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''Homo'' () is the
genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circumscription (taxonomy), circumscribing) and classifying gr ...
that emerged in the (otherwise extinct) genus ''
Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of ...

Australopithecus
'' that encompasses the extant species ''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread of , characterized by and large, complex brains. This has enabled the development of advanced , , and . Humans are highly social and tend to live in complex s composed of many ...

Homo sapiens
'' (
modern humans Early modern human (EMH) or anatomically modern human (AMH) are terms used to distinguish ''Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread of , characterized by and large, complex brains. This has enabled ...
), plus several extinct species classified as either
ancestral An ancestor, also known as a forefather, fore-elder or a forebear, is a parent A parent is a caregiver of the offspring In biology, offspring are the young born of living organism, organisms, produced either by a single organism or, in the ...
to or closely related to modern humans (depending on the species), most notably ''
Homo erectus ''Homo erectus'' (meaning "upright Body relative directions (also known as egocentric coordinates) are geometrical orientations relative to a body such as a human Humans (''Homo sapiens'') are the most populous and widespread spec ...

Homo erectus
'' and ''
Homo neanderthalensis Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an org ...

Homo neanderthalensis
''. The genus emerged with the appearance of ''
Homo habilis ''Homo habilis'' ("handy man") is a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the large ...

Homo habilis
'' just over 2 million years ago. ''Homo'', together with the genus ''
Paranthropus ''Paranthropus'' is a genus of extinct hominin which contains two widely accepted species: '' P. robustus'' and '' P. boisei''. However, the validity of ''Paranthropus'' is contested, and it is sometimes considered to be synonym (taxonomy), sy ...
'', is probably sister to ''
Australopithecus africanus ''Australopithecus africanus'' is an extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the endling, last individ ...

Australopithecus africanus
'', which itself had previously split from the lineage of ''
Pan Pan may refer to: Prefix *''Pan-'', a prefix A prefix is an affix which is placed before the stem of a word. Adding it to the beginning of one word changes it into another word. For example, when the prefix ''un-'' is added to the word ''happy ...
'', the
chimpanzees The chimpanzee (''Pan troglodytes''), also known simply as chimp, is a species of Hominidae, great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. The chimpanzee and ...

chimpanzees
. ''
Homo erectus ''Homo erectus'' (meaning "upright Body relative directions (also known as egocentric coordinates) are geometrical orientations relative to a body such as a human Humans (''Homo sapiens'') are the most populous and widespread spec ...

Homo erectus
'' appeared about 2 million years ago and, in several early migrations, spread throughout Africa (where it is dubbed ''
Homo ergaster ''Homo ergaster'' is an extinct species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular int ...

Homo ergaster
'') and Eurasia. It was likely the first human species to live in a
hunter-gatherer A hunter-gatherer is a human Humans (''Homo sapiens'') are the most populous and widespread species of primates, characterized by bipedality, opposable thumbs, hairlessness, and intelligence allowing the use of culture, language and tools. T ...
society and to control fire. An adaptive and successful species, ''Homo erectus'' persisted for more than a million years and gradually diverged into new species by around 500,000 years ago. ''Homo sapiens'' (
anatomically modern humans Early modern human (EMH) or anatomically modern human (AMH) are terms used to distinguish ''Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread of , characterized by and large, complex brains. This has enabled ...
) emerged close to 300,000 to 200,000 years ago, most likely in Africa, and ''
Homo neanderthalensis Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an org ...

Homo neanderthalensis
'' emerged at around the same time in Europe and Western Asia. ''H. sapiens'' dispersed from Africa in several waves, from possibly as early as 250,000 years ago, and certainly by 130,000 years ago, the so-called
Southern Dispersal In the context of the recent African origin of modern humans, the Southern Dispersal scenario (also the coastal migration or great coastal migration hypothesis) refers to the early human migrations, early migration along the southern coast of Asi ...
beginning about 70–50,000 years ago leading to the lasting colonisation of
Eurasia Eurasia () is the largest continent A continent is any of several large landmasses. Generally identified by convention (norm), convention rather than any strict criteria, up to seven geographical regions are commonly regarded as cont ...

Eurasia
and
Oceania Oceania (, , ) is a geographic region In geography Geography (from Greek: , ''geographia'', literally "earth description") is a field of science devoted to the study of the lands, features, inhabitants, and phenomena of the Eart ...

Oceania
by 50,000 years ago. Both in Africa and Eurasia, ''H. sapiens'' met with and interbred with
archaic humans A number of varieties of ''Homo ''Homo'' () is the genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classif ...
. Separate archaic (non-''sapiens'') human species are thought to have survived until around 40,000 years ago (
Neanderthal extinction Neanderthals Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ago. They most likely went ex ...
), with possible late survival of
hybrid species Hybrid speciation is a form of speciation Speciation is the evolution Evolution is change in the Heredity, heritable Phenotypic trait, characteristics of biological populations over successive generations. These characteristics are the G ...
as late as 12,000 years ago ( Red Deer Cave people).


Names and taxonomy

The
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as Latium. Through the power of the Roman Republic, it became ...

Latin
noun ''homō'' (genitive ''hominis'') means "human being" or "
man A man is an adult male Male (♂) is the sex of an organism that produces the gamete known as sperm. A male gamete can fuse with a larger female gamete, or ovum, in the process of fertilization. A male cannot sexual reproduction, reproduc ...
" in the generic sense of "human being, mankind". The
binomial name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed ...
''Homo sapiens'' was coined by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement Ennoblement is the conferring of nobility—the induction of an individual into the noble social class, class. Currently only a few kingdoms still grant nob ...

Carl Linnaeus
(1758). Names for other species of the genus were introduced beginning in the second half of the 19th century (''H. neanderthalensis'' 1864, ''H. erectus'' 1892). Even today, the genus ''Homo'' has not been strictly defined. Since the early human fossil record began to slowly emerge from the earth, the boundaries and definitions of the genus ''Homo'' have been poorly defined and constantly in flux. Because there was no reason to think it would ever have any additional members,
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement Ennoblement is the conferring of nobility—the induction of an individual into the noble social class, class. Currently only a few kingdoms still grant nob ...

Carl Linnaeus
did not even bother to define ''Homo'' when he first created it for humans in the 18th century. The discovery of Neanderthal brought the first addition. The genus ''Homo'' was given its taxonomic name to suggest that its member species can be classified as human. And, over the decades of the 20th century, fossil finds of pre-human and early human species from late
Miocene The Miocene ( ) is the first geological epoch In geochronology, an epoch is a subdivision of the geologic timescale that is longer than an age (geology), age but shorter than a period (geology), period. The current epoch is the Holocene Epoch of ...
and early
Pliocene The Pliocene ( ; also Pleiocene) epoch (geology), Epoch is the epoch in the geologic timescale that extends from 5.333 million to 2.58Pan Pan may refer to: Prefix *''Pan-'', a prefix A prefix is an affix which is placed before the stem of a word. Adding it to the beginning of one word changes it into another word. For example, when the prefix ''un-'' is added to the word ''happy ...
'', as one body of scientists argues that the two species of chimpanzee should be classed with genus ''Homo'' rather than ''Pan''. Even so, classifying the fossils of ''Homo'' coincides with evidence of: (1) competent human bipedalism in ''
Homo habilis ''Homo habilis'' ("handy man") is a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the large ...

Homo habilis
'' inherited from the earlier ''
Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of ...

Australopithecus
'' of more than four million years ago, as demonstrated by the
Laetoli footprints Laetoli is a site in Tanzania, dated to the Plio-Pleistocene and famous for its hominin footprints, preserved in volcanic ash. The site of the Laetoli footprints (Site G) is located 45 km south of Olduvai gorge. The location and tracks were di ...

Laetoli footprints
; and (2)  having begun by 2.5 million years ago. From the late-19th to mid-20th centuries, a number of new taxonomic names including new generic names were proposed for early human fossils; most have since been merged with ''Homo'' in recognition that ''
Homo erectus ''Homo erectus'' (meaning "upright Body relative directions (also known as egocentric coordinates) are geometrical orientations relative to a body such as a human Humans (''Homo sapiens'') are the most populous and widespread spec ...

Homo erectus
'' was a single species with a large geographic spread of early migrations. Many such names are now dubbed as "
synonyms A synonym is a word, morpheme A morpheme is the smallest meaningful lexical item in a language. A morpheme is not a word. The difference between a morpheme and a word is that a morpheme bound and free morphemes, sometimes does not stand alone ...
" with ''Homo'', including ''Pithecanthropus'', ''Protanthropus'', ''Sinanthropus'', ''Cyphanthropus'', ''Africanthropus'', ''Telanthropus'', ''Atlanthropus'', and ''Tchadanthropus''. Classifying the genus ''Homo'' into species and subspecies is subject to incomplete information and remains poorly done. This has led to using common names ("Neanderthal" and "Denisovan"), even in scientific papers, to avoid trinomial names or the ambiguity of classifying groups as ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer to a specific ...
'' (uncertain placement)—for example, ''H. neanderthalensis'' vs. ''H. sapiens neanderthalensis'', or ''H. georgicus'' vs. ''H. erectus georgicus''. Some recently extinct species in the genus ''Homo'' have only recently been discovered and do not as yet have consensus binomial names (see
Denisova hominin The Denisovans or Denisova hominins ) are an extinct species or subspecies of archaic human that ranged across Asia during the Lower Paleolithic, Lower and Middle Paleolithic. Denisovans are known from few remains, and, consequently, most of wha ...
and Red Deer Cave people). Since the beginning of the
Holocene The Holocene ( ) is the current geological epoch In geochronology, an epoch is a subdivision of the geologic timescale that is longer than an age (geology), age but shorter than a period (geology), period. The current epoch is the Holocene E ...
, it is likely that ''Homo sapiens'' (anatomically modern humans) has been the only extant species of ''Homo''.
John Edward Gray John Edward Gray, Fellow of the Royal Society, FRS (12 February 1800 – 7 March 1875) was a British zoology, zoologist. He was the elder brother of zoologist George Robert Gray and son of the pharmacologist and botanist Samuel Frederick G ...
(1825) was an early advocate of classifying taxa by designating tribes and families. Wood and Richmond (2000) proposed that
Hominini The Hominini form a taxonomic tribe of the subfamily Homininae Homininae (), also called "African hominids" or "African apes", is a subfamily of Hominidae. It includes two tribes, with their extant taxon, extant as well as extinct species: 1 ...

Hominini
("hominins") be designated as a
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant usage of the term is in the discipline of anthropology. The definition is contested, in part due to conflicting theoretical understa ...
that comprised all species of early humans and pre-humans ancestral to humans back to ''after'' the chimpanzee-human last common ancestor; and that Hominina be designated a subtribe of Hominini to include ''only'' the genus ''Homo'' — that is, ''not'' including the earlier upright walking hominins of the
Pliocene The Pliocene ( ; also Pleiocene) epoch (geology), Epoch is the epoch in the geologic timescale that extends from 5.333 million to 2.58Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of ...

Australopithecus
'', ''
Orrorin tugenensis ''Orrorin tugenensis'' is a postulated early species of Homininae, estimated at and discovered in 2000. It is not confirmed how ''Orrorin'' is related to Anatomically modern humans, modern humans. Its discovery was used to argue against the hypot ...

Orrorin tugenensis
'', ''
Ardipithecus ''Ardipithecus'' is a genus of an extinct hominine that lived during the Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Mess ...
'', or ''
Sahelanthropus ''Sahelanthropus tchadensis'' is an extinct species of the Homininae (African apes) dated to about , during the Miocene The Miocene ( ) is the first Epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocen ...
''. Designations alternative to Hominina existed, or were offered: ''Australopithecinae'' (Gregory & Hellman 1939) and ''Preanthropinae'' (Cela-Conde & Altaba 2002); and later, Cela-Conde and Ayala (2003) proposed that the four genera ''Australopithecus'', ''Ardipithecus'', ''Praeanthropus'', and ''Sahelanthropus'' be grouped with ''Homo'' within
Hominini The Hominini form a taxonomic tribe of the subfamily Homininae Homininae (), also called "African hominids" or "African apes", is a subfamily of Hominidae. It includes two tribes, with their extant taxon, extant as well as extinct species: 1 ...

Hominini
(sans Pan).


Evolution


''Australopithecus'' and the appearance of ''Homo''

Several species, including ''
Australopithecus garhi ''Australopithecus garhi'' is a species of australopithecine from the Bouri Formation in the Afar Region of Ethiopia 2.6–2.5 million years ago (mya) during the Early Pleistocene. The first remains were described in 1999 based on several ske ...
'', ''
Australopithecus sediba ''Australopithecus sediba'' is an extinct species of australopithecine recovered from Malapa Fossil Site, Cradle of Humankind, Malapa Cave, Cradle of Humankind, South Africa. It is known from a partial juvenile skeleton, the holotype MH1, and a ...

Australopithecus sediba
'', ''
Australopithecus africanus ''Australopithecus africanus'' is an extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the endling, last individ ...

Australopithecus africanus
'', and ''
Australopithecus afarensis ''Australopithecus afarensis'' is an extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the endling, last individua ...

Australopithecus afarensis
'', have been proposed as the ancestor or sister of the ''Homo'' lineage. These species have morphological features that align them with ''Homo'', but there is no consensus as to which gave rise to ''Homo''. Especially since the 2010s, the delineation of ''Homo'' in ''Australopithecus'' has become more contentious. Traditionally, the advent of ''Homo'' has been taken to coincide with the first use of
stone tool A stone tool is, in the most general sense, any tool made either partially or entirely out of Rock (geology), stone. Although stone tool-dependent societies and cultures still exist today, most stone tools are associated with prehistory, prehisto ...

stone tool
s (the
Oldowan The Oldowan (or Mode I) was a widespread stone tool A stone tool is, in the most general sense, any tool made either partially or entirely out of Rock (geology), stone. Although stone tool-dependent societies and cultures still exist today, ...

Oldowan
industry), and thus by definition with the beginning of the
Lower Palaeolithic 250px, Four views of an Acheulean handaxe The Lower Paleolithic (or Lower Palaeolithic) is the earliest subdivision of the Paleolithic The Paleolithic or Palaeolithic or Palæolithic (), also called the Old Stone Age, is a period in human pr ...

Lower Palaeolithic
. But in 2010, evidence was presented that seems to attribute the use of stone tools to ''Australopithecus afarensis'' around 3.3 million years ago, close to a million years before the first appearance of ''Homo''. "The oldest direct evidence of stone tool manufacture comes from
Gona Gona is a coastal village in Oro Province Oro Province, formerly (and officially still) Northern Province, is a coast The coast, also known as the coastline or seashore, is defined as the area where land meets the sea or ocean, or as a line th ...
(Ethiopia) and dates to between 2.6 and 2.5 million years (Myr) ago. ..Here we report stone-tool-inflicted marks on bones found during recent survey work in Dikika, Ethiopia .. showingunambiguous stone-tool cut marks for flesh removal .., datedto between 3.42 and 3.24 Myr ago ..Our discovery extends by approximately 800,000 years the antiquity of stone tools and of stone-tool-assisted consumption of ungulates by hominins; furthermore, this behaviour can now be attributed to Australopithecus afarensis."
LD 350-1 LD 350-1 is the earliest known specimen of the genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circums ...
, a fossil mandible fragment dated to 2.8 Mya, discovered in 2013 in , was described as combining "primitive traits seen in early ''Australopithecus'' with derived morphology observed in later ''Homo''. Some authors would push the development of ''Homo'' close to or even past 3 Mya. Others have voiced doubt as to whether ''Homo habilis'' should be included in ''Homo'', proposing an origin of ''Homo'' with ''Homo erectus'' at roughly 1.9 Mya instead. The most salient physiological development between the earlier australopithecine species and ''Homo'' is the increase in endocranial volume (ECV), from about in ''A. garhi'' to in ''H. habilis'' and further to in ''H. erectus'', in ''H. heidelbergensis'' and up to in ''H. neanderthalensis''. However, a steady rise in cranial capacity is observed already in ''Autralopithecina'' and does not terminate after the emergence of ''Homo'', so that it does not serve as an objective criterion to define the emergence of the genus.


''Homo habilis''

''
Homo habilis ''Homo habilis'' ("handy man") is a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the large ...

Homo habilis
'' emerged about 2.1 Mya. Already before 2010, there were suggestions that ''H. habilis'' should not be placed in genus ''Homo'' but rather in ''Australopithecus''. The main reason to include ''H. habilis'' in ''Homo'', its undisputed tool use, has become obsolete with the discovery of ''Australopithecus'' tool use at least a million years before ''H. habilis''. Furthermore, ''H. habilis'' was long thought to be the ancestor of the more gracile ''
Homo ergaster ''Homo ergaster'' is an extinct species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular int ...

Homo ergaster
'' (''
Homo erectus ''Homo erectus'' (meaning "upright Body relative directions (also known as egocentric coordinates) are geometrical orientations relative to a body such as a human Humans (''Homo sapiens'') are the most populous and widespread spec ...

Homo erectus
''). In 2007, it was discovered that ''H. habilis'' and ''H. erectus'' coexisted for a considerable time, suggesting that ''H. erectus'' is not immediately derived from ''H. habilis'' but instead from a common ancestor. With the publication of
Dmanisi skull 5 The Dmanisi skull, also known as Skull 5 or D4500, is one of five skulls discovered in Dmanisi, Georgia (country), Georgia and classified as early ''Homo erectus''. Described in a publication in October 2013, it is estimated to be about 1.8 mill ...
in 2013, it has become less certain that Asian ''H. erectus'' is a descendant of African ''H. ergaster'' which was in turn derived from ''H. habilis''. Instead, ''H. ergaster'' and ''H. erectus'' appear to be variants of the same species, which may have originated in either Africa or Asia and widely dispersed throughout Eurasia (including
Europe Europe is a continent A continent is any of several large landmass A landmass, or land mass, is a large region In geography Geography (from Greek: , ''geographia'', literally "earth description") is a field of scienc ...
,
Indonesia Indonesia ( ), officially the Republic of Indonesia ( id, Republik Indonesia, links=yes ), is a country in Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia or SEA, is t ...

Indonesia
,
China China (), officially the People's Republic of China (PRC; ), is a country in East Asia East Asia is the eastern region of Asia Asia () is Earth's largest and most populous continent, located primarily in the Eastern Hemisphere ...
) by 0.5 Mya.


''Homo erectus''

''Homo erectus'' has often been assumed to have developed anagenetically from ''Homo habilis'' from about 2 million years ago. This scenario was strengthened with the discovery of '' Homo erectus georgicus'', early specimens of ''H. erectus'' found in the
Caucasus The Caucasus (), or Caucasia (), is a region spanning Europe and Asia. It is situated between the Black Sea and the Caspian Sea and mainly occupied by Armenia, Azerbaijan, Georgia (country), Georgia, and parts of Southern Russia. It is home to ...
, which seemed to exhibit transitional traits with ''H. habilis''. As the earliest evidence for ''H. erectus'' was found outside of Africa, it was considered plausible that ''H. erectus'' developed in Eurasia and then migrated back to Africa. Based on fossils from the
Koobi Fora Koobi Fora refers primarily to a region around Koobi Fora Ridge, located on the eastern shore of Lake Turkana Lake Turkana (), formerly known as Lake Rudolf, is a lake in the Kenyan Rift Valley, in northern Kenya, with its far northern en ...
Formation, east of Lake Turkana in Kenya, Spoor et al. (2007) argued that ''H. habilis'' may have survived beyond the emergence of ''H. erectus'', so that the evolution of ''H. erectus'' would not have been anagenetically, and ''H. erectus'' would have existed alongside ''H. habilis'' for about half a million years (), during the early Calabrian. A separate South African species ''
Homo gautengensis ''Homo gautengensis'' is a species name proposed by anthropologist Darren Curnoe in 2010 for South African hominin The Hominini form a Tribe (biology), taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant gener ...
'' has been postulated as contemporary with ''Homo erectus'' in 2010.


Phylogeny

A taxonomy of ''Homo'' within the
great apes The Hominidae (), whose members are known as great apes or hominids (), are a taxonomic family In human society A society is a Social group, group of individuals involved in persistent Social relation, social interaction, or a larg ...
is assessed as follows, with ''Paranthropus'' and ''Homo'' emerging within ''Australopithecus'' (shown here
cladistically Cladistics (; ) is an approach to Taxonomy (biology), biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically ...

cladistically
granting ''Paranthropus'', ''Kenyanthropus'', and ''Homo''). The exact phylogeny within ''Australopithecus'' is still highly controversial. Approximate radiation dates of daughter clades are shown in millions of years ago (Mya). ''
Graecopithecus ''Graecopithecus'' is a hominid originally identified by a single mandible found in Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 milli ...
,'' ''
Sahelanthropus ''Sahelanthropus tchadensis'' is an extinct species of the Homininae (African apes) dated to about , during the Miocene The Miocene ( ) is the first Epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocen ...
'', ''
Orrorin ''Orrorin tugenensis'' is a postulated early species of Homininae Homininae (), also called "African hominids" or "African apes", is a subfamily of Hominidae. It includes two tribes, with their extant taxon, extant as well as extinct species: 1 ...
'', possibly sisters to ''Australopithecus'', are not shown here. Note that the naming of groupings is sometimes muddled as often certain groupings are presumed before any cladistic analysis is performed. Several of the ''Homo'' lineages appear to have surviving progeny through introgression into other lines. Genetic evidence indicates an archaic lineage separating from the other human lineages 1.5 million years ago, perhaps ''H. erectus'', may have interbred into the Denisovans about 55,000 years ago. Fossil evidence shows ''Homo erectus'' s.s. survived at least until 117,000 yrs ago, and the even more basal ''Homo floresiensis'' survived until 50,000 years ago. Moreover, a thigh bone, dated at 14,000 years, found in a Maludong cave ( Red Deer Cave people) strongly resembles very ancient species like early ''Homo erectus'' or the even more archaic lineage, ''Homo habilis'', which lived around 1.5 million year ago. A 1.5 million years Homo erectus-like lineage appears to have made its way into modern humans through the Denisovans and specifically into the Papuans and aboriginal Australians. The genomes of non-sub-Saharan African humans show what appear to be numerous independent introgression events involving Neanderthal and in some cases also Denisovans around 45,000 years ago. Likewise the genetic structure of sub-Saharan Africans seems to be indicative of introgression from a west Eurasian population some 3,000 years ago. Some evidence suggests that ''
Australopithecus sediba ''Australopithecus sediba'' is an extinct species of australopithecine recovered from Malapa Fossil Site, Cradle of Humankind, Malapa Cave, Cradle of Humankind, South Africa. It is known from a partial juvenile skeleton, the holotype MH1, and a ...

Australopithecus sediba
'' could be moved to the genus ''Homo'', or placed in its own genus, due to its position with respect to e.g. ''
Homo habilis ''Homo habilis'' ("handy man") is a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the large ...

Homo habilis
'' and ''
Homo floresiensis ''Homo floresiensis'' ("Flores Man"; nicknamed "Hobbit") is a species of small archaic human A number of varieties of ''Homo ''Homo'' () is the that emerged in the (otherwise extinct) genus ' that encompasses the extant species ' (), p ...

Homo floresiensis
''.


Dispersal

By about 1.8 million years ago, ''Homo erectus'' is present in both East Africa (''
Homo ergaster ''Homo ergaster'' is an extinct species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular int ...

Homo ergaster
'') and in Western Asia (''
Homo georgicus The Dmanisi hominins, Dmanisi people or Dmanisi man were a population of Early Pleistocene The Early Pleistocene is an unofficial sub-epoch in the international geologic timescale The geologic time scale (GTS) is a system of chronological dat ...
''). The ancestors of Indonesian ''
Homo floresiensis ''Homo floresiensis'' ("Flores Man"; nicknamed "Hobbit") is a species of small archaic human A number of varieties of ''Homo ''Homo'' () is the that emerged in the (otherwise extinct) genus ' that encompasses the extant species ' (), p ...

Homo floresiensis
'' may have left Africa even earlier. ''Homo erectus'' and related or derived
archaic human A number of varieties of ''Homo ''Homo'' () is the that emerged in the (otherwise extinct) genus ' that encompasses the extant species ' (), plus several extinct species classified as either to or closely related to modern humans (dependin ...
species over the next 1.5 million years spread throughout Africa and Eurasia (see:
Recent African origin of modern humans In paleoanthropology Paleoanthropology or paleo-anthropology is a branch of paleontology and biological anthropology, anthropology which seeks to understand the early development of anatomically modern humans, a process known as wikt:hominiza ...
). Europe is reached by about 0.5 Mya by ''
Homo heidelbergensis ''Homo heidelbergensis'' (also ''H. sapiens heidelbergensis'') is an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversit ...

Homo heidelbergensis
''. ''
Homo neanderthalensis Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an org ...

Homo neanderthalensis
'' and ''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread of , characterized by and large, complex brains. This has enabled the development of advanced , , and . Humans are highly social and tend to live in complex s composed of many ...

Homo sapiens
'' develop after about 300 kya. ''
Homo naledi '' Homo naledi'' is a species of Archaic humans, archaic human discovered in 2013 in the Rising Star Cave, Cradle of Humankind, South Africa dating to the Middle Pleistocene 335,000–236,000 years ago. The initial discovery comprises 1,550 spe ...

Homo naledi
'' is present in Southern Africa by 300 kya. ''H. sapiens'' soon after its first emergence spread throughout Africa, and to Western Asia in several waves, possibly as early as 250 kya, and certainly by 130 kya. In July 2019, anthropologists reported the discovery of 210,000 year old remains of a ''H. sapiens'' and 170,000 year old remains of a ''H. neanderthalensis'' in Apidima Cave,
Peloponnese The Peloponnese (), Peloponnesia, or Peloponnesus (; el, Πελοπόννησος, Pelopónnēsos, ) is a peninsula and geographic regions of Greece, geographic region in southern Greece. It is connected to the central part of the country by the ...
,
Greece Greece ( el, Ελλάδα, Elláda, ), officially the Hellenic Republic, is a country located in Southeastern Europe Southeast Europe or Southeastern Europe () is a geographical subregion A subregion is a part of a larger region In geogr ...

Greece
, more than 150,000 years older than previous ''H. sapiens'' finds in Europe. Most notable is the
Southern Dispersal In the context of the recent African origin of modern humans, the Southern Dispersal scenario (also the coastal migration or great coastal migration hypothesis) refers to the early human migrations, early migration along the southern coast of Asi ...
of ''H. sapiens'' around 60 kya, which led to the lasting peopling of Oceania and Eurasia by
anatomically modern humans Early modern human (EMH) or anatomically modern human (AMH) are terms used to distinguish ''Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread of , characterized by and large, complex brains. This has enabled ...
. ''H. sapiens'' interbred with archaic humans both in Africa and in Eurasia, in Eurasia notably with
Neanderthals Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an org ...
and
Denisovans The Denisovans or Denisova hominins ) are an extinct Extinction is the termination of a kind of organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system ...
. Among extant populations of ''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread of , characterized by and large, complex brains. This has enabled the development of advanced , , and . Humans are highly social and tend to live in complex s composed of many ...

Homo sapiens
'', the deepest temporal division is found in the
San people The San peoples (also Saan), or Bushmen, are members of various Khoe Maharishi International University (MIU), formerly Maharishi University of Management, is a private university in Fairfield, Iowa. It was founded in 1973 by Maharishi Mahesh ...

San people
of Southern Africa, estimated at close to 130,000 years, or possibly more than 300,000 years ago. Temporal division among non-Africans is of the order of 60,000 years in the case of
Australo-Melanesians Australo-Melanesians (also known as Australasians or Australomelanesoid race or Australoid race) is an outdated historical grouping of various people indigenous to Melanesia Melanesia (, ) is a subregion of Oceania in the southwestern Pacific ...
. Division of
Europeans Europeans are the focus of European ethnology, the field of anthropology related to the various indigenous groups that reside in the List of sovereign states and dependent territories in Europe, nations of Europe. Groups may be defined by commo ...
and
East Asians East Asian people (East Asians) are the people from East Asia East Asia is the eastern region of Asia, which is defined in both Geography, geographical and culture, ethno-cultural terms. The modern State (polity), states of East Asia incl ...
is of the order of 50,000 years, with repeated and significant admixture events throughout Eurasia during the
Holocene The Holocene ( ) is the current geological epoch In geochronology, an epoch is a subdivision of the geologic timescale that is longer than an age (geology), age but shorter than a period (geology), period. The current epoch is the Holocene E ...
. Archaic human species may have survived until the beginning of the
Holocene The Holocene ( ) is the current geological epoch In geochronology, an epoch is a subdivision of the geologic timescale that is longer than an age (geology), age but shorter than a period (geology), period. The current epoch is the Holocene E ...
( Red Deer Cave people), although they were mostly extinct or absorbed by the expanding ''H. sapiens'' populations by 40 kya (
Neanderthal extinction Neanderthals Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ago. They most likely went ex ...
).


List of lineages

The species status of '' H. rudolfensis'', '' H. ergaster'', '' H. georgicus'', '' H. antecessor'', '' H. cepranensis'', '' H. rhodesiensis'', '' H. neanderthalensis'',
Denisova hominin The Denisovans or Denisova hominins ) are an extinct species or subspecies of archaic human that ranged across Asia during the Lower Paleolithic, Lower and Middle Paleolithic. Denisovans are known from few remains, and, consequently, most of wha ...
, Red Deer Cave people, and ''H. floresiensis'' remains under debate. ''H. heidelbergensis'' and ''H. neanderthalensis'' are closely related to each other and have been considered to be human subspecies, subspecies of ''H. sapiens''. There has historically been a trend to postulate new human species based on as little as an individual fossil. A "minimalist" approach to human taxonomy recognizes at most three species, ''
Homo habilis ''Homo habilis'' ("handy man") is a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the large ...

Homo habilis
'' (2.1–1.5 Mya, membership in ''Homo'' questionable), ''
Homo erectus ''Homo erectus'' (meaning "upright Body relative directions (also known as egocentric coordinates) are geometrical orientations relative to a body such as a human Humans (''Homo sapiens'') are the most populous and widespread spec ...

Homo erectus
'' (1.8–0.1 Mya, including the majority of the age of the genus, and the majority of archaic varieties as Homo erectus subspecies, subspecies, including ''H. heidelbergensis'' as a late or transitional variety) and ''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread of , characterized by and large, complex brains. This has enabled the development of advanced , , and . Humans are highly social and tend to live in complex s composed of many ...

Homo sapiens
'' (300 kya to present, including ''H. neanderthalensis'' and other varieties as human subspecies, subspecies). "Species" does in this context not necessarily mean that hybridization and introgression were impossible at the time. However, it is often used as a convenient term, but it should be taken to mean to be a generic lineage at best, and clusters at worst. In general definitions and methodology of "species" delineation criteria are not generally agreed upon in anthropology or paleontology. Indeed, mammals can typically interbreed for 2 to 3 million years or longer, so all contemporary "species" in the genus ''Homo'' would potentially have been able to interbreed at the time, and introgression from beyond the genus ''Homo'' can not ''a priori'' be ruled out. It has been suggested that ''H. naledi'' may have been a hybrid with a late surviving ''Australipith'' (taken to mean beyond ''Homo'', ed.), despite the fact that these lineages generally are regarded as long extinct. As discussed above, many introgressions have occurred between lineages, with evidence of introgression after separation of 1.5 Million years.


See also

* List of human evolution fossils ''(with images)'' * Multiregional origin of modern humans


Footnotes


References


Bibliography

*


External links


Exploring the Hominid Fossil Record
(George Washington University, Center for the Advanced Study of Hominid Paleobiology at George Washington University)
Hominid species




*
Human Timeline (Interactive)
– Smithsonian Institution, Smithsonian, National Museum of Natural History (August 2016). {{Authority control Early species of Homo, * Humans, * Hominina Primate genera Human evolution Taxa named by Carl Linnaeus Mammal genera with one living species