Australopithecus

Australopithecus (/ˌɒstrələˈpɪθɪkəs, -l-/ OS-trə-lə-PITH-i-kəs, -⁠loh-;[1] from Latin australis, meaning 'southern', and Greek πίθηκος (pithekos), meaning 'ape'; singular: australopith) is a genus of hominins that existed in Africa from around 4.2[2] to 1.9 million years ago and from which the genus Homo, including modern humans, is considered to be descended. Australopithecus is a member of the subtribe Australopithecina,[3][4] which includes Paranthropus, Kenyanthropus,[5] Ardipithecus[5] and Praeanthropus,[6] though the term "australopithecine" is sometimes used to refer only to members of Australopithecus.

Australopithecus
Temporal range: Early PliocenePresent, 4.5 – 0 Mya (Range includes humans (Homo))
Mrs Ples Face.jpg
Mrs. Ples, an Australopithecus africanus specimen
Scientific classification e
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
Subgroups

Also called Praeanthropus

Cladistically included genera (traditionally excluded):

From palaeontological and archaeological evidence, Australopithecus apparently evolved in eastern Africa around 4.2 million years ago before spreading throughout the continent and eventually becoming extinct 1.9 million years ago (or 1.2 million years ago if Paranthropus is included).[7] While none of the groups normally directly assigned to this group survived, Australopithecus does not appear to be literally extinct (in the sense of having no living descendants), as the genus Homo probably emerged from late Australopithecus species[2] such as Australopithecus garhi,[8] Australopithecus africanus[9] and/or Australopithecus sediba.[10] During that time, a number of australopithecine species emerged, including the aforementioned three species as well as Australopithecus afarensis, Australopithecus anamensis, Australopithecus bahrelghazali and Australopithecus deyiremeda (proposed).

Some debate exists as to whether other hominid species of this time, such as Paranthropus robustus, Paranthropus boisei and Paranthropus aethiopicus (considered 'robust australopiths'), belong to a separate genus or Australopithecus (considered 'gracile australopiths)'. The genus name Paranthropus was proposed by South African palaeontologist Robert Broom upon the discovery of P. robustus in 1938.[2] Occasional suggestions have been made (by Cele-Conde et al. 2002 and 2007) that A. africanus should also be moved to Paranthropus.[2] On the basis of craniodental evidence, Strait and Grine (2004) suggest that Australopithecus is paraphyletic and that A. anamensis and A. garhi should be assigned to new genera.[11] Within Australopithecus, numerous researchers such as Leakey (2001), Ward (2001), White (2002), Alemseged (2006) and Kimbel (2009) suggest that A. bahrelghazali is simply a western version of A. afarensis and not a separate species.[12][13]

Australopithecus species played a significant part in human evolution, with most scientists in the field believing Homo was derived from Australopithecus[2] at some time between 3 and 2 million years ago.[14] 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.[15][16] 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).[17] One of the australopith species evolved into the genus Homo in Africa[18] and from early Homo species eventually into modern humans, H. sapiens sapiens.[18]

EvolutionEdit

Australopiths (A. anamensis) are probably descended from or closely related to Ardipithecus ramidus.[19] Some features of A. anamensis show similarities to features of both Ardipithecus ramidus (wide diastema, post-orbital constriction) and Sahelanthropus tchadensis (post-orbital constriction, shape of its mid-face and neurocranium), but also some dissimilarities.[19]

Gracile australopiths shared several traits with modern apes and humans, and were widespread throughout Eastern and Northern Africa around 3.5 million years ago. The earliest evidence of fundamentally bipedal hominids can be observed at the site of Laetoli in Tanzania. This site contains hominid footprints that are remarkably similar to those of modern humans and have been dated to as old as 3.6 million years.[20] 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.

 
Map of the fossil sites of the early australopithecines in Africa

Australopithecus anamensis, A. afarensis, and A. africanus are among the most famous of the extinct hominins. A. africanus was once considered to be ancestral to the genus Homo (in particular Homo erectus). However, fossils assigned to the genus Homo have been found that are older than A. africanus.[citation needed] Thus, the genus Homo either split off from the genus Australopithecus at an earlier date (the latest common ancestor being either A. afarensis[citation needed] or an even earlier form, possibly Kenyanthropus[citation needed]), or both developed from a yet possibly unknown common ancestor independently.[citation needed]

According to the Chimpanzee Genome Project, the human (Ardipithecus, Australopithecus and Homo) and chimpanzee (Pan troglodytes and Pan paniscus) lineages diverged from a common ancestor about five to six million years ago, assuming a constant rate of evolution. It is theoretically more likely for evolution to happen more slowly, as opposed to more quickly, from the date suggested by a gene clock (the result of which is given as a youngest common ancestor, i.e., the latest possible date of divergence.) However, hominins discovered more recently are somewhat older than the presumed rate of evolution would suggest.[21]

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.[21]

MorphologyEdit

The brains of most species of Australopithecus were roughly 35% of the size of a modern human brain[22] with an endocranial volume average of 466 c.c.[14] Although this is more than the average endocranial volume of chimpanzee brains (360 c.c.)[14] the earliest Australopiths (A. anamensis) appear to have been within the chimpanzee range,[19] whereas some later Australopith fossils have a larger endocranial volume than that of some early Homo fossils.[14]

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. In several species there is a considerable degree of sexual dimorphism, males being larger than females.[23] Modern humans do not display the same degree of sexual dimorphism as Australopithecus appears to have. 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. New research suggests, however, that australopithecines exhibited a lesser degree of sexual dimorphism than these figures suggest, but the issue is not settled.[23]

According to one scholar, A. Zihlman, Australopithecus body proportions closely resemble those of bonobos (Pan paniscus),[24] leading evolutionary biologists such as Jeremy Griffith to suggest that bonobos may be phenotypically similar to Australopithecus.[25] Furthermore, thermoregulatory models suggest that Australopithecus species were fully hair covered, more like chimpanzees and bonobos, and unlike humans.[26]

Species variationsEdit

Opinions differ as to whether the species A. aethiopicus, A. boisei, and A. robustus should be included within the genus Australopithecus, and no current consensus exists as to whether they should be placed in a distinct genus, Paranthropus,[27] which is suggested along with Homo to have developed as part of a clade with A. africanus as its basal root.[28] Until the last half-decade, the majority of the scientific community included all the species shown in the box at the top of this article in a single genus.[citation needed] The postulated genus Paranthropus was morphologically distinct from Australopithecus, and its specialized morphology implies that its behaviour may have been quite different from that of its ancestors, although it has been suggested that the distinctive features of A. aethiopicus, A. boisei, and A. robustus may have evolved independently. In reviewing both the literature and fossil record in 2007 Wood and Constantino recognised two competing hypotheses, one that aethiopicus, boisei and robustus are a distinct clade with boisei and robustus evolving from aethiopicus. The second is that there was some parallel evolution of two distinct lines, with boisei evolving from aethiopicus, but robustus evolving separately from A. africanus. (They also acknowledge that Skelton & McHenry in 1992 proposed that aethiopicus is separate from a boisei/robustus clade).[29]

Evolutionary roleEdit

The fossil record seems to indicate that Australopithecus is the common ancestor of the distinct group of hominids now called Paranthropus (the "robust australopiths"), and most likely the genus Homo, which includes modern humans. Although the intelligence of these early hominids was likely no more sophisticated than in modern apes, the bipedal stature is the key element that distinguishes the group from previous primates, which were quadrupeds. The morphology of Australopithecus upset what scientists previously believed — namely, that strongly increased brain size had preceded bipedalism.

If A. afarensis was the definite hominid that left the footprints at Laetoli, that strengthens the notion that A. afarensis had a small brain, but was a biped. Fossil evidence such as this makes it clear that bipedalism far predated large brains. However, it remains a matter of controversy as to how bipedalism first emerged (several concepts are still being studied). 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. However, many anthropologists argue that these advantages were not large enough to cause the emergence of bipedalism.

A recent study of primate evolution and morphology noted that all apes, both modern and fossil, show skeletal adaptations to erect posture of the trunk and that 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 suggested that erect, straight-legged walking originated as an adaptation to tree-dwelling. Studies of modern orangutans in Sumatra have shown that these apes use four legs when walking on large, stable branches, and swing underneath slightly smaller branches, but are bipedal and keep their legs very straight when walking on multiple flexible branches under 4 cm diameter, while also using their arms for balance and additional support. This enables them to get nearer to the edge of the tree canopy to get fruit or cross to another tree.[30]

The ancestors of gorillas and chimpanzees are suggested to have become more specialized in climbing vertical tree trunks, using a bent hip and bent knee posture that matches the knuckle-walking posture they use for ground travel. This was due to climate changes around 11 to 12 million years ago that affected forests in East and Central Africa, so periods occurred when openings prevented travel through the tree canopy, and at these times, ancestral hominids could have adapted the erect walking behavior for ground travel. Humans are closely related to these apes, and share features including wrist bones apparently strengthened for knuckle-walking.[31]

However, the view that human ancestors were knuckle-walkers is now questioned since the anatomy and biomechanics of knuckle-walking in chimpanzees and gorillas are different, suggesting that this ability evolved independently after the last common ancestor with the human lineage.[32] Further comparative analysis with other primates suggests that these wrist-bone adaptations support a palm-based tree walking.[32]

Radical changes in morphology took place before gracile australopiths evolved; the pelvis structure and feet are very similar to modern humans.[33] The teeth have small canines, but australopiths generally evolved a larger postcanine dentition with thicker enamel.[34]

Most species of Australopithecus were not any more adept at tool use than modern nonhuman primates[citation needed], yet modern African apes, chimpanzees, and most recently gorillas, have been known to use simple tools (i.e. cracking open nuts with stones and using long sticks to dig for termites in mounds), and chimpanzees have been observed using spears (not thrown) for hunting.[citation needed]

For a long time, no known stone tools were associated with A. afarensis, and paleoanthropologists commonly thought that stone artifacts only dated back to about 2.5 million years ago.[35] However, a 2010 study suggests the hominin species ate meat by carving animal carcasses with stone implements. This finding pushes back the earliest known use of stone tools among hominins to about 3.4 million years ago.[36]

In a pair of companion papers, the discoverers of A. garhi note that it is possible A. garhi used stone tools, as remains of this species and butchered animal remains were found close to each other.[37][38]

DentitionEdit

Australopithecines have thirty two teeth, like modern humans, but with an intermediate formation; between the great apes and humans. Their molars were parallel, like those of great apes, and they had a slight pre-canine diastema. But, 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.[39]

The molars of Australopithicus fit together in much the same way human's do, with low crowns and four low, rounded cusps used for crushing. They have cutting edges on the crests.[39]

Robust australopithecines (like A. boisei and A. robustus) had larger cheek, or buccal, teeth than the smaller – or gracile – species (like A. afarensis and A. africanus). It is possible that they had more tough, fibrous plant material in their diets while the smaller species of Australopithecus had more meat. But it is also possibly due to their generally larger build requiring more food. Their larger molars do support a slightly different diet, including some hard food.[39]

Australopithecines also had thick enamel, like those in genus Homo, while other great apes have markedly thinner enamel. One explanation for the thicker enamel is that these hominins were living more on the ground than in the trees and were foraging for tubers, nuts, and cereal grains. They would also have been eating a lot of gritty dirt with the food, which would wear at enamel, so thicker enamel would be advantageous. Or, it could simply indicate a change in diet. Robust australopithecines wore their molar surfaces down flat, unlike the more gracile species, who kept their crests, which certainly seems to suggest a different diet. The gracile Australopithecus had larger incisors, which indicates tearing and more meat in the diet, likely scavenged. The wear patterns on the tooth surfaces support a largely herbivorous diet.[39]

When we examine the buccal microwear patterns on the teeth of A. afarensis and A. anamensis, we see that A. afarensis did not consume a lot of grasses or seeds, but rather ate fruits and leaves, but A. anamensis did eat grasses and seeds in addition to fruits and leaves.[40]

DietEdit

 
Artistic interpretation of Australopithecus afarensis

In a 1979 preliminary microwear study of Australopithecus fossil teeth, anthropologist Alan Walker theorized that robust australopiths were largely frugivorous.[41] Australopithecus species mainly ate fruit, vegetables, small lizards, and tubers. 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.[42]

These observations have been interpreted as evidence that P. robustus may have fed on hard and brittle foods, such as some nuts and seeds.[42] More recently, new analyses based on three-dimensional renderings of wear facets have confirmed earlier work, but have also suggested that P. robustus ate hard foods primarily as a fallback resource, while A. africanus ate more mechanically tough foods.[43] A recent study looking at enamel fractures suggests A. africanus actually ate more hard foods than P. robustus, with double the frequency of antemortem chips.[44]

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.[45]

In 2005, fossils of animal bones with butchery marks dating 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.[46]

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.[47]

A study in 2018 found non-carious cervical lesions, caused by acid erosion, on the teeth of A. africanus suggesting the individual ate a lot of acidic fruits.[48]

History of studyEdit

The type specimen for genus Australopithecus 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 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 ancestor of humans.[49] Later, Scottish paleontologist Robert Broom and Dart set about to search for more early hominin specimens, and at several sites they found more A. africanus remains, as well as fossils of a species Broom named Paranthropus (which would now be recognised as P. robustus). Initially, anthropologists were largely hostile to the idea that these discoveries were anything but apes, though this changed during the late 1940s.[49] By 1950, Mayr was treating Australopithecus as a species of Homo, Homo transvaalensis, on the grounds that all bipedal apes should be treated as part of Homo.[28] However, the contra view taken by Robinson in 1954, excluding Australopiths from Homo, became the prevalent view in the 1950s.[28]

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 have continued to excavate the gorge, uncovering further evidence for australopithecines, as well as for Homo habilis and Homo erectus.[49] The scientific community took 20 years to widely accept Australopithecus as a member of the family tree.

Then, 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[50][51] 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,[52] though this is heavily disputed.

A. afarensis, A. anamensis, and A. bahrelghazali were split off into the genus Praeanthropus, but this genus been largely dismissed.[53]

PhylogenyEdit

A taxonomy of the Australopithecus within the great apes is assessed as follows, with Paranthropus and Homo emerging among the Australopithecus.[54] The genus Australopithecus with conventional definitions is assessed to be highly paraphyletic, i.e. it is not a natural group, and the genera Kenyanthropus, Paranthropus and Homo are included.[55][56][57] The exact phylogeny within Australopithecus is still highly controversial. Approximate radiation dates of daughter clades is shown in Millions of years ago (Mya). Sahelanthropus, Orrorin, and Ardipithecus, possibly sisters to Australopithecus, are not shown here.

Hominoidea (20.4 Mya)

Hylobatidae (gibbons)

Hominidae (15.7)

Ponginae (orangutans)

Homininae  (8.8)

Gorillini (gorillas)

Hominini  (6.3)

Panina (chimpanzees)

Hominina (4)

A. anamensis (†3.8)

A. afarensis

A. garhi

A. deyiremeda (†3.3)

A. africanus

Homo (2.5)

Homo habilis  

Paranthropus (†1.2)

Kenyanthropus platyops

Homo rudolfensis  

(1.9)

Homo ergaster  

Homo erectus  

(1.2)

Homo antecessor ssp.  

(0.8)

Homo heidelbergensis(†0.7)

Homo neanderthalensis (†0.25) 

Homo sapiens  

Inconsistent taxonomyEdit

Even though Australopithecus is classified as a "genus", several other genera appear to have emerged within it: Homo, Kenyanthropus and Paranthropus. This genus is thus regarded as an entrenched paraphyletic wastebasket taxon.[58][59][60][61] Resolving this into monophyletic groupings would require extensive renaming of species in the binomial nomenclature. Possibilities suggested have been to rename Homo sapiens to Australopithecus sapiens[62] (or even Pan sapiens[63][64]), or to rename all the Australopithecus species.[65]

Notable specimensEdit

  • KT-12/H1, an A. bahrelghazali mandibular fragment, discovered 1995 in Sahara, Chad
  • AL 129-1, an A. afarensis knee joint, discovered 1973 in Hadar, Ethiopia
  • Karabo, a juvenile male A. sediba, discovered in South Africa
  • Laetoli footprints, preserved hominin footprints in Tanzania
  • Lucy, a 40%-complete skeleton of a female A. afarensis, discovered 1974 in Hadar, Ethiopia
  • Selam, remains of a three-year-old A. afarensis female, discovered in Dikika, Ethiopia
  • STS 5 (Mrs. Ples), the most complete skull of an A. africanus ever found in South Africa
  • STS 14, remains of an A. africanus, discovered 1947 in Sterkfontein, South Africa
  • STS 71, skull of an A. africanus, discovered 1947 in Sterkfontein, South Africa
  • Taung Child, skull of a young A. africanus, discovered 1924 in Taung, South Africa

GalleryEdit

See alsoEdit

ReferencesEdit

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