Homininae is a subfamily of Hominidae that includes humans, gorillas, chimpanzees, bonobos and some extinct relatives; it comprises all hominids that arose after the split from orangutans (Ponginae). The Homininae cladogram has three main branches, which lead to gorillas, chimpanzees and bonobos, and humans. There are several extant species of chimpanzees and gorillas, but only one human species remains, although traces of several hypothetical species have been found with dates as recent as 12,000 years ago (Homo floresiensis, Homo denisova). Organisms in this class are described as hominine or hominines (not to be confused with hominin or hominini).
History of discoveries and classification
Until 1980, the family Hominidae contained only humans, with the great apes in the family Pongidae. Later discoveries led to a revision of classification, with Hominidae uniting the great apes (now in the sub-family Ponginae) and humans (in the sub-family Homininae). Further discoveries indicated that gorillas and chimpanzees are more closely related to humans than they are to orangutans, leading to their current placement in Homininae as well.
The subfamily Homininae can be further subdivided into three tribes, each with only a single living genus: Gorillini (gorillas), Panini (chimpanzees), and Hominini (humans and their extinct relatives). The early Late Miocene Nakalipithecus nakayamai, described in 2007, and perhaps also its contemporary Ouranopithecus, are basal members of this clade, not assignable to any of the three extant tribes. They suggest that the Homininae tribes diverged not earlier than about 8 million years ago (see Human evolutionary genetics).
Today, chimpanzees and gorillas live in tropical forests with acid soils that rarely preserved as fossils. Although no fossil gorillas have been reported, four chimpanzee teeth, about 500,000 years old, have been discovered in the East-African rift valley (Kapthurin Formation, Kenya), where many fossils from the human lineage (hominins)[Note 1] have previously been found. This shows that some chimpanzees lived close to Homo (H. erectus or H. rhodesiensis) at the time; the same is likely true for gorillas.
- Tribe Dryopithecini†
- Rudapithecus hungaricus
- Tribe Gorillini
- Chororapithecus † (placement debated)
- Chororapithecus abyssinicus
- Genus Gorilla
- Chororapithecus † (placement debated)
- Tribe Hominini
- Sahelanthropus† (hominin status highly problematic)
- Subtribe Panina
- Subtribe Hominina
- Kenyanthropus† (placement debated)
- Kenyanthropus platyops
- Homo – immediate ancestors of modern humans
- Homo gautengensis†
- Homo rudolfensis†
- Homo habilis†
- Homo floresiensis†
- Homo erectus†
- Homo ergaster†
- Homo antecessor†
- Homo heidelbergensis†
- Homo cepranensis†
- Denisovans (scientific name has not yet been assigned)†
- Homo neanderthalensis†
- Homo rhodesiensis†
- Homo sapiens
Evolution of bipedalism
Recent studies of Ardipithecus ramidus (4.4 million years old) and Orrorin tugenensis (6 million years old) suggest some degree of bipedalism. Australopithecus and early Paranthropus may have been bipedal. Very early hominins such as Ardipithecus ramidus may have possessed an arboreal type of bipedalism. and towards efficient walking and running in modern humans.
Brain size evolution
There has been a gradual increase in brain volume (brain size) as the ancestors of modern humans progressed along the timeline of human evolution, starting from about 600 cm3 in Homo habilis up to 1500 cm3 in Homo sapiens neanderthalensis. However, modern Homo sapiens have a brain volume slightly smaller (1250 cm3) than Neanderthals, women have a brain slightly smaller than men and the Flores hominids (Homo floresiensis), nicknamed hobbits, had a cranial capacity of about 380 cm3 (considered small for a chimpanzee), about a third of the Homo erectus average. It is proposed that they evolved from H. erectus as a case of insular dwarfism. In spite of their smaller brain, there is evidence that H. floresiensis used fire and made stone tools at least as sophisticated as those of their proposed ancestors H. erectus. In this case, it seems that for intelligence, the structure of the brain is more important than its size.
Evolution of family structure and sexuality
Sexuality is related to family structure and partly shapes it. The involvement of fathers in education is quite unique to humans, at least when compared to other Homininae. Concealed ovulation and menopause in women both also occur in a few other primates however, but are uncommon in other species. Testis and penis size seems to be related to family structure: monogamy or promiscuity, or harem, in humans, chimpanzees or gorillas, respectively. The levels of sexual dimorphism are generally seen as a marker of sexual selection. Studies have suggested that the earliest hominins were dimorphic and that this lessened over the course of the evolution of the genus Homo, correlating with humans becoming more monogamous, whereas gorillas, who live in harems, show a large degree of sexual dimorphism. Concealed (or "hidden") ovulation means that the phase of fertility is not detectable in women, whereas chimpanzees advertise ovulation via an obvious swelling of the genitals. Women can be partly aware of their ovulation along the menstrual phases, but men are essentially unable to detect ovulation in women. Most primates have semi-concealed ovulation, thus one can think that the common ancestor had semi-concealed ovulation, that was inherited by gorillas, and that later evolved in concealed ovulation in humans and advertised ovulation in chimpanzees. Menopause also occurs in rhesus monkeys, and possibly in chimpanzees, but does not in gorillas and is quite uncommon in other primates (and other mammal groups).
- Chimpanzee-human last common ancestor
- List of human evolution fossils (with images)
- The Third Chimpanzee
|Wikispecies has information related to: Homininae|
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- M. Goodman (1964). "Man’s place in the phylogeny of the primates as reflected in serum proteins". In S. L. Washburn. Classification and human evolution. Aldine, Chicago. pp. 204–234.
- M. Goodman (1974). "Biochemical Evidence on Hominid Phylogeny". Annual Review of Anthropology 3: 203–228. doi:10.1146/annurev.an.03.100174.001223.
- M. Goodman, D. A. Tagle, D. H. Fitch, W. Bailey, J. Czelusniak, B. F. Koop, P. Benson, J. L. Slightom (1990). "Primate evolution at the DNA level and a classification of hominoids". Journal of Molecular Evolution 30 (3): 260–266. doi:10.1007/BF02099995. PMID 2109087.
- McBrearty S, Jablonski N (2005). "First fossil chimpanzee". Nature 437 (7055): 105–8. doi:10.1038/nature04008. PMID 16136135.
- Haaramo, Mikko (2005-01-14). "Hominoidea". Mikko's Phylogeny Archive.
- Haaramo, Mikko (2007-11-10). "Hominidae". Mikko's Phylogeny Archive.
- Barras, Colin (2012-03-14). "Chinese human fossils unlike any known species". New Scientist. Retrieved 2012-03-15.
- "National Geographic". National Geographic Society. Retrieved 25 July 2009.
- Kivell TL, Schmitt D 2009 Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor. Proc Natl Acad Sci U S A. 2009 Aug 25;106(34):14241-6. PMID 19667206
- Brown P, Sutikna T, Morwood MJ, et al. (2004). "A new small-bodied hominin from the Late Pleistocene of Flores, Indonesia". Nature 431 (7012): 1055–61. doi:10.1038/nature02999. PMID 15514638.
- Davidson, Iain. "As large as you need and as small as you can'--implications of the brain size of Homo floresiensis, (Iain Davidson)". Une-au.academia.edu. Retrieved 2011-10-30.
- Diamond, Jared. The Third Chimpanzee.
- Diamond, Jared. Why is Sex Fun?.
- "Human Evolutionary Genetics" Jobling M.A., Hurles M., Tyler Smith C. 2004, Garland Science, New York