Australopithecine in the context of Hominini

Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped /ˈbaɪpɛd/, meaning 'two feet' (from Latin bis 'double' and pes 'foot'). Types of bipedal movement include walking or running (a bipedal gait) and hopping.

Several groups of modern species are habitual bipeds whose normal method of locomotion is two-legged. In the Triassic period some groups of archosaurs, a group that includes crocodiles and dinosaurs, developed bipedalism; among the dinosaurs, all the early forms and many later groups were habitual or exclusive bipeds; the birds are members of a clade of exclusively bipedal dinosaurs, the theropods. Within mammals, habitual bipedalism has evolved multiple times, with the macropods, kangaroo rats and mice, springhare, hopping mice, pangolins and hominin apes such as australopithecines, including humans, as well as many other extinct groups evolving the trait independently.

Homo sapiens is a distinct species of the hominid family of primates, which also includes all the great apes. Over their evolutionary history, humans gradually developed traits such as bipedalism, dexterity, and complex language, as well as interbreeding with other hominins (a tribe of the African hominid subfamily), indicating that human evolution was not linear but weblike. The study of the origins of humans involves several scientific disciplines, including physical and evolutionary anthropology, paleontology, and genetics; the field is also known by the terms anthropogeny, anthropogenesis, and anthropogony—with the latter two sometimes used to refer to the related subject of hominization.

Primates diverged from other mammals about 85 million years ago (mya), in the Late Cretaceous period, with their earliest fossils appearing over 55 mya, during the Paleocene. Primates produced successive clades leading to the ape superfamily, which gave rise to the hominid and the gibbon families; these diverged some 15–20 mya. African and Asian hominids (including orangutans) diverged about 14 mya. Hominins (including the Australopithecine and Panina subtribes) parted from the Gorillini tribe between 8 and 9 mya; Australopithecine (including the extinct biped ancestors of humans) separated from the Pan genus (containing chimpanzees and bonobos) 4–7 mya. The Homo genus is evidenced by the appearance of H. habilis over 2 mya, while anatomically modern humans emerged in Africa approximately 300,000 years ago.

Australopithecus africanus is an extinct species of australopithecine which lived between about 3.3 and 2.1 million years ago in the Late Pliocene to Early Pleistocene of South Africa. The species has been recovered from Taung, Sterkfontein, Makapansgat, and Gladysvale. The first specimen, the Taung child, was described by anatomist Raymond Dart in 1924, and was the first early hominin found. However, its closer relations to humans than to other apes would not become widely accepted until the middle of the century because most had believed humans evolved outside of Africa. It is unclear how A. africanus relates to other hominins, being variously placed as ancestral to Homo and Paranthropus, to just Paranthropus, or to just P. robustus. The specimen "Little Foot" is the most completely preserved early hominin, with 90% of the skeleton intact, and the oldest South African australopith. However, it is controversially suggested that it and similar specimens be split off into "A. prometheus".

A. africanus brain volume was about 420–510 cc (26–31 cu in). Like other early hominins, the cheek teeth were enlarged and had thick enamel. Male skulls may have been more robust than female skulls. Males may have been on average 140 cm (4 ft 7 in) in height and 40 kg (88 lb) in weight, and females 125 cm (4 ft 1 in) and 30 kg (66 lb). A. africanus was a competent biped, albeit less efficient at walking than humans. A. africanus also had several upper body traits in common with arboreal non-human apes. This is variously interpreted as either evidence of a partially or fully arboreal lifestyle, or as a non-functional vestige from a more apelike ancestor. The upper body of A. africanus is more apelike than that of the East African A. afarensis.

Homo habilis (lit. 'handy man') is an extinct species of archaic human from the Early Pleistocene of East and South Africa about 2.4 million years ago to 1.65 million years ago (mya). It is among the oldest species of archaic humans. Suggestions for pushing back the age to 2.8 Mya were made in 2015 based on the discovery of a jawbone. Upon species description in 1964, H. habilis was highly contested, with many researchers recommending it be synonymised with Australopithecus africanus, the only other early hominin known at the time, but H. habilis received more recognition as time went on and more relevant discoveries were made. By the 1980s, H. habilis was proposed to have been a human ancestor, directly evolving into Homo erectus, which directly led to modern humans. This viewpoint is now debated. Several specimens with insecure species identification were assigned to H. habilis, leading to arguments for splitting, namely into "H. rudolfensis" and "H. gautengensis" of which only the former has received wide support.

H. habilis brain size generally varied from 500 to 900 cm (31–55 cu in). The body proportions of H. habilis are only known from two highly fragmentary skeletons, and is based largely on assuming a similar anatomy to the earlier australopithecines. Because of this, it has also been proposed H. habilis be moved to the genus Australopithecus as Australopithecus habilis. However, the interpretation of H. habilis as a small-statured human with inefficient long-distance travel capabilities has been challenged. The presumed female specimen OH 62 is traditionally interpreted as having been 100–120 cm (3 ft 3 in – 3 ft 11 in) in height and 20–37 kg (44–82 lb) in weight assuming australopithecine-like proportions, but assuming humanlike proportions she would have been about 148 cm (4 ft 10 in) and 35 kg (77 lb). Nonetheless, Homo habilis may have been at least partially arboreal like what is postulated for australopithecines. Early hominins are typically reconstructed as having thick hair and marked sexual dimorphism with males much larger than females, though relative male and female size is not definitively known.

Australopithecus sediba is an extinct species of australopithecine recovered from Malapa Cave, Cradle of Humankind, South Africa. It is known from a partial juvenile skeleton, the holotype MH1, and a partial adult female skeleton, the paratype MH2. They date to about 1.98 million years ago in the Early Pleistocene, and coexisted with Paranthropus robustus and Homo ergaster / Homo erectus. Malapa Cave may have been a natural death trap, the base of a long vertical shaft which creatures could accidentally fall into. A. sediba was initially described as being a potential human ancestor, and perhaps the progenitor of Homo, but this is contested and it could also represent a late-surviving population or sister species of A. africanus which had earlier inhabited the area.

MH1 has a brain volume of about 350–440 cc, similar to other australopithecines. The face of MH1 is strikingly similar to Homo instead of other australopithecines, with a less pronounced brow ridge, cheek bones, and prognathism (the amount the face juts out), and there is evidence of a slight chin. However, such characteristics could be due to juvenility and lost with maturity. The teeth are quite small for an australopithecine. MH1 is estimated at 130 cm (4 ft 3 in) tall, which would equate to an adult height of 150–156 cm (4 ft 11 in – 5 ft 1 in). MH1 and MH2 were estimated to have been about the same weight at 30–36 kg (66–79 lb). Like other australopithecines, A. sediba is thought to have had a narrow and apelike upper chest, but a broad and humanlike lower chest. Like other australopithecines, the arm anatomy seems to suggest a degree of climbing and arboreal behaviour. The pelvis indicates A. sediba was capable of a humanlike stride, but the foot points to a peculiar gait not demonstrated in any other hominin involving hyperpronation of the ankle, and resultantly rotating the leg inwards while pushing off. This suite of adaptations may represent a compromise between habitual bipedalism and arboreality.

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 skeletal elements uncovered in the three years preceding. A. garhi was originally considered to have been a direct ancestor to Homo and the human line, but is now thought to have been an offshoot. Like other australopithecines, A. garhi had a brain volume of 450 cc (27 cu in); a jaw which jutted out (prognathism); relatively large molars and premolars; adaptations for both walking on two legs (bipedalism) and grasping while climbing (arboreality); and it is possible that, though unclear if, males were larger than females (exhibited sexual dimorphism). One individual, presumed female based on size, may have been 140 cm (4 ft 7 in) tall.

A. garhi is the first pre-Homo hominin postulated to have manufactured tools—using them in butchering—and may be counted among a growing body of evidence for pre-Homo stone tool industries (the ability to manufacture tools was previously believed to have separated Homo from predecessors). A. garhi possibly produced the Oldowan industry which was previously considered to have been invented by the later H. habilis, though this may have instead been produced by contemporary Homo.

Australopithecus afarensis is an extinct species of australopithecine which lived from about 3.9–2.9 million years ago (mya) in the Pliocene of East Africa. The first fossils were discovered in the 1930s, but major fossil finds would not take place until the 1970s. From 1972 to 1977, the International Afar Research Expedition—led by anthropologists Maurice Taieb, Donald Johanson and Yves Coppens—unearthed several hundreds of hominin specimens in Hadar, Ethiopia, the most significant being the exceedingly well-preserved skeleton AL 288-1 ("Lucy") and the site AL 333 ("the First Family"). Beginning in 1974, Mary Leakey led an expedition into Laetoli, Tanzania, and notably recovered fossil trackways. In 1978, the species was first described, but this was followed by arguments for splitting the wealth of specimens into different species given the wide range of variation which had been attributed to sexual dimorphism (normal differences between males and females). A. afarensis probably descended from A. anamensis and is hypothesised to have given rise to Homo, though the latter is debated.

A. afarensis had a tall face, a delicate brow ridge, and prognathism (the jaw jutted outwards). The jawbone was quite robust, similar to that of gorillas. The living size of A. afarensis is debated, with arguments for and against marked size differences between males and females. Lucy measured perhaps 105 cm (3 ft 5 in) in height and 25–37 kg (55–82 lb), but she was rather small for her species. In contrast, a presumed male was estimated at 165 cm (5 ft 5 in) and 45 kg (99 lb). A perceived difference in male and female size may simply be sampling bias. The leg bones as well as the Laetoli fossil trackways suggest A. afarensis was a competent biped, though somewhat less efficient at walking and slower at running than humans. The arm and shoulder bones have some similar aspects to those of orangutans and gorillas, which has variously been interpreted as either evidence of partial tree-dwelling (arboreality), or basal traits inherited from the chimpanzee–human last common ancestor with no adaptive functionality.

Australopithecus bahrelghazali is an extinct species of australopithecine discovered in 1995 at Koro Toro, Bahr el Gazel, Chad, existing around 3.5 million years ago in the Pliocene. It is the first and only australopithecine known from Central Africa, and demonstrates that this group was widely distributed across Africa as opposed to being restricted to East and southern Africa as previously thought. The validity of A. bahrelghazali has not been widely accepted, in favour of classifying the specimens as A. afarensis, a better known Pliocene australopithecine from East Africa, because of the anatomical similarity and the fact that A. bahrelghazali is known only from 3 partial jawbones and an isolated premolar. The specimens inhabited a lakeside grassland environment with sparse tree cover, possibly similar to the modern Okavango Delta, and similarly predominantly ate C₄ savanna foods—such as grasses, sedges, storage organs, or rhizomes—and to a lesser degree also C₃ forest foods—such as fruits, flowers, pods, or insects. However, the teeth seem ill-equipped to process C₄ plants, so its true diet is unclear.

Australopithecus deyiremeda is an extinct species of australopithecine from Woranso–Mille, Afar Region, Ethiopia, about 3.5 to 3.3 million years ago during the Pliocene. Because it is known only from three partial jawbones, it is unclear if these specimens indeed represent a unique species or belong to the much better-known A. afarensis. A. deyiremeda is distinguished by its forward-facing cheek bones and small cheek teeth compared to those of other early hominins. It is unclear if a partial foot specimen exhibiting a dextrous big toe (a characteristic unknown in any australopithecine) can be assigned to A. deyiremeda. A. deyiremeda lived in a mosaic environment featuring both open grasslands and lake- or riverside forests, and anthropologist Fred Spoor suggests it may have been involved in the Kenyan Lomekwi stone-tool industry typically assigned to Kenyanthropus. A. deyiremeda coexisted with A. afarensis, and they may have exhibited niche partitioning to avoid competing with each other for the same resources, such as by relying on different fallback foods during leaner times.

Paranthropus robustus is a species of robust australopithecine from the Early and possibly Middle Pleistocene of the Cradle of Humankind, South Africa, about 2.27 to 0.87 (or, more conservatively, 2 to 1) million years ago. It has been identified in Kromdraai, Swartkrans, Sterkfontein, Gondolin, Cooper's, and Drimolen Caves. Discovered in 1938, it was among the first early hominins described, and became the type species for the genus Paranthropus. However, it has been argued by some that Paranthropus is an invalid grouping and synonymous with Australopithecus, so the species is also often classified as Australopithecus robustus.

Robust australopithecines—as opposed to gracile australopithecines—are characterised by heavily built skulls capable of producing high stresses and bite forces, as well as inflated cheek teeth (molars and premolars). Males had more heavily built skulls than females. P. robustus may have had a genetic susceptibility for pitting enamel hypoplasia on the teeth, and seems to have had a dental cavity rate similar to non-agricultural modern humans. The species is thought to have exhibited marked sexual dimorphism, with males substantially larger and more robust than females. Based on 3 specimens, males may have been 132 cm (4 ft 4 in) tall and females 110 cm (3 ft 7 in). Based on 4 specimens, males averaged 40 kg (88 lb) in weight and females 30 kg (66 lb). The brain volume of the specimen SK 1585 is estimated to have been 476 cc, and of DNH 155 about 450 cc (for comparison, the brain volume of contemporary Homo varied from 500 to 900 cc). P. robustus limb anatomy is similar to that of other australopithecines, which may indicate a less efficient walking ability than modern humans, and perhaps some degree of arboreality (movement in the trees).

Paranthropus boisei is a species of australopithecine from the Early Pleistocene of East Africa about 2.5 to 1.15 million years ago. The holotype specimen, OH 5, was discovered by palaeoanthropologist Mary Leakey in 1959 at Olduvai Gorge, Tanzania and described by her husband Louis a month later. It was originally placed into its own genus as "Zinjanthropus boisei", but is now relegated to Paranthropus along with other robust australopithecines. However, it is also argued that Paranthropus is an invalid grouping and synonymous with Australopithecus, so the species is also often classified as Australopithecus boisei.

Robust australopithecines are characterised by heavily built skulls capable of producing high stresses and bite forces, and some of the largest molars with the thickest enamel of any known ape. P. boisei is the most robust of this group. Brain size was about 450–550 cc (27–34 cu in), similar to other australopithecines. Some skulls are markedly smaller than others, which is taken as evidence of sexual dimorphism where females are much smaller than males, though body size is difficult to estimate given only one specimen, OH 80, definitely provides any bodily elements. The presumed male OH 80 may have been 156 cm (5 ft 1 in) tall and 61.7 kg (136 lb) in weight, and the presumed female KNM-ER 1500 124 cm (4 ft 1 in) tall (though its species designation is unclear). The arm and hand bones of OH 80 and KNM-ER 47000 suggest P. boisei was arboreal to a degree.

Ardipithecus ramidus is a species of australopithecine from the Afar region of Early Pliocene Ethiopia 4.4 million years ago (Ma). The species A. ramidus is the type species for the genus Ardipithecus. There is an older species in this same genus, Ardipithecus kadabba that was discovered more recently.

A. ramidus, unlike modern hominids, has adaptations for both walking on two legs (bipedality) and life in the trees (arboreality), as it has a divergent big toe and evidence of bipedality. This combination of a big toe that would facilitate climbing suggests that Ardipithecus was not as efficient at bipedality as humans or even Australopithecus (a genus that did not have a divergent big toe), nor as good at arboreality as non-human great apes.