Hominidae

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Hominidae [1]
Temporal range: Miocene–present, 17–0  Ma
Hominidae (extant species).jpg
The eight extant hominid species, one row per genus (humans, chimpanzees, gorillas, orangutans)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Parvorder: Catarrhini
Superfamily: Hominoidea
Family: Hominidae
Gray, 1825 [2]
Type genus
Homo
Linnaeus, 1758
Subfamilies

sister: Hylobatidae

Distribution of the Great Apes.png
Distribution of great ape species
Synonyms
  • Pongidae Elliot, 1913
  • Gorillidae Frechkop, 1943
  • Panidae Ciochon, 1983

The Hominidae ( /hɒˈmɪnɪdiː/ ), whose members are known as the great apes [note 1] or hominids ( /ˈhɒmɪnɪdz/ ), are a taxonomic family of primates that includes eight extant species in four genera: Pongo (the Bornean, Sumatran and Tapanuli orangutan); Gorilla (the eastern and western gorilla); Pan (the chimpanzee and the bonobo); and Homo , of which only modern humans (Homo sapiens) remain. [1]

Contents

Numerous revisions in classifying the great apes have caused the use of the term hominid to change over time. The original meaning of "hominid" referred only to humans (Homo) and their closest extinct relatives. However, by the 1990s humans, apes, and their ancestors were considered to be "hominids".

The earlier restrictive meaning has now been largely assumed by the term hominin , which comprises all members of the human clade after the split from the chimpanzees (Pan). The current meaning of "hominid" includes all the great apes including humans. Usage still varies, however, and some scientists and laypersons still use "hominid" in the original restrictive sense; the scholarly literature generally shows the traditional usage until the turn of the 21st century. [5]

Within the taxon Hominidae, a number of extant and extinct genera are grouped with the humans, chimpanzees, and gorillas in the subfamily Homininae; others with orangutans in the subfamily Ponginae (see classification graphic below). The most recent common ancestor of all Hominidae lived roughly 14 million years ago, [6] when the ancestors of the orangutans speciated from the ancestral line of the other three genera. [7] Those ancestors of the family Hominidae had already speciated from the family Hylobatidae (the gibbons), perhaps 15 to 20 million years ago. [7] [8]

Due to the close genetic relationship between humans and the other great apes, certain animal rights organizations, such as the Great Ape Project, argue that nonhuman great apes are persons and should be given basic human rights. Twenty-nine countries have instituted research bans to protect great apes from any kind of scientific testing. [9]

Evolution

Sumatran orangutan (Pongo abelii) Man of the woods.JPG
Sumatran orangutan (Pongo abelii)

In the early Miocene, about 22 million years ago, there were many species of tree-adapted primitive catarrhines from East Africa; the variety suggests a long history of prior diversification. Fossils from 20 million years ago include fragments attributed to Victoriapithecus , the earliest Old World monkey. Among the genera thought to be in the ape lineage leading up to 13 million years ago are Proconsul , Rangwapithecus , Dendropithecus , Limnopithecus , Nacholapithecus , Equatorius , Nyanzapithecus , Afropithecus , Heliopithecus, and Kenyapithecus , all from East Africa.

At sites far distant from East Africa, the presence of other generalized non-cercopithecids, that is, non-monkey primates, of middle Miocene age— Otavipithecus from cave deposits in Namibia, and Pierolapithecus and Dryopithecus from France, Spain and Austria—is further evidence of a wide diversity of ancestral ape forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the early and middle Miocene. The most recent of these far-flung Miocene apes (hominoids) is Oreopithecus , from the fossil-rich coal beds in northern Italy and dated to 9 million years ago.

Molecular evidence indicates that the lineage of gibbons (family Hylobatidae), the "lesser apes", diverged from that of the great apes some 18–12 million years ago, and that of orangutans (subfamily Ponginae) diverged from the other great apes at about 12 million years. There are no fossils that clearly document the ancestry of gibbons, which may have originated in a still-unknown South East Asian hominoid population; but fossil proto-orangutans, dated to around 10 million years ago, may be represented by Sivapithecus from India and Griphopithecus from Turkey. [10] Species close to the last common ancestor of gorillas, chimpanzees and humans may be represented by Nakalipithecus fossils found in Kenya and Ouranopithecus fossils found in Greece. Molecular evidence suggests that between 8 and 4 million years ago, first the gorillas (genus Gorilla), and then the chimpanzees (genus Pan) split off from the line leading to humans. Human DNA is approximately 98.4% identical to that of chimpanzees when comparing single nucleotide polymorphisms (see human evolutionary genetics). [11] The fossil record, however, of gorillas and chimpanzees is limited; both poor preservation—rain forest soils tend to be acidic and dissolve bone—and sampling bias probably contribute most to this problem.

Other hominins probably adapted to the drier environments outside the African equatorial belt; and there they encountered antelope, hyenas, elephants and other forms becoming adapted to surviving in the East African savannas, particularly the regions of the Sahel and the Serengeti. The wet equatorial belt contracted after about 8 million years ago, and there is very little fossil evidence for the divergence of the hominin lineage from that of gorillas and chimpanzees—which split was thought to have occurred around that time. The earliest fossils argued by some to belong to the human lineage are Sahelanthropus tchadensis (7 Ma) and Orrorin tugenensis (6 Ma), followed by Ardipithecus (5.5–4.4 Ma), with species Ar. kadabba and Ar. ramidus.

Taxonomy

Terminology

Humans are one of the four extant hominid genera. Akha cropped hires.JPG
Humans are one of the four extant hominid genera.

The classification of the great apes has been revised several times in the last few decades; these revisions have led to a varied use of the word "hominid" over time. The original meaning of the term referred to only humans and their closest relatives—what is now the modern meaning of the term "hominin". The meaning of the taxon Hominidae changed gradually, leading to a modern usage of "hominid" that includes all the great apes including humans.

A number of very similar words apply to related classifications:

A cladogram indicating common names (cf. more detailed cladogram below):

Hominoidea

Hylobatidae
   gibbons

Hominidae
hominids, great apes
hominoids, apes

Extant and fossil relatives of humans

A fossil hominid exhibit at The Museum of Osteology, Oklahoma City, Oklahoma Fossil hominids.jpg
A fossil hominid exhibit at The Museum of Osteology, Oklahoma City, Oklahoma

Hominidae was originally the name given to the family of humans and their (extinct) close relatives, with the other great apes (that is, the orangutans, gorillas and chimpanzees) all being placed in a separate family, the Pongidae. However, that definition eventually made Pongidae paraphyletic because at least one great ape species (the chimpanzees) proved to be more closely related to humans than to other great apes. Most taxonomists today encourage monophyletic groups—this would require, in this case, the use of Pongidae to be restricted to just one closely related grouping. Thus, many biologists now assign Pongo (as the subfamily Ponginae) to the family Hominidae. The taxonomy shown here follows the monophyletic groupings according to the modern understanding of human and great ape relationships.

Humans and close relatives including the tribes Hominini and Gorillini form the subfamily Homininae (see classification graphic below). (A few researchers go so far as to refer the chimpanzees and the gorillas to the genus Homo along with humans.) [15] [16] [17] But, those fossil relatives more closely related to humans than the chimpanzees represent the especially close members of the human family, and without necessarily assigning subfamily or tribal categories. [18]

Many extinct hominids have been studied to help understand the relationship between modern humans and the other extant hominids. Some of the extinct members of this family include Gigantopithecus , Orrorin , Ardipithecus , Kenyanthropus , and the australopithecines Australopithecus and Paranthropus . [19]

The exact criteria for membership in the tribe Hominini under the current understanding of human origins are not clear, but the taxon generally includes those species that share more than 97% of their DNA with the modern human genome, and exhibit a capacity for language or for simple cultures beyond their 'local family' or band. The theory of mind concept—including such faculties as empathy, attribution of mental state, and even empathetic deception—is a controversial criterion; it distinguishes the adult human alone among the hominids. Humans acquire this capacity after about four years of age, whereas it has not been proven (nor has it been disproven) that gorillas or chimpanzees ever develop a theory of mind. [20] This is also the case for some New World monkeys outside the family of great apes, as, for example, the capuchin monkeys.

However, even without the ability to test whether early members of the Hominini (such as Homo erectus , Homo neanderthalensis , or even the australopithecines) had a theory of mind, it is difficult to ignore similarities seen in their living cousins. Orangutans have shown the development of culture comparable to that of chimpanzees, [21] and some[ who? ] say the orangutan may also satisfy those criteria for the theory of mind concept. These scientific debates take on political significance for advocates of great ape personhood.

Phylogeny

Below is a cladogram with extinct species. [22] [23] [24] [ failed verification ] It is indicated approximately how many million years ago (Mya) the clades diverged into newer clades. [25]

Hominidae chart inverted.svg

Taxonomy of Hominoidea (emphasis on family Hominidae): After an initial separation from the main line by the Hylobatidae (gibbons) some 18 million years ago, the line of Ponginae broke away, leading to the orangutan; later, the Homininae split into the tribes Hominini (led to humans and chimpanzees) and Gorillini (led to gorillas).

Extant

There are eight living species of great ape which are classified in four genera. The following classification is commonly accepted: [1]

Fossil

Replica of the skull sometimes known as "Nutcracker Man", found by Mary Leakey Paranthropus boisei skull.jpg
Replica of the skull sometimes known as "Nutcracker Man", found by Mary Leakey

In addition to the extant species and subspecies, archaeologists, paleontologists, and anthropologists have discovered and classified numerous extinct great ape species as below, based on the taxonomy shown. [27]

Family Hominidae

Description

Gorilla Gorilla 019.jpg
Gorilla

The great apes are tailless primates, with the smallest living species being the bonobo at 30 to 40 kilograms (66 to 88 lb) in weight, and the largest being the eastern gorillas, with males weighing 140 to 180 kilograms (310 to 400 lb). In all great apes, the males are, on average, larger and stronger than the females, although the degree of sexual dimorphism varies greatly among species. Hominid teeth are similar to those of the Old World monkeys and gibbons, although they are especially large in gorillas. The dental formula is 2.1.2.32.1.2.3. Human teeth and jaws are markedly smaller for their size than those of other apes, which may be an adaptation to not only having supplanted with extensive tool use the role of jaws in hunting and fighting, but also eating cooked food since the end of the Pleistocene. [32] [33]

Behavior

Although most living species are predominantly quadrupedal, they are all able to use their hands for gathering food or nesting materials, and, in some cases, for tool use. [34] They build complex sleeping platforms, also called nests, in trees to sleep in at night, but chimpanzees and gorillas also build terrestrial nests, and gorillas can also sleep on the bare ground. [35]

All species are omnivorous, [36] although chimpanzees and orangutans primarily eat fruit. When gorillas run short of fruit at certain times of the year or in certain regions, they resort to eating shoots and leaves, often of bamboo, a type of grass. Gorillas have extreme adaptations for chewing and digesting such low-quality forage, but they still prefer fruit when it is available, often going miles out of their way to find especially preferred fruits. Humans, since the Neolithic revolution, have consumed mostly cereals and other starchy foods, including increasingly highly processed foods, as well as many other domesticated plants (including fruits) and meat.

Gestation in great apes lasts 8–9 months, and results in the birth of a single offspring, or, rarely, twins. The young are born helpless, and require care for long periods of time. Compared with most other mammals, great apes have a remarkably long adolescence, not being weaned for several years, [37] and not becoming fully mature for eight to thirteen years in most species (longer in orangutans and humans). As a result, females typically give birth only once every few years. There is no distinct breeding season. [34]

Gorillas and chimpanzees live in family groups of around five to ten individuals, although much larger groups are sometimes noted. Chimpanzees live in larger groups that break up into smaller groups when fruit becomes less available. When small groups of female chimpanzees go off in separate directions to forage for fruit, the dominant males can no longer control them and the females often mate with other subordinate males. In contrast, groups of gorillas stay together regardless of the availability of fruit. When fruit is hard to find, they resort to eating leaves and shoots.

This fact is related to gorillas' greater sexual dimorphism relative to that of chimpanzees; that is, the difference in size between male and female gorillas is much greater than that between male and female chimpanzees. This enables gorilla males to physically dominate female gorillas more easily. In both chimpanzees and gorillas, the groups include at least one dominant male, and young males leave the group at maturity.

Due to the close genetic relationship between humans and the other great apes, certain animal rights organizations, such as the Great Ape Project, argue that nonhuman great apes are persons and, per the Declaration on Great Apes, should be given basic human rights. In 1999, New Zealand was the first country to ban any great ape experimentation, and now 29 countries have currently instituted a research ban to protect great apes from any kind of scientific testing.

On 25 June 2008, the Spanish parliament supported a new law that would make "keeping apes for circuses, television commercials or filming" illegal. [38] On 8 September 2010, the European Union banned the testing of great apes. [39]

Conservation

The following table lists the estimated number of great ape individuals living outside zoos.

SpeciesEstimated
number		Conservation
status		Refs
Bornean orangutan 61,234 Critically endangered [40]
Sumatran orangutan 6,667 Critically endangered [41]
Tapanuli orangutan 800 Critically endangered [42]
Western gorilla 200,000 Critically endangered [43]
Eastern gorilla 6,000 Critically endangered [43]
Chimpanzee 200,000 Endangered [44] [45]
Bonobo 10,000 Endangered [44]
Human 8,040,640,000N/A [46]

See also

Notes

  1. ↑ "Great ape" is a common name rather than a taxonomic label, and there are differences in usage, even by the same author. The term may or may not include humans, as when Dawkins writes "Long before people thought in terms of evolution ... great apes were often confused with humans" [3] and "gibbons are faithfully monogamous, unlike the great apes which are our closer relatives." [4]

Related Research Articles

<i>Ardipithecus</i> Extinct genus of hominins

Ardipithecus is a genus of an extinct hominine that lived during the Late Miocene and Early Pliocene epochs in the Afar Depression, Ethiopia. Originally described as one of the earliest ancestors of humans after they diverged from the chimpanzees, the relation of this genus to human ancestors and whether it is a hominin is now a matter of debate. Two fossil species are described in the literature: A. ramidus, which lived about 4.4 million years ago during the early Pliocene, and A. kadabba, dated to approximately 5.6 million years ago. Initial behavioral analysis indicated that Ardipithecus could be very similar to chimpanzees, however more recent analysis based on canine size and lack of canine sexual dimorphism indicates that Ardipithecus was characterised by reduced aggression, and that they more closely resemble bonobos.

<span class="mw-page-title-main">Human evolution</span> Evolutionary process leading to anatomically modern humans

Human evolution is the evolutionary process within the history of primates that led to the emergence of Homo sapiens as a distinct species of the hominid family that includes all the great apes. This process involved the gradual development of traits such as human bipedalism, dexterity, and complex language, as well as interbreeding with other hominins, indicating that human evolution was not linear but weblike. The study of the origins of humans, variously known by the terms anthropogeny, anthropogenesis, or anthropogony, involves several scientific disciplines, including physical and evolutionary anthropology, paleontology, and genetics.

<span class="mw-page-title-main">Homininae</span> Subfamily of mammals

Homininae, also called "African hominids" or "African apes", is a subfamily of Hominidae. It includes two tribes, with their extant as well as extinct species: 1) the tribe Hominini ―and 2) the tribe Gorillini (gorillas). Alternatively, the genus Pan is sometimes considered to belong to its own third tribe, Panini. Homininae comprises all hominids that arose after orangutans split from the line of great apes. The Homininae cladogram has three main branches, which lead to gorillas and to humans and chimpanzees. There are two living species of Panina and two living species of gorillas, but only one extant human species. Traces of extinct Homo species, including Homo floresiensis have been found with dates as recent as 40,000 years ago. Organisms in this subfamily are described as hominine or hominines.

<i>Orrorin</i> Postulated early hominin discovered in Kenya

Orrorin is an extinct genus of primate within Homininae from the Miocene Lukeino Formation and Pliocene Mabaget Formation, both of Kenya.

<i>Australopithecus</i> Genus of hominin ancestral to modern humans

Australopithecus is a genus of early hominins that existed in Africa during the Pliocene and Early Pleistocene. The genera Homo, Paranthropus, and Kenyanthropus evolved from some Australopithecus species. Australopithecus is a member of the subtribe Australopithecina, which sometimes also includes Ardipithecus, though the term "australopithecine" is sometimes used to refer only to members of Australopithecus. Species include A. garhi, A. africanus, A. sediba, A. afarensis, A. anamensis, A. bahrelghazali and A. deyiremeda. Debate exists as to whether some Australopithecus species should be reclassified into new genera, or if Paranthropus and Kenyanthropus are synonymous with Australopithecus, in part because of the taxonomic inconsistency.

<i>Sahelanthropus</i> Extinct hominid from Miocene Africa

Sahelanthropus tchadensis is an extinct species of hominid dated to about 7 million years ago, during the Miocene epoch. The species, and its genus Sahelanthropus, was announced in 2002, based mainly on a partial cranium, nicknamed Toumaï, discovered in northern Chad.

<span class="mw-page-title-main">Ape</span> Branch of primates

Apes are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia, which together with its sister group Cercopithecidae form the catarrhine clade, cladistically making them monkeys. Apes do not have tails due to a mutation of the TBXT gene. In traditional and non-scientific use, the term ape can include tailless primates taxonomically considered Cercopithecidae, and is thus not equivalent to the scientific taxon Hominoidea. There are two extant branches of the superfamily Hominoidea: the gibbons, or lesser apes; and the hominids, or great apes.

<span class="mw-page-title-main">Pongidae</span> Obsolete ape taxon

Pongidae, or the pongids is an obsolete primate taxon containing chimpanzees, gorillas and orangutans. By this definition pongids were also called "great apes". This taxon is not used today but is of historical significance. The great apes are currently classified as Hominidae. This entry addresses the old usage of pongid.

<i>Homo</i> Genus of hominins that includes humans and their closest extinct relatives

Homo is a genus of great ape that emerged from the genus Australopithecus and encompasses the extant species Homo sapiens and a number of extinct species classified as either ancestral or closely related to modern humans. These include Homo erectus and Homo neanderthalensis. The oldest member of the genus is Homo habilis, with records of just over 2 million years ago. Homo, together with the genus Paranthropus, is probably most closely related to the species Australopithecus africanus within Australopithecus. The closest living relatives of Homo are of the genus Pan, with the ancestors of Pan and Homo estimated to have diverged around 5.7-11 million years ago during the Late Miocene.

Paleoanthropology or paleo-anthropology is a branch of paleontology and anthropology which seeks to understand the early development of anatomically modern humans, a process known as hominization, through the reconstruction of evolutionary kinship lines within the family Hominidae, working from biological evidence and cultural evidence.

<span class="mw-page-title-main">Ponginae</span> Orangutan subfamily of apes

Ponginae, also known as the Asian hominids, is a subfamily in the family Hominidae. Once a diverse lineage of Eurasian apes, the subfamily has only one extant genus, Pongo (orangutans), which contains three extant species; the Sumatran orangutan, the Tapanuli orangutan and the Bornean orangutan. All three species are listed as critically endangered by the International Union for Conservation of Nature (IUCN).

<i>Sivapithecus</i> Genus of extinct Asian ape

Sivapithecus is a genus of extinct apes. Fossil remains of animals now assigned to this genus, dated from 12.2 million years old in the Miocene, have been found since the 19th century in the Sivalik Hills of the Indian subcontinent as well as in Kutch. Any one of the species in this genus may have been the ancestor to the modern orangutans.

<span class="mw-page-title-main">Knuckle-walking</span> Form of quadrupedal walking using the knuckles

Knuckle-walking is a form of quadrupedal walking in which the forelimbs hold the fingers in a partially flexed posture that allows body weight to press down on the ground through the knuckles. Gorillas and chimpanzees use this style of locomotion, as do anteaters and platypuses.

<span class="mw-page-title-main">Hominini</span> Tribe of mammals

The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera Homo (humans) and Pan and in standard usage excludes the genus Gorilla (gorillas).

Orthograde is a term derived from Greek ὀρθός, orthos + Latin gradi that describes a manner of walking which is upright, with the independent motion of limbs. Both New and Old World monkeys are primarily arboreal, and they have a tendency to walk with their limbs swinging in parallel to one another. This differs from the manner of walking demonstrated by the apes.

Human taxonomy is the classification of the human species within zoological taxonomy. The systematic genus, Homo, is designed to include both anatomically modern humans and extinct varieties of archaic humans. Current humans have been designated as subspecies Homo sapiens sapiens, differentiated, according to some, from the direct ancestor, Homo sapiens idaltu.

<span class="mw-page-title-main">Australopithecine</span> Extinct subtribe of the Hominini tribe, and members of the human clade

The australopithecines, formally Australopithecina or Hominina, are generally any species in the related genera of Australopithecus and Paranthropus. It may also include members of Kenyanthropus, Ardipithecus, and Praeanthropus. The term comes from a former classification as members of a distinct subfamily, the Australopithecinae. They are now classified within the Australopithecina subtribe of the Hominini tribe. All these related species are now sometimes collectively termed australopithecines, australopiths or homininans. They are the extinct, close relatives of modern humans and, together with the extant genus Homo, comprise the human clade. Members of the human clade, i.e. the Hominini after the split from the chimpanzees, are now called Hominina.

The chimpanzee–human last common ancestor (CHLCA) is the last common ancestor shared by the extant Homo (human) and Pan genera of Hominini. Estimates of the divergence date vary widely from thirteen to five million years ago.

<i>Graecopithecus</i> Extinct genus of hominids

Graecopithecus is an extinct genus of hominid that lived in southeast Europe during the late Miocene around 7.2 million years ago. Originally identified by a single lower jaw bone bearing teeth found in Pyrgos Vasilissis, Athens, Greece, in 1944, other teeth were discovered from Azmaka quarry in Bulgaria in 2012. With only little and badly preserved materials to reveal its nature, it is considered as "the most poorly known European Miocene hominoids." The creature was popularly nicknamed 'El Graeco' by scientists.

<span class="mw-page-title-main">Evolution of primates</span> Origin and diversification of primates through geologic time

The evolutionary history of the primates can be traced back 57-90 million years. One of the oldest known primate-like mammal species, Plesiadapis, came from North America; another, Archicebus, came from China. Other similar basal primates were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and Eocene. Purgatorius is the genus of the four extinct species believed to be the earliest example of a primate or a proto-primate, a primatomorph precursor to the Plesiadapiformes, dating to as old as 66 million years ago.

References

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