Australopithecus Temporal range: | |
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Mrs. Ples, an Australopithecus africanus specimen | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Suborder: | Haplorhini |
Infraorder: | Simiiformes |
Family: | Hominidae |
Subfamily: | Homininae |
Tribe: | Hominini |
Subtribe: | Australopithecina |
Genus: | † Australopithecus R.A. Dart, 1925 |
Type species | |
† Australopithecus africanus Dart, 1925 | |
Species | |
Classically excluded but cladistically included: |
Australopithecus ( /ˌɒstrələˈpɪθɪkəs,-loʊ-/ , OS-trə-lə-PITH-i-kəs, -loh-; [1] from Latin australis 'southern',and Ancient Greek πίθηκος (pithekos) 'ape' [2] ) is a genus of early hominins that existed in Africa during the Pliocene and Early Pleistocene. The genera Homo (which includes modern humans), Paranthropus , and Kenyanthropus evolved from some Australopithecus species. Australopithecus is a member of the subtribe Australopithecina, [3] [4] which sometimes also includes Ardipithecus , [5] 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. [6] [7]
Furthermore, because e.g. A. africanus is more closely related to for instance humans, or their ancestors at the time, than e.g. A. anamensis and many more Australopithecus branches, Australopithecus cannot be consolidated into a coherent grouping without also including the Homo genus and other genera.
The earliest known member of the genus, A. anamensis, existed in eastern Africa around 4.2 million years ago. Australopithecus fossils become more widely dispersed throughout eastern and southern Africa (the Chadian A. bahrelghazali indicates the genus was much more widespread than the fossil record suggests), before eventually becoming pseudo-extinct 1.9 million years ago (or 1.2 to 0.6 million years ago if Paranthropus is included). While none of the groups normally directly assigned to this group survived, Australopithecus gave rise to living descendants, as the genus Homo emerged from an Australopithecus species [6] [8] [9] [10] [11] [ excessive citations ] at some time between 3 and 2 million years ago. [12]
Australopithecus possessed two of three duplicated genes derived from SRGAP2 roughly 3.4 and 2.4 million years ago ( SRGAP2B and SRGAP2C ), the second of which contributed to the increase in number and migration of neurons in the human brain. [13] [14] Significant changes to the hand first appear in the fossil record of later A. afarensis about 3 million years ago (fingers shortened relative to thumb and changes to the joints between the index finger and the trapezium and capitate). [15]
The first Australopithecus specimen, the type specimen, was discovered in 1924 in a lime quarry by workers at Taung, South Africa. The specimen was studied by the Australian anatomist Raymond Dart, who was then working at the University of the Witwatersrand in Johannesburg. The fossil skull was from a three-year-old bipedal primate (nicknamed Taung Child) that he named Australopithecus africanus . The first report was published in Nature in February 1925. Dart realised that the fossil contained a number of humanoid features, and so he came to the conclusion that this was an early human ancestor. [16] Later, Scottish paleontologist Robert Broom and Dart set out to search for more early hominin specimens, and several more A. africanus remains from various sites. Initially, anthropologists were largely hostile to the idea that these discoveries were anything but apes, though this changed during the late 1940s. [16]
In 1950, evolutionary biologist Ernst Walter Mayr said that all bipedal apes should be classified into the genus Homo, and considered renaming Australopithecus to Homo transvaalensis. [17] However, the contrary view taken by Robinson in 1954, excluding australopiths from Homo, became the prevalent view. [17] The first australopithecine fossil discovered in eastern Africa was an A. boisei skull excavated by Mary Leakey in 1959 in Olduvai Gorge, Tanzania. Since then, the Leakey family has continued to excavate the gorge, uncovering further evidence for australopithecines, as well as for Homo habilis and Homo erectus . [16] The scientific community took 20 more years to widely accept Australopithecus as a member of the human family tree.
In 1997, an almost complete Australopithecus skeleton with skull was found in the Sterkfontein caves of Gauteng, South Africa. It is now called "Little Foot" and it is around 3.7 million years old. It was named Australopithecus prometheus [18] [19] which has since been placed within A. africanus. Other fossil remains found in the same cave in 2008 were named Australopithecus sediba , which lived 1.9 million years ago. A. africanus probably evolved into A. sediba, which some scientists think may have evolved into H. erectus, [20] though this is heavily disputed.
In 2003, Spanish writer Camilo José Cela Conde and evolutionary biologist Francisco J. Ayala proposed resurrecting the genus Praeanthropus to house Orrorin , A. afarensis, A. anamensis, A. bahrelghazali, and A. garhi, [21] but this genus has been largely dismissed. [22]
With the apparent emergence of the genera Homo, Kenyanthropus , and Paranthropus in the genus Australopithecus, taxonomy runs into some difficulty, as the name of species incorporates their genus. According to cladistics, groups should not be left paraphyletic, where it is kept not consisting of a common ancestor and all of its descendants. [23] [24] [25] [26] [27] [28] Resolving this problem would cause major ramifications in the nomenclature of all descendent species. Possibilities suggested have been to rename Homo sapiens to Australopithecus sapiens [29] (or even Pan sapiens [30] [31] ), or to move some Australopithecus species into new genera. [7]
In 2002 and again in 2007, Cele-Conde et al. suggested that A. africanus be moved to Paranthropus. [6] On the basis of craniodental evidence, Strait and Grine (2004) suggest that A. anamensis and A. garhi should be assigned to new genera. [32] It is debated whether or not A. bahrelghazali should be considered simply a western variant of A. afarensis instead of a separate species. [33] [34]
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A. anamensis may have descended from or was closely related to Ardipithecus ramidus . [35] A. anamensis shows some similarities to both Ar. ramidus and Sahelanthropus . [35]
Australopiths shared several traits with modern apes and humans, and were widespread throughout Eastern and Northern Africa by 3.5 million years ago (MYA). The earliest evidence of fundamentally bipedal hominins is a 3.6 MYA fossil trackway in Laetoli, Tanzania, which bears a remarkable similarity to those of modern humans. The footprints have generally been classified as australopith, as they are the only form of prehuman hominins known to have existed in that region at that time. [36]
According to the Chimpanzee Genome Project, the human–chimpanzee last common ancestor existed about five to six million years ago, assuming a constant rate of mutation. However, hominin species dated to earlier than the date could call this into question. [37] Sahelanthropus tchadensis , commonly called "Toumai", is about seven million years old and Orrorin tugenensis lived at least six million years ago. Since little is known of them, they remain controversial among scientists since the molecular clock in humans has determined that humans and chimpanzees had a genetic split at least a million years later.[ citation needed ] One theory suggests that the human and chimpanzee lineages diverged somewhat at first, then some populations interbred around one million years after diverging. [37]
The brains of most species of Australopithecus were roughly 35% of the size of a modern human brain [38] with an endocranial volume average of 466 cc (28.4 cu in). [12] Although this is more than the average endocranial volume of chimpanzee brains at 360 cc (22 cu in) [12] the earliest australopiths (A. anamensis) appear to have been within the chimpanzee range, [35] whereas some later australopith specimens have a larger endocranial volume than that of some early Homo fossils. [12]
Most species of Australopithecus were diminutive and gracile, usually standing 1.2 to 1.4 m (3 ft 11 in to 4 ft 7 in) tall. It is possible that they exhibited a considerable degree of sexual dimorphism, males being larger than females. [39] In modern populations, males are on average a mere 15% larger than females, while in Australopithecus, males could be up to 50% larger than females by some estimates. However, the degree of sexual dimorphism is debated due to the fragmentary nature of australopith remains. [39] One paper finds that A. afarensis had a level of dimorphism close to modern humans. [40]
According to A. Zihlman, Australopithecus body proportions closely resemble those of bonobos (Pan paniscus), [41] leading evolutionary biologist Jeremy Griffith to suggest that bonobos may be phenotypically similar to Australopithecus. [42] Furthermore, thermoregulatory models suggest that australopiths were fully hair covered, more like chimpanzees and bonobos, and unlike humans. [43]
The fossil record seems to indicate that Australopithecus is ancestral to Homo and modern humans. It was once assumed that large brain size had been a precursor to bipedalism, but the discovery of Australopithecus with a small brain but developed bipedality upset this theory. Nonetheless, it remains a matter of controversy as to how bipedalism first emerged. The advantages of bipedalism were that it left the hands free to grasp objects (e.g., carry food and young), and allowed the eyes to look over tall grasses for possible food sources or predators, but it is also argued that these advantages were not significant enough to cause the emergence of bipedalism.[ citation needed ] Earlier fossils, such as Orrorin tugenensis , indicate bipedalism around six million years ago, around the time of the split between humans and chimpanzees indicated by genetic studies. This suggests that erect, straight-legged walking originated as an adaptation to tree-dwelling. [44] Major changes to the pelvis and feet had already taken place before Australopithecus. [45] It was once thought that humans descended from a knuckle-walking ancestor, [46] but this is not well-supported. [47]
Australopithecines have thirty-two teeth, like modern humans. Their molars were parallel, like those of great apes, and they had a slight pre-canine gap (diastema). Their canines were smaller, like modern humans, and with the teeth less interlocked than in previous hominins. In fact, in some australopithecines, the canines are shaped more like incisors. [48] The molars of Australopithecus fit together in much the same way those of humans do, with low crowns and four low, rounded cusps used for crushing. They have cutting edges on the crests. [48] However, australopiths generally evolved a larger postcanine dentition with thicker enamel. [49] Australopiths in general had thick enamel, like Homo, while other great apes have markedly thinner enamel. [48] Robust australopiths wore their molar surfaces down flat, unlike the more gracile species, who kept their crests. [48]
Australopithecus species are thought to have eaten mainly fruit, vegetables, and tubers, and perhaps easy-to-catch animals such as small lizards. Much research has focused on a comparison between the South African species A. africanus and Paranthropus robustus. Early analyses of dental microwear in these two species showed, compared to P. robustus, A. africanus had fewer microwear features and more scratches as opposed to pits on its molar wear facets. [50] Microwear patterns on the cheek teeth of A. afarensis and A. anamensis indicate that A. afarensis predominantly ate fruits and leaves, whereas A. anamensis included grasses and seeds (in addition to fruits and leaves). [51] The thickening of enamel in australopiths may have been a response to eating more ground-bound foods such as tubers, nuts, and cereal grains with gritty dirt and other small particulates which would wear away enamel. Gracile australopiths had larger incisors, which indicates tearing food was important, perhaps eating scavenged meat. Nonetheless, the wearing patterns on the teeth support a largely herbivorous diet. [48]
In 1992, trace-element studies of the strontium/calcium ratios in robust australopith fossils suggested the possibility of animal consumption, as they did in 1994 using stable carbon isotopic analysis. [52] In 2005, fossil animal bones with butchery marks dating to 2.6 million years old were found at the site of Gona, Ethiopia. This implies meat consumption by at least one of three species of hominins occurring around that time: A. africanus, A. garhi, and/or P. aethiopicus. [53] In 2010, fossils of butchered animal bones dated 3.4 million years old were found in Ethiopia, close to regions where australopith fossils were found. [54]
Robust australopithecines (Paranthropus) had larger cheek teeth than gracile australopiths, possibly because robust australopithecines had more tough, fibrous plant material in their diets, whereas gracile australopiths ate more hard and brittle foods. [48] However, such divergence in chewing adaptations may instead have been a response to fallback food availability. In leaner times, robust and gracile australopithecines may have turned to different low-quality foods (fibrous plants for the former, and hard food for the latter), but in more bountiful times, they had more variable and overlapping diets. [55] [56] In a 1979 preliminary microwear study of Australopithecus fossil teeth, anthropologist Alan Walker theorized that robust australopiths ate predominantly fruit (frugivory). [57]
A study in 2018 found non-carious cervical lesions, caused by acid erosion, on the teeth of A. africanus , probably caused by consumption of acidic fruit. [58]
It is debated if the Australopithecus hand was anatomically capable of producing stone tools. [59] A. garhi was associated with large mammal bones bearing evidence of processing by stone tools may indicate australopithecine tool production. [60] [61] [62] [63] Stone tools dating to roughly the same time as A. garhi (about 2.6 mya) were later discovered at the nearby Gona and Ledi-Geraru sites, but the appearance of Homo at Ledi-Geraru (LD 350-1) casts doubt on australopithecine authorship. [64]
In 2010, cut marks dating to 3.4 mya on a bovid leg were found at the Dikaka site, which were at first attributed to butchery by A. afarensis, [65] but because the fossil came from a sandstone unit (and were modified by abrasive sand and gravel particles during the fossilisation process), the attribution to butchery is dubious. [66]
In 2015, the Lomekwi culture was discovered at Lake Turkana dating to 3.3 mya, possibly attributable to Kenyanthropus [67] or A. deyiremeda. [68]
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.
Kenyanthropus is a genus of extinct hominin identified from the Lomekwi site by Lake Turkana, Kenya, dated to 3.3 to 3.2 million years ago during the Middle Pliocene. It contains one species, K. platyops, but may also include the 2 million year old Homo rudolfensis, or K. rudolfensis. Before its naming in 2001, Australopithecus afarensis was widely regarded as the only australopithecine to exist during the Middle Pliocene, but Kenyanthropus evinces a greater diversity than once acknowledged. Kenyanthropus is most recognisable by an unusually flat face and small teeth for such an early hominin, with values on the extremes or beyond the range of variation for australopithecines in regard to these features. Multiple australopithecine species may have coexisted by foraging for different food items, which may be reason why these apes anatomically differ in features related to chewing.
Orrorin is an extinct genus of primate within Homininae from the Miocene Lukeino Formation and Pliocene Mabaget Formation, both of Kenya.
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. 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. A. afarensis probably descended from A. anamensis and is hypothesised to have given rise to Homo, though the latter is debated.
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.
Sterkfontein is a set of limestone caves of special interest in paleoanthropology located in Gauteng province, about 40 kilometres (25 mi) northwest of Johannesburg, South Africa in the Muldersdrift area close to the town of Krugersdorp. The archaeological sites of Swartkrans and Kromdraai are in the same area. Sterkfontein is a South African National Heritage Site and was also declared a World Heritage Site in 2000. The area in which it is situated is known as the Cradle of Humankind. The Sterkfontein Caves are also home to numerous wild African species including Belonogaster petiolata, a wasp species of which there is a large nesting presence.
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".
Australopithecus anamensis is a hominin species that lived approximately between 4.2 and 3.8 million years ago and is the oldest known Australopithecus species, living during the Plio-Pleistocene era.
Paranthropus aethiopicus is an extinct species of robust australopithecine from the Late Pliocene to Early Pleistocene of East Africa about 2.7–2.3 million years ago. However, it is much debated whether or not Paranthropus is an invalid grouping and is synonymous with Australopithecus, so the species is also often classified as Australopithecus aethiopicus. Whatever the case, it is considered to have been the ancestor of the much more robust P. boisei. It is debated if P. aethiopicus should be subsumed under P. boisei, and the terms P. boisei sensu lato and P. boisei sensu stricto can be used to respectively include and exclude P. aethiopicus from P. boisei.
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. One individual, presumed female based on size, may have been 140 cm tall.
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 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.
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.
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.
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 homininians. 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.
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.
The Middle Awash is a paleoanthropological research area in the northwest corner of Gabi Rasu in the Afar Region along the Awash River in Ethiopia's Afar Depression. It is a unique natural laboratory for the study of human origins and evolution and a number of fossils of the earliest hominins, particularly of the Australopithecines, as well as some of the oldest known Olduwan stone artifacts, have been found at the site—all of late Miocene, the Pliocene, and the very early Pleistocene times, that is, about 5.6 million years ago (mya) to 2.5 mya. It is broadly thought that the divergence of the lines of the earliest humans (hominins) and of chimpanzees (hominids) was completed near the beginning of that time range, or sometime between seven and five mya. However, the larger community of scientists provide several estimates for periods of divergence that imply a greater range for this event, see CHLCA: human-chimpanzee split.
Post-canine megadontia is a relative enlargement of the molars and premolars compared to the size of the incisors and canines. This phenomenon is seen in some early hominid ancestors such as Paranthropus aethiopicus.
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 is interpreted as having 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.
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 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.
Forms such as Ardipithecus, Sahelanthropus, and Orrorin have also been admitted to the pantheon, though this has clearly been facilitated by their great age. And in a nod to history, the venerable genus Paranthropus has been grandfathered in for use by those who think it useful. But except for the widely dismissed revival of Praeanthropus, there has been little real rethinking of the hugely minimalist hominid taxonomy, generic as well as specific, that Mayr foisted on us all those years ago...