2/13/2021 Homo - Wikipedia

Homo
Homo (from Latin homō 'man') is the genus that emerged in the (otherwise extinct) genus Australopithecus that encompasses the extant species Homo sapiens (modern humans), plus several extinct species classified as either ancestral to or closely related to modern humans (depending on the species), most notably Homo erectus and Homo neanderthalensis. The genus emerged with the
appearance of Homo habilis just over 2 million years ago.[1] Homo, together with the genus Paranthropus, is probably sister to Australopithecus africanus, which itself had previously split from the lineage of Pan, the chimpanzees.[2][3] Homo
Temporal range: Piacenzian-Present,
Homo erectus appeared about 2 million years ago and, in several early migrations, spread throughout Africa (where it is dubbed Homo ergaster) and Eurasia. It was likely the first human species to live in a hunter-gatherer 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 2.8–0 Ma
years ago.[4] ↓

PreꞒ Ꞓ O S D C P T J K PgN
Homo sapiens (anatomically modern humans) emerged close to 300,000 to 200,000 years ago,[5]
most likely in Africa, and 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 beginning about 70–
50,000 years ago[6][7][8][9] leading to the lasting colonisation of Eurasia and Oceania by 50,000 years ago. Both in Africa and Eurasia, H. sapiens met with and interbred with[10][11] archaic humans. Separate archaic (non-sapiens) human species are thought to have survived until around 40,000 years ago (Neanderthal extinction), with possible late survival of hybrid species as late as 12,000 years ago
(Red Deer Cave people).

Contents
Names and taxonomy
Evolution
Australopithecus
Homo habilis Homo ergaster/erectus skull

Homo erectus Scientific classification

Phylogeny Kingdom: Animalia
Dispersal
Phylum: Chordata
List of lineages
Class: Mammalia
See also
Order: Primates
Footnotes
Suborder: Haplorhini
References
External links Infraorder: Simiiformes
Family: Hominidae

Names and taxonomy Subfamily: Homininae

Tribe: Hominini

See Homininae for an overview of taxonomy. Genus: Homo

Linnaeus, 1758
The Latin noun homō (genitive hominis) means "human being" or "man" in the generic sense of "human being, mankind".[a] The binomial name Homo sapiens was coined by Carl Linnaeus (1758).[13][b] Names for other species of the genus were introduced beginning in the second half of the 19th century (H. neanderthalensis 1864, H. erectus 1892).
Type species
Even today, the genus Homo has not been strictly defined.[15][16][17]
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 did not even bother to define Homo when he first created it for Homo sapiens
humans in the 18th century. The discovery of Neanderthal brought the first addition.
Linnaeus, 1758
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 and early Pliocene times produced a rich mix for debating classifications. There is continuing debate on delineating Homo from Australopithecus—or, indeed, delineating Homo Species
from Pan, 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 inherited from the earlier Australopithecus of more than four million years ago, as demonstrated by the Laetoli footprints; and (2) human tool culture
having begun by 2.5 million years ago. Homo sapiens
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 was a single species with a large geographic spread of early migrations. Many such names are now dubbed as "synonyms" with Homo, including Pithecanthropus,[18] †Homo antecessor
Protanthropus,[19] Sinanthropus,[20] Cyphanthropus,[21] Africanthropus,[22] Telanthropus,[23] Atlanthropus,[24] and Tchadanthropus.[25][26] †Homo erectus
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 (uncertain placement)—for example, H. neanderthalensis vs. H. sapiens neanderthalensis, or †Homo ergaster
H. georgicus vs. H. erectus georgicus.[27] Some recently extinct species in the genus Homo are only recently discovered and do not as yet have consensus binomial names (see Denisova hominin and Red Deer Cave people).[28] Since the beginning of the Holocene, it is likely that Homo sapiens (anatomically modern humans) has been the only extant species of Homo. †Homo floresiensis
John Edward Gray (1825) was an early advocate of classifying taxa by designating tribes and families.[29] Wood and Richmond (2000) proposed that Hominini ("hominins") be designated as a tribe 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 †Homo habilis
include only the genus Homo — that is, not including the earlier upright walking hominins of the Pliocene such as Australopithecus, Orrorin tugenensis, Ardipithecus, or Sahelanthropus.[30] Designations alternative to Hominina existed, or were offered: Australopithecinae (Gregory & Hellman 1939) and Preanthropinae (Cela-Conde & Altaba 2002);[31][32][33] and later, Cela-Conde and Ayala (2003) †Homo heidelbergensis
proposed that the four genera Australopithecus, Ardipithecus, Praeanthropus, and Sahelanthropus be grouped with Homo within Hominini (sans pan).[34]
†Homo luzonensis
Evolution †Homo rudolfensis
†Homo naledi
See Hominini and Chimpanzee–human last common ancestor for the separation of Australopithecina and Panina.
†Homo neanderthalensis

Australopithecus For other species or subspecies

suggested, see below.
Several species, including Australopithecus garhi, Australopithecus sediba, Australopithecus africanus, and Australopithecus afarensis, have been proposed as the ancestor or sister of the Homo lineage.[35][36] These species have morphological features that align them with Homo, but there is no consensus as to which gave rise to Homo.
Synonyms
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 tools (the Oldowan industry), and thus by definition with the beginning of the 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.[37] LD 350-1, a fossil mandible fragment dated to 2.8 Mya, discovered in 2015 in Afar, Ethiopia, was described as combining "primitive traits seen in early Australopithecus with derived morphology observed in later Homo.[38] Some authors would push the development of Homo close to or even past 3 Mya.[39] Synonyms
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.[40] Africanthropus Dreyer, 1935

The most salient physiological development between the earlier australopithecine species and Homo is the increase in endocranial volume (ECV), from about 460 cm3 (28 cu in) in A. garhi to 660 cm3 (40 cu in) in H. habilis and further to 760 cm3 (46 cu in) in H. erectus, 1,250 cm3 (76 cu in) in H. heidelbergensis and up to 1,760 cm3 (107 cu in) in H. neanderthalensis. However, a steady rise in Atlanthropus Arambourg, 1954
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.[41] Cyphanthropus Pycraft, 1928
Palaeanthropus Bonarelli, 1909
Homo habilis
Palaeoanthropus Freudenberg,
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.[42][43] 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.[37] Furthermore, H. habilis was long thought 1927
to be the ancestor of the more gracile Homo ergaster (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.[44] With the publication of Dmanisi skull 5 in 2013, it has become less certain that Asian H. erectus is a descendant of African H. Pithecanthropus Dubois, 1894
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[45] and widely dispersed throughout Eurasia (including Europe, Indonesia, China) by 0.5 Mya.[46]
Protanthropus Haeckel, 1895
Sinanthropus Black, 1927
Homo erectus
Tchadanthropus Coppens, 1965
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, which seemed to exhibit transitional traits with H. habilis. As the earliest evidence for H. erectus was found outside of Africa, it was considered
Telanthropus Broom & Anderson
plausible that H. erectus developed in Eurasia and then migrated back to Africa. Based on fossils from the Koobi Fora 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 (1.9 to 1.4 million years ago), during the early Calabrian.[47] 1949

A separate South African species Homo gautengensis has been postulated as contemporary with Homo erectus in 2010.[48]

Phylogeny
A taxonomy of Homo within the great apes is assessed as follows, with Paranthropus and Homo emerging within Australopithecus (shown here cladistically granting Paranthropus, Kenyanthropus, and Homo).[49][50][3][51][52][53][54][55][56][57][58][59][60][61] The exact phylogeny within Australopithecus is still highly controversial. Approximate radiation dates of daughter clades
are shown in millions of years ago (Mya).[62] Graecopithecus, Sahelanthropus, Orrorin, possibly sisters to Australopithecus, are not shown here. Note that the naming of groupings is sometimes muddled as often certain groupings are presumed before a cladistic analyses is performed.[55]

  Evolutionary tree chart emphasizing the subfamily Homininae and

  Hylobatidae (gibbons) the tribe Hominini. After diverging from the line to Ponginae the
early Homininae split into the tribes Hominini and Gorillini. The
  early Hominini split further, separating the line to Homo from the
  Ponginae (orangutans)
lineage of Pan. Currently, tribe Hominini designates the subtribes
Hominina, containing genus Homo; Panina, genus Pan; and
Hominoidea   Australopithecina, with several extinct genera—the subtribes are
  Gorillini (gorillas)
(20.4 Mya) Hominidae not labelled on this chart.
(15.7) Homininae  
(8.8)   Panina (chimpanzees)
Hominini
(7.8)  
  Australopithecines (incl. Australopithecus, Kenyanthropus, Paranthropus, Homo)

 
  †Ardipithecus ramidus

 
  Australopithecus anamensis s.s. (†3.8)

 
  †Australopithecus afarensis

 
  †Australopithecus garhi A model of the evolution of the genus Homo over the
last 2 million years (vertical axis). The rapid "Out of
  Africa" expansion of H. sapiens is indicated at the top
  Australopithecus deyiremeda (†3.4) of the diagram, with admixture indicated with
Neanderthals, Denisovans, and unspecified archaic
  African hominins. Late survival of robust
  Kenyanthropus platyops (†3.3) australopithecines (Paranthropus) alongside Homo
until 1.2 Mya is indicated in purple.
 
  Australopithecus africanus (†2.1)
 
   
  Paranthropus (†1.2)

  Homo habilis (†1.5)

 

  Homo rudolfensis (†1.9)

 

  Homo ergaster (†1.4)

 
Australopithecines
(7.3 Mya) Australopithecus s.l.
(3.9)     African Homo erectus s.s. (†)
     
   
      Asian Homo erectus s.s. ((†)0.1)
     
 
 
Homo   Red Deer Cave people (†0.01)
(3.4) H. erectus s.l.
(1.9)
    Homo antecessor (†0.8)
   

  H. neanderthalensis ((†)0.05)
 
(1.2) Neandersovans
  (0.3)
   
(0.5)   Denisova people ((†)0.05)

  Homo sapiens
 

 
  Australopithecus sediba (†2.0)

 
Homo floresiensis (†0.05)
 

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.[63][64][65][54][66][67] 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.[68][69] 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.[54] 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.[70][66] Likewise the genetic structure of sub-Saharan Africans seems to be indicative of introgression from a west Eurasian population some 3,000 years ago.[71][72]

Some evidence suggests that 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 and Homo floresiensis.[56][73]

Dispersal

By about 1.8 million years ago, Homo erectus is present in both East Africa (Homo ergaster) and in Western Asia (Homo georgicus). The ancestors of Indonesian Homo floresiensis may have left Africa even earlier.[74]

Homo erectus and related or derived archaic human species over the next 1.5 million years spread throughout Africa and Eurasia[75][76] (see: Recent African origin of modern humans). Europe is reached by about 0.5 Mya by Homo heidelbergensis.

Homo neanderthalensis and Homo sapiens develop after about 300 kya. 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, Greece, more than 150,000 years older than
previous H. sapiens finds in Europe.[77][78][79]

Most notable is the Southern Dispersal of H. sapiens around 60 kya, which led to the lasting peopling of Oceania and Eurasia by anatomically modern humans.[80] H. sapiens interbred with archaic humans both in Africa and in Eurasia, in Eurasia notably with Neanderthals and Denisovans.[81]

Among extant populations of Homo sapiens, the deepest temporal division is found in the San people of Southern Africa, estimated at close to 130,000 years,[82] or possibly more than 300,000 years ago.[83] Temporal division among non-Africans is of the order of 60,000 years in the case of Australo-Melanesians. Division of Europeans and East Asians is of the order of 50,000 years, Successive dispersals of Homo erectus (yellow),
with repeated and significant admixture events throughout Eurasia during the Holocene.
Homo neanderthalensis (ochre) and Homo
sapiens (red).
Archaic human species may have survived until the beginning of the Holocene (Red Deer Cave people), although they were mostly extinct or absorbed by the expanding H. sapiens populations by 40 kya (Neanderthal extinction).

List of lineages
The species status of H. rudolfensis, H. ergaster, H. georgicus, H. antecessor, H. cepranensis, H. rhodesiensis, H. neanderthalensis, Denisova hominin, 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 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 (2.1–1.5 Mya, membership in Homo questionable), Homo erectus (1.8–0.1 Mya, including the majority of the age of the genus, and the majority of archaic varieties as subspecies,[84] including H. heidelbergensis as a late or transitional
variety[85]) and Homo sapiens (300 kya to present, including H. neanderthalensis and other varieties as 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[86] or longer,[87] 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.[88] It has been suggested that H. naledi may have been a hybrid with a late surviving Australipith (taken to
mean beyond Homo, ed.),[89] 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.

https://en.wikipedia.org/wiki/Homo 1/3
2/13/2021 Homo - Wikipedia

Comparative table of Homo lineages

Cranial capacity Discovery/

Temporal range
Lineages Habitat Adult height Adult mass Fossil record publication
(kya) (cm3)
of name
H. habilis 1960
2,100–1,500[c][d] Tanzania 110–140 cm (3 ft 7 in – 4 ft 7 in) 33–55 kg (73–121 lb) 510–660 Many
membership in Homo uncertain 1964
H. rudolfensis 1972
1,900 Kenya 700 2 sites
membership in Homo uncertain 1986
H. gautengensis 2010
1,900–600 South Africa 100 cm (3 ft 3 in) 3 individuals[92][e]
also classified as H. habilis 2010
1891
H. erectus 1,900–140[93][f][94][g] Africa, Eurasia 180 cm (5 ft 11 in) 60 kg (130 lb) 850 (early) – 1,100 (late) Many[h][i]
1892
H. ergaster 1949
1,800–1,300[96] East and Southern Africa 700–850 Many
African H. erectus 1975
1994
H. antecessor 1,200–800 Western Europe 175 cm (5 ft 9 in) 90 kg (200 lb) 1,000 2 sites
1997
H. heidelbergensis 1907
600–300[j] Europe, Africa 180 cm (5 ft 11 in) 90 kg (200 lb) 1,100–1,400 Many
early H. neanderthalensis 1908
H. cepranensis 1994
c. 450[97] Italy 1,000 1 skull cap
a single fossil, possibly H. erectus 2003
H. rhodesiensis 1921
c. 300 Zambia 1,300 Single or very few
early H. sapiens 1921
2013
H. naledi c. 300[98] South Africa 150 cm (4 ft 11 in) 45 kg (99 lb) 450 15 individuals
2015
H. sapiens ——
c. 300–present[k] Worldwide 150–190 cm (4 ft 11 in – 6 ft 3 in) 50–100 kg (110–220 lb) 950–1,800 (extant)
(anatomically modern humans) 1758
55–70 kg (121–154 lb) 1829
H. neanderthalensis 240–40[101][l] Europe, Western Asia 170 cm (5 ft 7 in) 1,200–1,900 Many
(heavily built) 1864
H. floresiensis 2003
190–50 Indonesia 100 cm (3 ft 3 in) 25 kg (55 lb) 400 7 individuals
classification uncertain 2004
H. tsaichangensis 2008(?)
c. 100[m] Taiwan 1 individual
possibly H. erectus or Denisova 2015
2007
H. luzonensis c. 67[104][105] Philippines 3 individuals
2019
2000
Denisova hominin 40 Siberia 2 sites
2010[n]
Red Deer Cave people
15–12[o][106] Southwest China Very few
possible H. sapiens subspecies or hybrid

See also
List of human evolution fossils (with images)
Multiregional origin of modern humans

Footnotes
a. The word "human" itself is from Latin humanus, an adjective formed on the root of homo, thought to derive from a Proto-Indo-European word for "earth" reconstructed as *dhǵhem-.[12]
b. Note that in 1959, Carl Linnaeus was designated as the lectotype for Homo sapiens[14]:4 which means that following the nomenclatural rules, Homo sapiens was validly defined as the animal species to which Linnaeus belonged.
c. Confirmed H. habilis fossils are dated to between 2.1 and 1.5 million years ago. This date range overlaps with the emergence of Homo erectus.[90][91]
d. Hominins with "proto-Homo" traits may have lived as early as 2.8 million years ago, as suggested by a fossil jawbone classified as transitional between Australopithecus and Homo discovered in 2015.
e. A species proposed in 2010 based on the fossil remains of three individuals dated between 1.9 and 0.6 million years ago. The same fossils were also classified as H. habilis, H. ergaster or Australopithecus by other anthropologists.
f. H. erectus may have appeared some 2 million years ago. Fossils dated to as much as 1.8 million years ago have been found both in Africa and in Southeast Asia, and the oldest fossils by a narrow margin (1.85 to 1.77 million years ago) were found in the Caucasus, so that it is unclear whether H. erectus emerged in Africa and migrated to Eurasia, or if, conversely, it evolved in Eurasia and migrated back to Africa.
g. Homo erectus soloensis, found in Java, is considered the latest known survival of H. erectus. Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed back the date of its extinction of H. e. soloensis to 143,000 years ago at the latest, more likely before 550,000 years ago. [95]
h. Now also included in H. erectus are Peking Man (formerly Sinanthropus pekinensis) and Java Man (formerly Pithecanthropus erectus).
i. H. erectus is now grouped into various subspecies, including Homo erectus erectus, Homo erectus yuanmouensis, Homo erectus lantianensis, Homo erectus nankinensis, Homo erectus pekinensis, Homo erectus palaeojavanicus, Homo erectus soloensis, Homo erectus tautavelensis, Homo erectus georgicus. The distinction from descendant species such as Homo ergaster, Homo floresiensis, Homo antecessor, Homo
heidelbergensis and indeed Homo sapiens is not entirely clear.
j. The type fossil is Mauer 1, dated to ca. 0.6 million years ago. The transition from H. heidelbergensis to H. neanderthalensis between 300 and 243 thousand years ago is conventional, and makes use of the fact that there is no known fossil in this period. Examples of H. heidelbergensis are fossils found at Bilzingsleben (also classified as Homo erectus bilzingslebensis).
k. The age of H. sapiens has long been assumed to be close to 200,000 years, but since 2017 there have been a number of suggestions extending this time to has high as 300,000 years. In 2017, fossils found in Jebel Irhoud (Morocco) suggest that Homo sapiens may have speciated by as early as 315,000 years ago.[99] Genetic evidence has been adduced for an age of roughly 270,000 years.[100]
l. The first humans with "proto-Neanderthal traits" lived in Eurasia as early as 0.6 to 0.35 million years ago (classified as H. heidelbergensis, also called a chronospecies because it represents a chronological grouping rather than being based on clear morphological distinctions from either H. erectus or H. neanderthalensis). There is a fossil gap in Europe between 300 and 243 kya, and by convention, fossils younger than 243 kya are
called "Neanderthal".[102]
m. younger than 450 kya, either between 190–130 or between 70–10 kya[103]
n. provisional names Homo sp. Altai or Homo sapiens ssp. Denisova.
o. Bølling–Allerød warming period

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External links
Exploring the Hominid Fossil Record (http://www.bradshawfoundation.com/origins/index.php) (Center for the Advanced Study of Hominid Paleobiology at George Washington University)
Hominid species (http://www.talkorigins.org/faqs/homs/species.html)
Prominent Hominid Fossils (http://www.talkorigins.org/faqs/homs/specimen.html)
Mikko's Phylogeny archive (http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/synapsida/eutheria/primates/hominoidea/hominidae_1.html)
"Homo" (https://www.eol.org/pages/42268) at the Encyclopedia of Life
Human Timeline (Interactive) (http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive) – Smithsonian, National Museum of Natural History (August 2016).

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