300-260 Ma Shortly after the appearance of the first amniotes, two branches split off. One branch is the Sauropsida, from which come the reptiles, including birds. The other branch is Synapsida, from which come modern mammals. Both had temporal fenestrae, a pair of holes in their skulls behind the eyes, which were used to increase the space for jaw muscles. Synapsids had one opening on each side, while diapsids (a branch of Sauropsida) had two. An early, inefficient version of diaphragm may have evolved in synapsids.

The earliest synapsids, or "proto-mammals," are the pelycosaurs. The pelycosaurs were the first animals to have temporal fenestrae. Pelycosaurs were not therapsids 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.[21] They had lost gastralia and, possibly, scales.

260-230 Ma
Cynognathus

One subgroup of therapsids, the cynodonts, lose pineal eye and lumbar ribs and very likely became warm-blooded. The lower respiratory tract forms intricate branches in the lung parenchyma, ending in highly vascularized alveoli. Erythrocytes and thrombocytes lose their nuclei while lymphatic systems and advanced immunity emerge. They may have also had thicker 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 orbits. Their hindlimbs became erect and their posterior bones of the jaw progressively shrunk to the region of the columella.[22]

230-170 Ma
Repenomamus

From Eucynodontia came the first mammals. Most early mammals were small shrew-like animals that fed on insects and had transitioned to nocturnality to avoid competition with the dominant archosaurs — this led to the loss of the vision of red and ultraviolet light (ancestral tetrachromacy of vertebrates reduced to dichromacy). Although there is no evidence in the fossil record, it is likely that these animals had a constant body temperature, hair and milk glands for their young (the glands stemmed from the milk line). The 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 prostate gland and a pair of holes opening to the columella and nearby shrinking jaw bones; new eardrums 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, canines, premolars and molars; mammals become diphyodont and possess developed diaphragms and males have internal penises. All mammals have four chambered hearts (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 the platypus and echidna. Recent genome sequencing of the platypus indicates that its sex genes are closer to those of birds than to those of the therian (live birthing) mammals. Comparing this to other mammals, it can be inferred that the first mammals to gain 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).

170-120 Ma
Juramaia sinensis

Evolution of live birth (viviparity), with early therians probably having pouches for keeping their undeveloped young like in modern marsupials. Nipples stemmed out of the therian milk lines. The 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 stapes) in the 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 (cochlea) to advance their hearing, with therians further evolving an external pinna and erect forelimbs. Female placentalian mammals do not have pouches and epipubic bones but instead have a developed placenta which penetrates the uterus walls (unlike marsupials), allowing a longer gestation; they also have separated urinary and genital openings.[23]

100-90 Ma Last common ancestor of rodents, rabbits, ungulates, carnivorans, bats, shrews and humans (base of the clade Boreoeutheria; males now have external testicles).

Primates

[edit]
Date Event
90–66 Ma
Plesiadapis
Carpolestes simpsoni

A group of small, nocturnal, arboreal, insect-eating mammals called Euarchonta begins a speciation that will lead to the orders of primates, treeshrews and flying 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-primates Plesiadapiformes. 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 the Cretaceous–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 is Carpolestes simpsoni, having grasping digits but not forward-facing eyes.

66-56 Ma Primates diverge into suborders Strepsirrhini (wet-nosed primates) and Haplorrhini (dry-nosed primates). Brain expands and cerebrum divides into 4 pairs of lobes. The 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 prosimians; modern examples include lemurs and lorises. The haplorrhines include the two living groups: prosimian tarsiers, and simian monkeys, including apes. The Haplorrhini metabolism lost the ability to produce 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.
50-35 Ma
Aegyptopithecus

Simians split into infraorders Platyrrhini and Catarrhini. They fully transitioned to diurnality and lacked any claw and tapetum lucidum (which evolved many times in various vertebrates). They possibly evolved at least some of the paranasal sinuses, and transitioned from estrous cycle to 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 raft of vegetation or via a land bridge (the hypothesis now favored[27]). Catarrhines mostly stayed in Africa as the two continents drifted apart. Possible early ancestors of catarrhines include Aegyptopithecus and Saadanius.

35-20 Ma
Proconsul

Catarrhini splits into 2 superfamilies, Old World monkeys (Cercopithecoidea) and apes (Hominoidea). Human trichromatic 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 of Old World monkey and ape characteristics. Proconsul's monkey-like features include thin tooth enamel, a light build with a narrow chest and short forelimbs, and an arboreal quadrupedal lifestyle. Its ape-like features are its lack of a tail, ape-like elbows, and a slightly larger brain relative to body size.

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

Hominidae

[edit]
Date Event
20-15 Ma Hominidae (great ape ancestors) speciate from the ancestors of the gibbon (lesser apes) between c. 20 to 16 Ma. They largely reduced their ancestral snout and lost the uricase enzyme (present in most organisms).[28]
16-12 Ma Homininae ancestors speciate from the ancestors of the orangutan between c. 18 to 14 Ma.[29]

Pierolapithecus catalaunicus is thought to be a common ancestor of humans and the other great apes, or at least a species that brings us closer to a common ancestor than any previous fossil discovery. It had the special adaptations for tree climbing as do present-day humans and other great apes: a wide, flat rib cage, a stiff lower spine, flexible wrists, and shoulder blades that lie along its back.

12 Ma Danuvius guggenmosi is the first-discovered Late Miocene great ape with preserved long 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 Ma The clade currently represented by humans and the genus Pan (chimpanzees and bonobos) splits from the ancestors of the gorillas between c. 12 to 8 Ma.[31]
8-6 Ma
Sahelanthropus tchadensis

Hominini: The latest common ancestor of humans and chimpanzees is estimated to have lived between roughly 10 to 5 million years ago. Both chimpanzees and humans have a larynx that repositions during the first two years of life to a spot between the pharynx and the lungs, indicating that the common ancestors have this feature, a precondition for vocalized speech in humans. Speciation may have begun shortly after 10 Ma, but late admixture between the lineages may have taken place until after 5 Ma. Candidates of Hominina or Homininae species which lived in this time period include Graecopithecus (c. 7 Ma), Sahelanthropus tchadensis (c. 7 Ma), Orrorin tugenensis (c. 6 Ma).

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[32] during the early Pliocene, and A. kadabba, dated to approximately 5.6 million years ago[33] (late Miocene). A. ramidus had a small brain, measuring between 300 and 350 cm3. This is about the same size as the modern bonobo and female chimpanzee brain; it is somewhat smaller than the brain of australopithecines like Lucy (400 to 550 cm3) and slightly over a fifth the size of the modern Homo sapiens brain.

Ardipithecus was arboreal, meaning it lived largely in the forest where it competed with other forest animals for food, no doubt including the contemporary ancestor of the chimpanzees. Ardipithecus was probably bipedal as evidenced by its bowl shaped pelvis, the angle of its foramen magnum and its thinner wrist bones, though its feet were still adapted for grasping rather than walking for long distances.

4-3.5 Ma
Reconstruction of "Lucy"

A member of the 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 Australopithecus and the genus Homo. Compared to the modern and extinct great apes, A. afarensis had reduced canines and molars, although they were still relatively larger than in modern humans. A. afarensis also has a relatively small brain size (380–430 cm3) and a prognathic (anterior-projecting) face.

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

3.5–3.0 Ma Kenyanthropus platyops, a possible ancestor of Homo, emerges from the Australopithecus. Stone tools are deliberately constructed, possibly by Kenyanthropus platyops or Australopithecus afarensis.[34]
3 Ma The bipedal australopithecines (a genus of the subtribe Hominina) evolve in the savannas of Africa being hunted by Megantereon. Loss of body hair occurs from 3 to 2 Ma, in parallel with the development of full bipedalism and slight enlargement of the brain.[35]

Homo

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Date Event
2.8–2.0 Ma

Early Homo appears in East Africa, speciating from australopithecine ancestors. The Lower Paleolithic is defined by the beginning of use of stone tools. Australopithecus garhi was using stone tools at about 2.5 Ma. Homo habilis is the oldest species given the designation Homo, by Leakey et al. in 1964. H. habilis is intermediate between Australopithecus afarensis and H. erectus, and there have been suggestions to re-classify it within genus Australopithecus, as Australopithecus habilis.

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

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 hominins found in Georgia by 300,000 years, although whether these hominins were an early species in the genus Homo or another hominin species is unknown.[37]

1.9–0.8 Ma
Reconstruction of a female H. erectus

Homo erectus derives from early Homo or late Australopithecus.

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 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.[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 of Homo sapiens and Neanderthals.[40][41] At present estimate, humans have approximately 20,000–25,000 genes and share 99% of their DNA with the now extinct Neanderthal[42] and 95–99% of their DNA with their closest living evolutionary relative, the chimpanzees.[43][44] The human variant of the FOXP2 gene (linked to the control of speech) has been found to be identical in Neanderthals.[45]

0.8–0.3 Ma
Reconstruction of Homo heidelbergensis

Divergence of Neanderthal and Denisovan lineages from a common ancestor.[46] Homo heidelbergensis (in Africa also known as 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 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 Homo antecessor.[47][48] Brain expansion (enlargement) between 0.8 and 0.2 Ma may have occurred due to the extinction of most African 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 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.[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 of H. erectus). By 200 ka, humans had stopped their brain expansion.

Homo sapiens

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Date Event
300–130 ka
Reconstruction of early Homo sapiens from Jebel Irhoud, Morocco c. 315 000 years BP

Neanderthals and Denisovans emerge from the northern Homo heidelbergensis lineage around 500-450 ka while sapients emerge from the southern lineage around 350-300 ka.[50]

Fossils attributed to H. sapiens, along with stone tools, dated to approximately 300,000 years ago, found at Jebel Irhoud, Morocco[51] yield the earliest fossil evidence for anatomically modern Homo sapiens. Modern human presence in East 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 a H. sapiens in Apidima Cave, Peloponnese, Greece.[53][54][55]

Patrilineal and matrilineal 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 the Awash River Valley (near present-day Herto village, Ethiopia) practiced excarnation.[59]

130–80 ka Marine Isotope Stage 5 (Eemian).

Modern human presence in Southern Africa and West Africa.[60] Appearance of mitochondrial haplogroup (mt-haplogroup) L2.

80–50 ka MIS 4, beginning of the Upper Paleolithic.

Early evidence for behavioral modernity.[61] Appearance of mt-haplogroups M and N. Southern Dispersal migration out of Africa, Proto-Australoid peopling of Oceania.[62] Archaic admixture from Neanderthals in Eurasia,[63][64] from Denisovans 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 of Oase 2 (c. 40 ka)

Behavioral modernity develops by this time or earlier, according to the "great leap forward" theory.[70] Extinction of Homo floresiensis.[71] M168 mutation (carried by all non-African males). Appearance of mt-haplogroups U and K. Peopling of Europe, peopling of the North Asian Mammoth steppe. Paleolithic art. 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 J and X.

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

Last Glacial Maximum; Epipaleolithic / Mesolithic / Holocene. Peopling of the Americas. Appearance of: Y-Haplogroup R1a; mt-haplogroups V and T. Various recent 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 surviving archaic humans at the beginning of the Holocene (12 ka). Accelerated divergence due to selection pressures in populations participating in the Neolithic Revolution after 12 ka, e.g. East Asian types of ADH1B associated with rice domestication,[74] or lactase 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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References

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
The timeline of human evolution outlines the major biological, behavioral, and cultural developments that trace the lineage from early hominins to modern Homo sapiens, spanning roughly 7 million years from the divergence of the human-chimpanzee last common ancestor to the present day. This chronological sequence highlights key adaptations such as bipedalism, enlarged brain size, tool manufacture, controlled use of fire, and global migrations, primarily originating in Africa and driven by environmental pressures like climate fluctuations. The process involved multiple hominin species coexisting and interbreeding, with Homo sapiens emerging as the sole surviving species around 40,000 years ago after interactions with relatives like Neanderthals and Denisovans. Human evolution began in the late Miocene epoch, approximately 6–7 million years ago, when the human lineage split from that of chimpanzees in Africa, marked by the appearance of early hominins like Sahelanthropus tchadensis and Orrorin tugenensis, which exhibited initial signs of upright walking and reduced canine teeth. By 4.4 million years ago, Ardipithecus ramidus demonstrated a mix of bipedal locomotion and arboreal capabilities, with a brain size of about 300–350 cubic centimeters, reflecting a transitional phase from forested to more open habitats. The genus Australopithecus, emerging around 4–2 million years ago, solidified bipedalism as a defining trait, as seen in Australopithecus afarensis (e.g., the "Lucy" fossil from 3.2 million years ago), with brain volumes of 390–515 cubic centimeters and adaptations for scavenging in savanna environments. The advent of the genus Homo around 2.8–2.3 million years ago represented a pivotal shift, with species like Homo habilis introducing stone tool use (Oldowan tools by 2.6 million years ago) and gradual brain expansion to support increased meat consumption and social complexity. Homo erectus, appearing by 1.9 million years ago, achieved further milestones including the Acheulean hand axes (1.7 million years ago), control of fire (around 800,000 years ago), and the first major migrations out of Africa to Eurasia by 1.8 million years ago, with brain sizes reaching up to 1,250 cubic centimeters. Later hominins, such as Homo heidelbergensis (700,000–300,000 years ago) and Neanderthals (430,000–40,000 years ago), developed advanced hunting strategies, symbolic behaviors like cave art (evident by 64,000 years ago), and interbreeding with early Homo sapiens. Anatomically modern Homo sapiens first appeared in Africa around 300,000 years ago, as evidenced by fossils from Jebel Irhoud, Morocco, featuring a mix of modern and archaic traits alongside sophisticated Middle Stone Age tools. Early dispersals out of Africa occurred as early as 210,000–100,000 years ago to sites in Israel and China, but the major wave around 60,000–50,000 years ago led to global colonization, behavioral modernity (e.g., burials and art by 50,000 years ago), and the extinction of other hominins. By 12,000 years ago, the onset of agriculture marked the transition to complex societies, culminating in cultural achievements like writing (3,400 BCE) and space exploration (e.g., the 1969 moon landing). This timeline underscores the dynamic, mosaic nature of human evolution, shaped by genetic admixture, environmental adaptation, and cumulative cultural innovation.

Evolutionary Foundations

Taxonomic Ranks in Human Ancestry

The taxonomic classification of human ancestry employs a hierarchical system originally developed by Carl Linnaeus in the 18th century, organizing organisms into nested ranks based on shared characteristics and evolutionary relationships. This Linnaean taxonomy includes eight primary ranks, from broadest to most specific: domain, kingdom, phylum, class, order, family, genus, and species. For humans (Homo sapiens), the classification begins at the domain Eukarya, encompassing all organisms with complex cells containing nuclei; kingdom Animalia, comprising multicellular, heterotrophic organisms capable of locomotion; phylum Chordata, characterized by a notochord, dorsal nerve cord, pharyngeal slits, and post-anal tail at some life stage; class Mammalia, featuring hair, mammary glands, and endothermy; order Primates, distinguished by forward-facing eyes, grasping hands, and large brains; family Hominidae, including great apes and humans; genus Homo, defined by advanced tool use and bipedalism; and species sapiens, noted for high cognitive abilities and cultural complexity. While the traditional Linnaean system relies on a fixed hierarchy of ranks to categorize organisms by morphological similarities, the cladistic approach, developed in the mid-20th century, emphasizes evolutionary branching patterns derived from shared derived traits (synapomorphies) and common ancestry, often visualized through cladograms or phylogenetic trees. In cladistics, taxa are defined as clades—monophyletic groups consisting of an ancestor and all its descendants—rather than artificial ranks that may group unrelated species (paraphyly) or split natural groups (polyphyly). For human ancestry, a simplified cladogram of the lineage might branch as follows: the family Hominidae splits into subfamilies Ponginae (orangutans) and Homininae (African apes and humans), with Homininae further dividing into Gorillini (gorillas) and Hominini (chimpanzees, bonobos, and hominins); within Hominini, the genus Pan (chimpanzees and bonobos) branches from the Homo lineage leading to modern humans. This cladistic framework reveals paraphyletic groups in older taxonomies, such as the traditional "Pongidae" (excluding humans), which omitted the human branch from great ape ancestry, whereas modern Hominidae is strictly monophyletic, encompassing all great apes and humans as descendants of a common ancestor. Key concepts in this classification include monophyly, where a taxon includes an ancestor and all descendants (e.g., Hominidae as a monophyletic family uniting great apes and humans based on shared traits like large body size and taillessness), and paraphyly, where a group excludes some descendants (e.g., earlier definitions of "apes" excluding humans, rendering it paraphyletic). These principles apply directly to human ancestors, such as the tribe Hominini, which is monophyletic and includes the genus Homo alongside extinct hominins like Australopithecus, reflecting their exclusive common ancestry separate from other primates. Recent genomic studies in the 2020s have reinforced and refined this taxonomic structure, with DNA analyses confirming the monophyly of Hominidae and the close Pan-Homo clade while prompting reclassifications of certain fossil hominins based on integrated genetic and morphological evidence. For instance, ancient DNA from Denisovans and Neanderthals has clarified their positions within the monophyletic genus Homo, supporting revisions that integrate archaic humans more fully into the human lineage tree without altering higher ranks. This progression traces human ancestry from early eukaryotes through chordates to Homo sapiens, providing a framework for understanding evolutionary branching.

Dating Methods and Evidence Sources

The timeline of human evolution relies on a combination of absolute and relative dating methods to establish chronological frameworks, drawing from diverse evidence sources such as fossils, genetic material, and archaeological artifacts. Absolute dating provides numerical ages, while relative methods determine the sequence of events without precise years. These techniques are essential for correlating findings across sites and refining our understanding of hominin development. Radiometric dating measures the decay of radioactive isotopes in geological samples, governed by the exponential decay law expressed as N=N0eλtN = N_0 e^{-\lambda t}, where NN is the current amount of the isotope, N0N_0 is the initial amount, λ\lambda is the decay constant, and tt is time elapsed. Carbon-14 dating, applicable to organic remains up to approximately 50,000 years old, exploits the decay of 14^{14}C (half-life of 5,730 years) formed in the atmosphere and absorbed by living organisms. It has dated recent hominin fossils and associated artifacts, such as those from Neanderthal sites. Potassium-argon (K-Ar) dating targets volcanic rocks interlayered with fossils, measuring the decay of 40^{40}K to 40^{40}Ar (half-life of 1.3 billion years), and is widely used for early hominin sites like Olduvai Gorge, providing ages from 100,000 to over 4 billion years. Uranium-lead (U-Pb) dating, effective for carbonates and speleothems in cave sites, tracks the decay chains of 238^{238}U and 235^{235}U to stable lead isotopes (half-lives of 4.5 and 0.7 billion years, respectively), and has dated South African hominid-bearing deposits to between 1 and 3 million years ago. Relative dating techniques establish sequences without numerical values, relying on geological and biological principles. Stratigraphy uses the law of superposition, where deeper layers are older, to order sedimentary deposits containing hominin remains. Biostratigraphy correlates layers based on fossil assemblages, assuming similar faunas indicate contemporaneous deposits, while index fossils—short-lived, widespread species—serve as markers for specific time intervals in paleoanthropological contexts. Evidence for human evolution encompasses fossil records, including skeletal bones and dental remains that preserve morphological traits; genetic data from mitochondrial DNA (mtDNA), tracing maternal lineages, and Y-chromosome analysis for paternal history; and archaeological artifacts such as stone tools and hearths indicating behavioral evolution. Post-2020 advances in paleogenomics have enhanced ancient DNA (aDNA) extraction from degraded samples, enabling the recovery of full genomes from Neanderthals and Denisovans to reveal interbreeding events, such as gene flow contributing 1-4% Neanderthal ancestry in non-African modern humans. Dating methods have inherent limitations, including error margins from statistical uncertainties (often ±1-5% for radiometric techniques) and systematic issues like sample contamination. Radiocarbon dating, for instance, is susceptible to contamination by modern carbon from soil or handling, potentially skewing results by thousands of years, necessitating rigorous pretreatment protocols. K-Ar and U-Pb methods require closed-system assumptions, where excess argon or uranium migration can introduce errors up to 10% in volcanic or carbonate contexts. These challenges are mitigated through cross-validation with multiple methods, ensuring robust timelines for human evolutionary events.

Origins of Complex Life

Unicellular Life and Early Eukaryotes

The origins of life on Earth trace back to the Hadean Eon, approximately 4.1 to 3.8 billion years ago (bya), when abiogenesis—the emergence of life from non-living chemical compounds—likely occurred amid a volatile environment of cooling crust, forming oceans, and intense meteorite bombardment. Recent theories, supported by 2023–2025 research, propose that alkaline hydrothermal vents on the seafloor provided the necessary conditions for this process, offering mineral-rich, energy-gradient environments where organic molecules could concentrate and react to form primitive metabolic cycles and self-replicating systems. For instance, studies have demonstrated how these vents could stabilize RNA precursors and facilitate protometabolic stages, such as carbon fixation, potentially kickstarting life's chemistry as early as 4.0 bya. Updated evidence from microfossils in 3.7 bya rocks in Greenland further corroborates rapid abiogenesis shortly after Earth's formation, with biogenic carbon signatures indicating microbial activity in ancient sediments. A central hypothesis for this early phase is the RNA world, where RNA molecules served dual roles as both genetic material and catalysts, preceding the DNA-protein systems of modern cells and enabling the first replicators around 3.8 bya. This scenario posits that RNA's ability to store information and perform enzymatic functions allowed for the evolution of increasingly complex biochemistry in prebiotic soups or vent microenvironments. The last universal common ancestor (LUCA) of all life likely emerged as a prokaryotic organism approximately 4.2 billion years ago, giving rise to the two primary prokaryotic domains: Bacteria and Archaea, which dominated early Earth for billions of years. These single-celled organisms lacked a nucleus and membrane-bound organelles, relying on circular DNA in the cytoplasm for genetic storage, and reproduced asexually via binary fission. Prokaryotes thrived in diverse niches, with bacteria encompassing oxygenic photosynthesizers like cyanobacteria and archaea adapting to extreme conditions such as high temperatures and salinity, laying the groundwork for metabolic innovations. The Great Oxidation Event (GOE), around 2.4 bya, marked a pivotal shift when cyanobacteria's oxygenic photosynthesis dramatically increased atmospheric oxygen levels from trace amounts to about 1–10%, oxidizing the oceans and atmosphere while causing a mass extinction of anaerobic life but ultimately enabling aerobic respiration and more energy-efficient metabolisms essential for complex life. This oxygenation, driven by prokaryotic activity, created selective pressures that favored organisms capable of handling reactive oxygen species. Eukaryotes, placing humans within the domain Eukarya, emerged around 2.1–1.8 bya through endosymbiosis, where an archaeal host cell engulfed an alpha-proteobacterium that evolved into the mitochondrion, providing aerobic energy production via oxidative phosphorylation. This event is evidenced by mitochondrial genomes retaining bacterial-like features, such as circular DNA and 70S ribosomes, and phylogenetic analyses dating the alphaproteobacterial ancestor to approximately 1.9 bya. Unlike prokaryotes, eukaryotic cells feature a membrane-bound nucleus enclosing linear DNA organized into chromosomes, diverse organelles like the endoplasmic reticulum and Golgi apparatus for compartmentalized functions, and larger sizes (10–100 μm versus 1–5 μm in prokaryotes), allowing for greater cellular complexity. The origins of sexual reproduction in early eukaryotes, likely arising around 2.0 bya in their last common ancestor, introduced meiosis and genetic recombination, enhancing diversity and adaptability in response to environmental stresses like rising oxygen levels post-GOE. This process evolved from prokaryotic gene transfer mechanisms but incorporated fusion of haploid cells to form diploid zygotes, a trait nearly universal among eukaryotes and crucial for repairing DNA damage from oxidative stress. These unicellular and early eukaryotic innovations set the stage for the evolutionary lineage leading to human ancestry by fostering genetic variability and energy efficiency in an increasingly oxygenated world.

Emergence of Multicellular Animals

The emergence of multicellular animals, or metazoans, marked a pivotal transition in evolutionary history, building upon the foundational capabilities of unicellular eukaryotes that had developed complex internal structures and energy production mechanisms hundreds of millions of years earlier. This shift toward multicellularity enabled cells to specialize and cooperate, laying the groundwork for the diverse body plans seen in animal lineages ancestral to humans. Fossil and molecular evidence indicates that the first multicellular animals appeared between approximately 1.0 and 0.6 billion years ago (bya), with soft-bodied forms proliferating during the Ediacaran Period (635–541 million years ago, mya). Key innovations facilitating this transition included the evolution of cell adhesion molecules, such as cadherins, which allowed stable attachments between cells, and mechanisms for cell differentiation that enabled functional specialization within aggregates. These developments likely arose from pre-existing eukaryotic genes involved in cell signaling and cytoskeletal regulation, permitting the formation of tissues and rudimentary organs. Additionally, the emergence of Hox genes—transcription factors that pattern anterior-posterior body axes—provided a genetic toolkit for organizing complex morphologies, with evidence suggesting their origins in the common ancestor of all animals shortly after the advent of multicellularity. The Ediacaran biota, dating from about 575 to 541 mya, represents the earliest widespread assemblage of these soft-bodied multicellular forms, including disk-like and frond-shaped organisms preserved in situ on ancient seabeds. Recent fossil discoveries from Ediacaran sites in China have further illuminated precursor stages of multicellularity, including embryo-like holozoan fossils from the Weng'an biota (~609 mya) that exhibit cellular division patterns suggestive of early metazoan development, and 2024 reports of enigmatic sail-shaped structures from Yunnan Province indicating diverse soft-bodied experimentation before the Cambrian. The Cambrian Explosion, beginning around 541 mya, saw a rapid diversification of multicellular animals, characterized by the evolution of mineralized hard parts like shells and exoskeletons, which enhanced protection and predation capabilities among early metazoans. This event introduced major animal phyla, including Porifera (sponges), the earliest diverging group with fossil biomarkers and spicule evidence dating back over 650 mya; Cnidaria (such as jellyfish and corals), with molecular and trace fossil indications of radial symmetry emerging by the late Ediacaran; and early bilaterians, exemplified by the mollusk-like Kimberella (~555 mya), which displayed bilateral symmetry and possible grazing behavior. Environmental factors played a crucial role in driving these innovations, particularly the aftermath of Snowball Earth glaciations (~720–635 mya and ~650–635 mya), which created nutrient-scarce, viscous ocean conditions that favored multicellular aggregates for efficient resource acquisition over solitary cells. Concurrently, rising oceanic oxygenation levels around 600 mya provided the metabolic support necessary for larger, more active multicellular forms, alleviating oxygen diffusion limitations in cell clusters and enabling the metabolic demands of differentiation and movement. These pressures culminated in the metazoan radiation, setting the evolutionary stage for the vertebrate lineages that would eventually lead to humans.

Vertebrate and Tetrapod Development

Chordates and Early Vertebrates

The phylum Chordata encompasses a diverse group of animals characterized by the presence of a notochord, dorsal hollow nerve cord, pharyngeal slits, post-anal tail, and endostyle at some stage of development, marking a pivotal stage in the evolution toward vertebrates. Fossil evidence indicates that chordates first appeared during the Cambrian period around 530 million years ago (mya), coinciding with the diversification of early multicellular animals and the emergence of these defining anatomical features that supported filter-feeding lifestyles in marine environments. Early chordates, such as those represented by fossils like Pikaia gracilis, exhibited a flexible notochord for basic locomotion and pharyngeal slits for suspension feeding, laying the groundwork for subsequent evolutionary innovations within the phylum. Chordates are divided into three major subphyla: Cephalochordata (lancelets), Urochordata (tunicates), and Vertebrata, with vertebrates representing the most speciose and morphologically complex group that dominated aquatic ecosystems through much of evolutionary history. Lancelets, such as Branchiostoma, retain chordate characteristics throughout adulthood and serve as models for ancestral forms, while tunicates display these traits primarily in their larval stages before undergoing metamorphosis into sessile adults. Vertebrates, emerging later, replaced the notochord with a vertebral column and diversified extensively, with fish-like forms achieving ecological dominance in marine and freshwater habitats from the Ordovician onward, outcompeting many non-chordate invertebrates through superior mobility and feeding efficiency. Key evolutionary transitions within chordates included the origin of neural crest cells, the development of jaws in gnathostomes, and the appearance of paired fins, each enhancing vertebrate capabilities for active life. Neural crest cells, unique to vertebrates, arose around 500 mya as migratory cells derived from the dorsal neural tube, contributing to the formation of peripheral nerves, sensory structures, and craniofacial elements that distinguished early vertebrates from invertebrate chordates. Jaws evolved in gnathostomes approximately 420 mya during the Silurian-Devonian boundary, transforming the anterior pharyngeal arches into hinged structures that enabled biting and predation, a shift from the jawless filter-feeding of agnathans. Paired fins, first evident in early jawed fishes around 420-400 mya, originated from lateral plate mesoderm and provided stability and maneuverability for swimming, precursors to tetrapod limbs without implying terrestrial adaptation. Recent genomic studies have strengthened the phylogenetic links between tunicates and vertebrates, confirming their close relationship as sister groups within Chordata. These findings, building on whole-genome sequencing, highlight conserved genetic modules for pharyngeal and neural structures that predate vertebrate-specific innovations. Early vertebrates, particularly jawless agnathans appearing around 480 mya, developed adaptations for active predation, including enhanced sensory systems that improved detection and pursuit of prey in dimly lit aquatic environments. These included lateral line organs for sensing water movements, rudimentary eyes for phototaxis, and electroreceptive ampullae for navigating murky waters, all facilitated by neural crest-derived components that heightened responsiveness to environmental cues. Such sensory advancements, coupled with muscular enhancements from the notochord and early vertebral elements, enabled a transition from passive filter-feeding to opportunistic hunting, driving the radiation of vertebrate lineages during the Devonian period.

Transition to Tetrapods

The transition from aquatic vertebrates to land-dwelling tetrapods marked a pivotal evolutionary milestone during the Late Devonian period, approximately 375 to 360 million years ago (mya), enabling the invasion of terrestrial environments by vertebrate life. This shift is exemplified by transitional forms known as "fishapods," such as Tiktaalik roseae, discovered in the Canadian Arctic on Ellesmere Island, which possessed robust pectoral fins with internal bones homologous to tetrapod limbs, a flexible neck, and a flat skull adapted for shallow-water navigation. By around 365 mya, more advanced tetrapods like Acanthostega emerged, featuring digits and limb girdles capable of supporting weight outside water, though these early forms retained gills and fin-like tails indicative of their aquatic origins. These fossils illustrate a gradual fin-to-limb evolution, with sarcopterygian fish serving as precursors through their lobed fins reinforced by endochondral bones. Key adaptations facilitated this terrestrial transition, including the modification of swim bladders into functional lungs for air breathing, allowing survival in oxygen-poor swamp waters. Sturdy limbs evolved from pectoral and pelvic fins, with polydactylous (many-toed) patterns in early tetrapods providing enhanced propulsion and weight-bearing capacity on mudflats. Dermal bones, such as those forming robust rib cages and skull reinforcements, offered structural support against gravity and desiccation stresses, building on the endoskeletal foundations of earlier vertebrates. Sensory shifts, including robust spiracles for improved olfaction and audition in air, further aided navigation in emergent habitats. The environmental context involved the colonization of freshwater rivers and coastal swamps amid rising sea levels and the spread of early vascular plants, which stabilized sediments and boosted atmospheric oxygen to about 15-20% by the mid-Devonian. This greening of continents created niches for finned vertebrates to venture onto land for foraging or escaping predators, though many early tetrapods remained semi-aquatic. A notable gap in the fossil record, known as Romer's gap (approximately 360-345 mya), follows the initial tetrapod appearance, characterized by scarce remains possibly due to taphonomic biases in non-marine deposits or a temporary decline in diversity post-Devonian mass extinction events. Recent paleontological work in Arctic Canada has refined timelines for the fin-to-limb transition; for instance, the 2022 description of Qikiqtania wakei from Nunavut, dated to about 375 mya, reveals a closely related elpistostegalian with paddle-like fins suited for swimming rather than walking, highlighting evolutionary experimentation and mosaic trait acquisition within the group. Similarly, reanalysis of Elpistostege watsoni specimens from the same region in 2020 demonstrated a wrist-like radial bone structure in its pectoral fin, bridging fish and tetrapod forelimbs more precisely. By the early Carboniferous as early as ~355 million years ago, tetrapod lineages diversified into amniotes, egg-laying vertebrates fully adapted to land with waterproof skin and amniotic membranes, representing a critical offshoot toward drier terrestrial habitats.

Mammalian and Primate Lineage

Origin and Diversification of Mammals

The origins of mammals trace back to the synapsid lineage, which diverged from other amniotes during the late Carboniferous period around 312 million years ago (mya), with early representatives known as pelycosaurs exhibiting basal features such as a single temporal fenestra in the skull for enhanced jaw musculature. These primitive synapsids, including forms like Dimetrodon, dominated terrestrial ecosystems in the early Permian but lacked many derived mammalian characteristics. By approximately 275 mya, in the late Carboniferous to early Permian, therapsids emerged as a more advanced synapsid group, showing progressive mammalian-like adaptations such as differentiated teeth and more efficient locomotion, setting the stage for mammalian evolution. True mammals, defined by features like a mammalian jaw joint and ear structure, first appeared in the late Triassic around 225 mya, as evidenced by fossils like Brasilodon from Brazil, which display advanced dental patterns indicative of a fully mammalian dentition. Key traits that distinguished early mammals from their synapsid ancestors included endothermy, enabling internal heat regulation and active lifestyles; the development of hair or fur for insulation; mammary glands for nourishing offspring with milk; and the reconfiguration of the middle ear to three ossicles (malleus, incus, and stapes), derived from reptilian jaw bones (quadrate and articular) that detached to improve auditory sensitivity. Endothermy likely evolved stepwise in therapsids during the Permian, supported by bone histology showing rapid growth rates, while hair follicles may have originated as early as the late Permian in cynodont therapsids, inferred from skin impressions with follicular pits. Mammary glands arose from modified apocrine sweat glands associated with hair follicles, providing a proto-lactation system for moisture and antimicrobial secretions before true milk production in the Triassic. The middle ear evolution involved the migration of postdentary elements from the jaw to the ear region over the Permian-Triassic transition, enhancing high-frequency hearing crucial for small, nocturnal mammals. During the Mesozoic era, mammals remained small and ecologically marginal, overshadowed by dinosaurs, but underwent initial diversification with groups like multituberculates—rodent-like herbivores with specialized teeth for grinding plants, persisting from the Jurassic to Eocene—and monotremes, egg-laying mammals whose origins trace to the late Triassic or early Jurassic, as indicated by bone microstructure in fossils like Kryoryctes suggesting semiaquatic adaptations. Other groups, such as docodontans, also diversified in the Jurassic, with the oldest known specimen, Nujalikodon cassiopeiae from Greenland (~200 mya, as of 2025), indicating early presence in high-latitude environments. Multituberculates achieved notable diversity, occupying niches from insectivory to herbivory across Laurasia and Gondwana. The Cretaceous-Paleogene (K-Pg) extinction event around 66 mya, which eliminated non-avian dinosaurs, triggered a rapid mammalian radiation, allowing survivors to exploit vacant ecological roles and leading to the proliferation of larger body sizes and varied diets in the Paleocene. Within this context, the divergence between placental (eutherian) and marsupial (metatherian) lineages occurred around 160 mya in the Middle Jurassic, as evidenced by the eutherian fossil Juramaia sinensis from China, which exhibits a mosaic of primitive and derived traits bridging early therians. This split marked a pivotal point in mammalian reproductive evolution, with placentals developing extended gestation and marsupials emphasizing pouch-based development.

Evolution of Primates

The evolution of primates began in the aftermath of the Cretaceous-Paleogene extinction event approximately 66 million years ago (mya), when early mammal lineages diversified to occupy newly available ecological niches. Plesiadapiforms, considered stem primates or close relatives, emerged during the Paleocene epoch (66–55 mya) and exhibited proto-primate traits such as forward-directed orbits and specialized dentition for grasping and processing food, though they lacked key defining features like nails. These arboreal mammals bridged the gap between non-primate euarchontoglires and crown primates, adapting to forested environments through enhanced manual dexterity. True primates, or euprimates, appeared in the early Eocene around 55 mya, coinciding with global warming and the expansion of angiosperm-dominated forests. This radiation marked the origin of the order Primates, characterized by hallmark adaptations that facilitated life in complex arboreal settings. Forward-facing eyes enabled stereoscopic vision for depth perception during leaping and foraging, while grasping hands and feet with opposable digits allowed precise manipulation of branches and food items. Enlarged brain sizes relative to body mass supported improved sensory integration and problem-solving, and flat nails replaced claws, enhancing grip and grooming behaviors. These traits collectively positioned primates as adept exploiters of three-dimensional habitats, distinct from other mammalian orders. The primate lineage diverged into two major suborders, Strepsirrhini (wet-nosed primates, including lemurs, lorises, and galagos) and Haplorhini (dry-nosed primates, encompassing tarsiers and anthropoids), around 71 million years ago. Strepsirrhines retained more primitive features, such as a rhinarium and nocturnal habits, reflecting early primate ancestry. Haplorhines, in contrast, showed advanced visual acuity and reduced olfaction, setting the stage for further specialization. Within haplorhines, anthropoids (monkeys, apes, and humans) diverged from tarsier-like ancestors around 60–65 mya, though crown haplorhines are estimated at 66.2 mya based on molecular clocks. Anthropoid diversification accelerated in the Oligocene (34–23 mya), with the split between New World monkeys (Platyrrhini) and Old World monkeys plus apes (Catarrhini) occurring around 42–43 mya, as revised by 2024 genomic analyses incorporating expanded reference genomes. This divergence likely followed the rafting of platyrrhine ancestors across the Atlantic from African catarrhines, enabling independent radiations in South American forests. Old World lineages, including cercopithecoids (Old World monkeys) and hominoids (apes), adapted to diverse Old World habitats. Early anthropoids shifted from predominantly nocturnal lifestyles to diurnal activity, enhancing color vision for detecting ripe fruits and reducing reliance on insectivory in favor of frugivory. This dietary transition, supported by enhanced trichromatic vision in some lineages, drove further ecological specialization and brain expansion.

Hominid and Human Emergence

Hominidae Family and Early Hominins

The Hominidae family, encompassing great apes and humans, originated in Africa during the Miocene epoch approximately 14 to 7 million years ago, evolving from earlier ape-like ancestors adapted to forested environments. Genetic and fossil evidence indicates that the lineage diverged from orangutans around 14 million years ago, followed by the split from gorillas approximately 8.8 million years ago, and from chimpanzees and bonobos between 6.5 and 5.7 million years ago. These divergences marked the emergence of the African hominid clade, characterized by adaptations such as increased body size and flexible shoulder joints suited for brachiation, building on the grasping limb heritage from earlier primates. A defining trait of early hominins within Hominidae was the evolution of bipedalism, first evidenced in Sahelanthropus tchadensis around 7 million years ago through cranial features like a forward-positioned foramen magnum and postcranial elements such as a femur indicating upright locomotion. Additional hominin characteristics included reduced canine teeth, reflecting decreased male-male aggression and a shift toward omnivory, alongside larger molars with thick enamel for processing tough, fibrous vegetation. These dental changes appeared early in the hominin lineage, distinguishing it from other great apes with larger canines for display and tearing. Subsequent genera illustrate the diversification of bipedal hominins. Orrorin tugenensis, dated to about 6 million years ago in Kenya, shows femoral morphology consistent with partial bipedalism, including a thickened cortex and shortened femoral neck for weight-bearing during upright walking. Ardipithecus, spanning 5.8 to 4.4 million years ago in Ethiopia, combined bipedal adaptations in the pelvis and feet with arboreal traits like opposable big toes, suggesting a mosaic lifestyle in woodland settings. The genus Australopithecus, from 4 to 2 million years ago, further refined bipedalism; A. afarensis, exemplified by the 3.2-million-year-old "Lucy" skeleton from Ethiopia, possessed a curved spine, angled femur, and arched foot for efficient terrestrial travel, while retaining climbing capabilities. Ecologically, these hominins adapted to expanding savannas during Miocene-Pliocene climate shifts, where decreasing forest cover favored bipedalism for efficient long-distance travel and foraging across open grasslands. This habitat transition likely promoted dietary flexibility, with tool precursors emerging by 3.4 million years ago in A. afarensis, as indicated by cut marks on animal bones suggesting use of sharp stones for scavenging meat and processing plants.

Genus Homo and Archaic Humans

The genus Homo emerged in Africa around 2.8 million years ago, marking a pivotal shift in human evolution with the development of more advanced tool use and increased brain size compared to earlier hominins. Recent discoveries as of 2025, including fossils from Ethiopia's Afar region dated 2.6–3 million years ago, indicate co-existence of Australopithecus and early Homo species. The earliest species, Homo habilis, dated to approximately 2.3–1.4 million years ago, is recognized as the first known toolmaker, associated with the Oldowan stone tool industry, which consisted of simple choppers and flakes used for processing food. Fossils from sites like Olduvai Gorge in Tanzania reveal a brain capacity ranging from 500 to 900 cubic centimeters, larger than that of contemporaneous australopiths, alongside a more rounded skull and reduced facial prognathism. These traits suggest enhanced cognitive abilities, though H. habilis retained a body size and limb proportions similar to earlier bipedal ancestors. Following H. habilis, Homo erectus appeared around 1.9 million years ago and persisted until about 110,000 years ago, representing a major adaptive leap with brain sizes expanding to 600–1,100 cubic centimeters. This species is credited with inventing the Acheulean tool tradition, featuring symmetrical bifacial handaxes and cleavers that required greater planning and skill, likely aiding in butchery and woodworking. Evidence for controlled fire use by H. erectus dates to approximately 1 million years ago, as indicated by burned sediments and hearths at sites like Wonderwerk Cave in South Africa, which facilitated cooking, warmth, and predator deterrence. H. erectus was the first hominin to migrate out of Africa, with fossils in Dmanisi, Georgia, dated to 1.8 million years ago, and subsequent dispersals reaching as far as Java and China by 1.6 million years ago, driven by environmental changes and resource exploitation. Later archaic humans within the genus Homo diversified further, with Homo heidelbergensis emerging around 700,000–200,000 years ago in Africa and Europe, exhibiting even larger brains (up to 1,400 cubic centimeters) and robust builds adapted to varied climates. This species refined Acheulean tools and showed evidence of hunting large game, bridging earlier Homo lineages to later forms. Neanderthals (Homo neanderthalensis), evolving around 400,000–40,000 years ago primarily in Europe and western Asia, displayed advanced adaptations including large nasal cavities for cold air humidification and cultural behaviors like burial practices. Denisovans, known from genetic and limited fossil evidence dated to 200,000–50,000 years ago in Asia, shared a common ancestor with Neanderthals around 400,000 years ago and exhibited high-altitude adaptations, such as the EPAS1 gene variant aiding Tibetan populations today. Genomic analyses indicate Denisovan interbreeding extended to Southeast Asia, with up to 5% Denisovan ancestry in modern populations like the Ayta Magbukon in the Philippines, reflecting multiple admixture waves. Multiple waves of Homo dispersal out of Africa shaped archaic human distributions, beginning with H. erectus around 1.8 million years ago via the Levantine corridor and southern routes, followed by later migrations of H. heidelbergensis and Neanderthal ancestors into Eurasia by 700,000 years ago. Genomic evidence reveals interbreeding between these archaic groups and early modern humans, with non-African populations carrying 1–2% Neanderthal DNA from events around 50,000–60,000 years ago, and higher Denisovan contributions (up to 6%) in East Asian and Oceanian groups from encounters in Asia. Such admixture provided adaptive advantages, including immune system enhancements and metabolic adjustments, while highlighting the reticulated nature of human evolution.

Modern Homo sapiens

Anatomically modern Homo sapiens emerged in Africa around 300,000 years ago, with the earliest known fossils discovered at Jebel Irhoud in Morocco, dated to approximately 315,000 years old through thermoluminescence dating of associated artifacts and sediments. These early individuals exhibit a mix of modern and archaic features, including a globular braincase and facial morphology transitional toward the fully modern form characterized by a high forehead, rounded skull vault, prominent chin, and reduced brow ridges. The average brain volume of modern Homo sapiens is approximately 1,350–1,400 cm³, larger than that of earlier hominins and enabling advanced cognitive capacities. Evidence of behavioral modernity, including symbolic art, complex tools, and ornaments, first appears in Africa around 100,000 years ago, with further developments evident during the Upper Paleolithic in Eurasia around 50,000 years ago, coinciding with innovations like blade technologies and cave paintings that reflect abstract thinking and social complexity. Between approximately 70,000 and 50,000 years ago, small groups of Homo sapiens migrated out of Africa, dispersing across Eurasia and eventually replacing or assimilating archaic populations such as Neanderthals through a combination of demographic expansion, competitive advantages, and interbreeding events. Genetic evidence indicates that non-African populations carry 1–2% Neanderthal DNA from these admixture episodes, which occurred primarily in Eurasia around 50,000–60,000 years ago, contributing adaptive alleles for immunity and skin pigmentation. This Out-of-Africa expansion involved genetic bottlenecks, reducing effective population sizes and leading to decreased diversity in non-African genomes compared to African ones, with founder effects shaping regional variations. Further dispersals reached Australia by about 65,000 years ago and the Americas around 15,000 years ago via Beringia, facilitated by coastal and inland routes during the Late Pleistocene. The Neolithic Revolution, beginning around 12,000 years ago in the Fertile Crescent, marked a pivotal shift as Homo sapiens transitioned from foraging to agriculture and animal domestication, enabling population growth and sedentary societies. This period also saw ongoing evolution, including adaptations like lactase persistence in pastoralist populations, driven by positive selection on mutations in the LCT gene that allowed adult milk digestion, emerging independently in Europe, Africa, and Asia within the last 10,000 years. Recent genomic studies highlight additional archaic influences, such as elevated Denisovan admixture in Oceanian populations—up to 4–6% in some Papuan and Australian groups—revealed through 2025 analyses of genes like MUC19, which show recurrent introgression events potentially aiding adaptations to tropical environments. These findings underscore the dynamic interplay of migration, admixture, and natural selection in shaping contemporary human diversity.

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