Homo
Homo | |
---|---|
Notable members of Homo. Clockwise from top left: An approximate reconstruction of a Neanderthal (Homo neanderthalensis) skeleton, a modern human (Homo sapiens) female with a child, a reconstructed Homo habilis skull, and a replica skull of Peking Man (subspecies of Homo erectus). | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Suborder: | Haplorhini |
Infraorder: | Simiiformes |
Family: | Hominidae |
Subfamily: | Homininae |
Tribe: | Hominini |
Subtribe: | Hominina
|
Genus: | Homo Linnaeus, 1758 |
Type species | |
Homo sapiens
, 1758 | |
Species | |
For other species or subspecies suggested, see below. | |
Synonyms | |
Synonyms
|
Homo (from
Homo erectus appeared about 2 million years ago and spread throughout
Names and taxonomy
The Latin noun homō (genitive hominis) means "human being" or "man" in the generic sense of "human being, mankind".[c] The binomial name Homo sapiens was coined by Carl Linnaeus (1758).[d][15] Names for other species of the genus were introduced from the second half of the 19th century (H. neanderthalensis 1864, H. erectus 1892).
The genus Homo has not been strictly defined, even today.[16][17][18] Since the early human fossil record began to slowly emerge from the earth, the boundaries and definitions of the genus 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 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
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 regarded as "synonyms" with Homo, including Pithecanthropus,[20] Protanthropus,[21] Sinanthropus,[22] Cyphanthropus,[23] Africanthropus,[24] Telanthropus,[25] Atlanthropus,[26] and Tchadanthropus.[27][28]
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 , it is likely that Homo sapiens (anatomically modern humans) has been the only extant species of Homo.
Evolution
Australopithecus and the appearance of Homo
Several species, including Australopithecus garhi, Australopithecus sediba, Australopithecus africanus, and Australopithecus afarensis, have been proposed as the ancestor or sister of the Homo lineage.[36][37] 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
The most salient physiological development between the earlier australopithecine species and Homo is the increase in
Homo habilis
Homo habilis emerged about 2.1 Mya. Already before 2010, there were suggestions that H. habilis should not be placed in the genus Homo but rather in Australopithecus.[43][44] 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.[38] Furthermore, H. habilis was long thought 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.[45] With the publication of Dmanisi skull 5 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[46] and widely dispersed throughout Eurasia (including Europe, Indonesia, China) by 0.5 Mya.[47]
Homo erectus
Homo erectus has often been assumed to have developed
A separate South African species Homo gautengensis has been postulated as contemporary with H. erectus in 2010.[50]
Phylogeny
million years ago ) |
A taxonomy of Homo within the
Hominoidea
|
|
(20.4 Mya) |
Cladogram based on Dembo et al. (2016):[56]
Australopithecines
|
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(7.3 Mya) |
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.[61][53][62] Fossil evidence shows H. erectus s.s. survived at least until 117,000 yrs ago, and the even more basal H. floresiensis survived until 50,000 years ago. A 1.5-million-year H. erectus-like lineage appears to have made its way into modern humans through the Denisovans and specifically into the Papuans and aboriginal Australians.[53] 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.[63][62] The genetic structure of some sub-Saharan African groups seems to be indicative of introgression from a west Eurasian population some 3,000 years ago.[57][64]
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. H. habilis and H. floresiensis.[55][65]
Dispersal
By about 1.8 million years ago, H. erectus is present in both East Africa (H. ergaster) and in Western Asia (
Homo erectus and related or derived
Homo neanderthalensis and H. 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.[68][69][70]
Most notable is the
Among extant populations of H. sapiens, the deepest temporal division is found in the
Archaic human species may have survived until the beginning of the Holocene, 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
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, H. habilis (2.1–1.5 Mya, membership in Homo questionable), H. erectus (1.8–0.1 Mya, including the majority of the age of the genus, and the majority of archaic varieties as
Lineages | Temporal range (kya) |
Habitat | Adult height | Adult mass | Cranial capacity (cm3) |
Fossil record | Discovery/ publication of name |
---|---|---|---|---|---|---|---|
H. habilis membership in Homo uncertain |
2,100–1,500[h][i] | Tanzania | 110–140 cm (3 ft 7 in – 4 ft 7 in) | 33–55 kg (73–121 lb) | 510–660 | Many | 1960 1964 |
H. rudolfensis membership in Homo uncertain |
1,900 | Kenya | 700 | 2 sites | 1972 1986 | ||
H. gautengensis also classified as H. habilis |
1,900–600 | South Africa | 100 cm (3 ft 3 in) | 3 individuals[86][j] | 2010 2010 | ||
H. erectus | 1,900–140[87][k][88][l] | Eurasia
|
180 cm (5 ft 11 in) | 60 kg (130 lb) | 850 (early) – 1,100 (late) | Many[m][n] | 1891 1892 |
H. ergaster African H. erectus |
1,800–1,300[90] | East and Southern Africa | 700–850 | Many | 1949 1975 | ||
H. antecessor | 1,200–800 | Western Europe | 175 cm (5 ft 9 in) | 90 kg (200 lb) | 1,000 | 2 sites | 1994 1997 |
H. heidelbergensis early H. neanderthalensis |
600–300[o] | Europe, Africa | 180 cm (5 ft 11 in) | 90 kg (200 lb) | 1,100–1,400 | Many | 1907 1908 |
H. cepranensis a single fossil, possibly H. heidelbergensis |
c. 450[91] | Italy | 1,000 | 1 skull cap | 1994 2003 | ||
H. naledi | 335—236[92] | South Africa | 150 cm (4 ft 11 in) | 45 kg (99 lb) | 450 | 15 individuals | 2013 2015 |
H. longi | 309–138[93] | Northeast China | 1,420[94] | 1 individual | 1933 2021 | ||
H. rhodesiensis early H. sapiens |
c. 300 | Zambia
|
1,300 | Single or very few | 1921 1921 | ||
H. sapiens (anatomically modern humans) |
c. 300–present[p] | Worldwide | 150–190 cm (4 ft 11 in – 6 ft 3 in) | 50–100 kg (110–220 lb) | 950–1,800 | (extant) | —— 1758 |
Denisova hominin
|
c. 285 - c. 51 | Siberia | 2 sites | 2000 2010[q] | |||
H. neanderthalensis |
240–40[97][r] | Europe, Western Asia | 170 cm (5 ft 7 in) | 55–70 kg (121–154 lb) (heavily built) |
1,200–1,900 | Many | 1829 1864 |
H. floresiensis classification uncertain |
190–50 | Indonesia | 100 cm (3 ft 3 in) | 25 kg (55 lb) | 400 | 7 individuals | 2003 2004 |
Nesher Ramla Homo classification uncertain |
140–120 | Israel | several individuals | 2021 | |||
H. tsaichangensis possibly H. erectus or Denisova |
c. 100[s] | Taiwan | 1 individual | 2008(?) 2015 | |||
H. luzonensis |
c. 67[100][101] | Philippines | 3 individuals | 2007 2019 |
See also
Footnotes
- ^ a b The conventional estimate on the age of H. habilis is at roughly 2.1 to 2.3 million years.[1][2] Suggestions for pushing back the age to 2.8 Mya were made in 2015 based on the discovery of a jawbone.[3]
- 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 the H. e. soloensis extinction date back to 143,000 years ago at the latest, more likely before 550,000 years ago.[6]
- ^ 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-.[13]
- lectotype for Homo sapiens,[14] which means that following the nomenclatural rules, Homo sapiens was validly defined as the animal species to which Linnaeus belonged.
- ^ Cela-Conde & Ayala (2003) recognize five genera within Hominina: Ardipithecus, Australopithecus (including Paranthropus), Homo (including Kenyanthropus), Praeanthropus (including Orrorin), and Sahelanthropus.[34]
- ^ The line to the earliest members of Homo were derived from Australopithecus, a genus that had separated from the chimpanzee–human last common ancestor by late Miocene or early Pliocene times.[51]
- Dmanisi Man, raising the possibility that the ancestors of H. floresiensis left Africa before the appearance of H. erectus, possibly even becoming the first hominins to do so and evolved further in Asia.[55]
- ^ 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.[84][85]
- ^ 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.
- ^ 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.
- ^ 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.
- 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. [89]
- ^ Now also included in H. erectus are Peking Man (formerly Sinanthropus pekinensis) and Java Man (formerly Pithecanthropus erectus).
- Homo erectus georgicus. The distinction from descendant species such as Homo ergaster, Homo floresiensis, Homo antecessor, Homo heidelbergensisand indeed Homo sapiens is not entirely clear.
- ^ 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).
- ^ 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 as 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.[95] Genetic evidence has been adduced for an age of roughly 270,000 years.[96]
- ^ provisional names Homo sp. Altai or Homo sapiens ssp. Denisova.
- ^ 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".[98]
- ^ younger than 450 kya, either between 190–130 or between 70–10 kya[99]
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- ^ See:
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the adaptive coherence of Homo would be compromised if H. habilis is included in Homo. Thus, if these arguments are accepted the origins of the genus Homo are coincident in time and place with the emergence of H. erectus, not H. habilis.
- PMID 27298460.
A fresh look at brain size, hand morphology and earliest technology suggests that a number of key Homo attributes may already be present in generalized species of Australopithecus, and that adaptive distinctions in Homo are simply amplifications or extensions of ancient hominin trends. [...] the adaptive shift represented by the ECV of Australopithecus is at least as significant as the one represented by the ECV of early Homo, and that a major 'grade-level' leap in brain size with the advent of H. erectus is probably illusory.
- ^ Wood & Richmond (2000), p. 41: "A recent reassessment of cladistic and functional evidence concluded that there are few, if any, grounds for retaining H. habilis in Homo, and recommended that the material be transferred (or, for some, returned) to Australopithecus (Wood & Collard, 1999)."
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A partial maxilla assigned to H. habilis reliably demonstrates that this species survived until later than previously recognized, making an anagenetic relationship with H. erectus unlikely. The discovery of a particularly small calvaria of H. erectus indicates that this taxon overlapped in size with H. habilis, and may have shown marked sexual dimorphism. The new fossils confirm the distinctiveness of H. habilis and H. erectus, independently of overall cranial size, and suggest that these two early taxa were living broadly sympatrically in the same lake basin for almost half a million years.
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Lordkipanidze D, Ponce de León MS, Margvelashvili A, Rak Y, Rightmire GP, Vekua A, Zollikofer CP (October 2013). "A complete skull from Dmanisi, Georgia, and the evolutionary biology of early Homo". S2CID 20435482.
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Mounier, A.; Marchal, F.; Condemi, S. (March 2009). "Is Homo heidelbergensis a distinct species? New insight on the Mauer mandible". PMID 19249816.
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Bibliography
- Wood, Bernard; Richmond, Brian G. (July 2000). "Human evolution: taxonomy and paleobiology". PMID 10999270.
External links
- Exploring the Hominid Fossil Record (Center for the Advanced Study of Hominid Paleobiology at George Washington University)
- Hominid species
- Prominent Hominid Fossils
- Mikko's Phylogeny archive
- "Homo" at the Encyclopedia of Life
- Human Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).