| Australopithecus africanus | |
|---|---|
.JPG) | |
| Mrs. Ples, at theUniversity of Zurich | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Primates |
| Suborder: | Haplorhini |
| Infraorder: | Simiiformes |
| Family: | Hominidae |
| Subfamily: | Homininae |
| Tribe: | Hominini |
| Genus: | †Australopithecus |
| Species: | †A. africanus |
| Binomial name | |
| †Australopithecus africanus Dart, 1925 | |
| Synonyms | |
Australopithecus africanus is anextinctspecies ofaustralopithecine which lived between about 3.3 and 2.1 million years ago in theLate Pliocene toEarly Pleistocene ofSouth Africa.[1] The species has been recovered fromTaung,Sterkfontein,Makapansgat, andGladysvale. The first specimen, theTaung child, wasdescribed by anatomistRaymond Dart in 1924, and was the first earlyhominin found. However, its closer relations to humans than to otherapes would not become widely accepted until the middle of the century because most had believed humans evolved outside of Africa. It is unclear howA. africanus relates to other hominins, being variously placed as ancestral toHomo andParanthropus, to justParanthropus, or to justP. 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, thecheek teeth were enlarged and had thickenamel. 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 competentbiped, 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 ofA. africanus is more apelike than that of the East AfricanA. afarensis.
A. africanus, unlike most other primates, seems to have exploitedC4 foods such as grasses, seeds,rhizomes, undergroundstorage organs, or potentially creatures higher up on the food chain. Nonetheless, the species had a highly variable diet, making it ageneralist. It may have eaten lower quality, harder foods, such as nuts, in leaner times. To survive, children may have needed nursing during such periods until reaching perhaps 4 to 5 years of age. The species appears to have beenpatrifocal, with females more likely to leave the group than males.A. africanus lived in agallery forest surrounded by more open grasslands or bushlands. South African australopithecine remains probably accumulated in caves due to predation by large carnivores (namelybig cats), and the Taung child appears to have been killed by abird of prey.A. africanus probably went extinct due to major climatic variability and volatility and possibly competition withHomo andP. robustus.
In 1924, Australian anatomist ProfessorRaymond Dart, since 1923 working in South-Africa, was informed by one of his students, Josephine Salmons, that monkey fossils (ofPapio izodi) had been discovered by shotfirer M.G. de Bruyn in alimestonequarry inTaung, South Africa, operated by the Northern Lime Company. Knowing that Scottish geologist ProfessorRobert Burns Young was at the time carrying out excavations in the area in search ofarchaic human remains likeHomo rhodesiensis fromKabwe, Zambia (at the time Broken Hill,Northern Rhodesia) discovered in 1921, he asked his colleague to send him some primate remains from the quarry.[2]: 284–285
On 24 November 1924, Dart received two boxes with fossils collected by De Bruyn. In them, he noticed a natural brain endocast and a face of a, now known to be 2.8 million year old, juvenile skull, theTaung child, that he immediately recognised as atransitional fossil between apes and humans. Most notably, it had a small brain size yet was, as shown by the position of theforamen magnum,bipedal. Dart, after hastily freeing the fossil from its matrix, already in January 1925 named the specimen as a new genus and species:Australopithecus africanus.[2]: 284–285
At the time of discovery,great apes were classified into thefamilyPongidae encompassing all non-human fossil apes, andHominidae encompassing humans and ancestors. Dart felt the Taung child fit into neither, and erected the family "Homo-simiadæ" ("man-apes").[2]: 285–286 This family name was soon abandoned, and Dart proposed "Australopithecidae" in 1929. In 1933, South African palaeoanthropologistRobert Broom suggested movingA. africanus into Hominidae, which at the time contained only humans and their ancestors.[2]: 285
A. africanus was the first evidence that humans evolved in Africa, asCharles Darwin had postulated in his 1871The Descent of Man. However, Dart's claim of the Taung child as the transitional stage between apes and humans was at odds with the-then popular model of human evolution which held that large brain size and humanlike characteristics had developed rather early on, and that large brain size evolved before bipedalism. Resultantly,A. africanus was generally cast aside as a member of thegorilla orchimpanzee lineages, most notably by SirArthur Keith.[2]: 285–288
This view was perpetuated byCharles Dawson's 1912 hoaxPiltdown Man hailing from Britain. Further, the discovery of the humanlikePeking Man (Homo erectus pekinensis) in China also seemed to place the origins of humankind outside of Africa. Humanlike characteristics of the Taung child were attributed to the specimen's juvenile status, meaning they would disappear with maturity. Nonetheless, Dart and Broom continued to argue thatAustralopithecus was far removed from chimpanzees, showing several physical and claiming some behavioural similarities with humans.[2]: 285–288 To this extent, Dart made note of the amalgamations of large mammal bone fragments in australopithecine-bearing caves which are now attributed tohyena activity.[3] However, Dart proposed that the bones were instead evidence of what he named the "osteodontokeratic culture" produced by australopithecine hunters, who manufactured weapons using thelong bones, teeth, and horns of large hoofed prey:[4]
On this thesis man's predecessors differed from living apes in being confirmed killers: carnivorous creatures, that seized living quarries by violence, battered them to death, tore apart their broken bodies, dismembered them limb from limb, slaking their ravenous thirst with the hot blood of victims and greedily devouring livid writhing flesh.
Broom was one of the few scientists defending the close human affinities ofAustralopithecus africanus. In 1936, he was informed by two of Dart's students, Trevor R. Jones and G. Schepers, that human-like remains had been discovered in theSterkfontein Cave quarries. On 9 August 1936, he asked G.W. Barlow to provide him with any finds. On 17 August 1936 he received an adult skull including a natural endocast, specimen Sts 60. However, Broom classified it as a new species, "A. transvaalensis",[5][6] and in 1938 moved it into a new genus as "Plesianthropus transvaalensis". He also discovered the robust australopithecineParanthropus robustus, showing evidence of a wide diversity ofEarly Pleistocene "man-apes".[7] BeforeWorld War II, several more sites boreA. africanus fossils. A detailed monograph by Broom and palaeoanthropologist Gerrit Willem Hendrik Schepers in 1946 regarding these australopithecines from South Africa, as well as several papers by British palaeoanthropologist SirWilfrid Le Gros Clark, had turned around scientific opinion, garnering wide support forA. africanus' classification as a human ancestor.[2]: 289–290 In 1947, the most complete skull was discovered, STS 5 ("Mrs. Ples").[8] Wider acceptance ofA. africanus prompted re-evaluation of Piltdown Man in 1953, revealing its falsehood.[2]: 290
In 1949, Dart recommended splitting a presumed-female facial fragment fromMakapansgat, South Africa, (MLD 2) into a new species as "A. prometheus".[9] In 1954, he referred another presumed-female specimen from Makapansgat (a jawbone fragment).[10] However, in 1953, South African palaeontologistJohn Talbot Robinson believed that splitting species and genera on such fine hairs was unjustified, and that australopithecine remains from East Africa recovered over the previous couple of decades were indistinguishable from "Plesianthropus"/A. africanus. Based on this, in 1955, Dart agreed with synonymising "A. prometheus" withA. africanus because they are already quite similar to each other, and if speciation did not occur across a continent, then it quite unlikely occurred over a couple tens of kilometres according to Dart.[11] The East African remains would be split off intoA. afarensis in 1978.[12] In 2008, palaeoanthropologistRonald J. Clarke recommended reviving "A. prometheus" to house the StW 573 nearly-complete skeleton ("Little Foot"), StS 71 cranium, StW 505 cranium, StW 183maxilla, StW 498 maxilla and jawbone, StW 384 jawbone, StS 1palate, and MLD 2.[13] In 2018, palaeoanthropologistsLee Rogers Berger andJohn D. Hawks considered "A. prometheus" anomen nudum ("naked name"), and has not been properly described with diagnostic characteristics which separate it fromA. africanus.[14] At the time, these remains were dated to 3.3 million years ago in theLate Pliocene. In 2019, Clarke and South African palaeoanthropologist Kathleen Kuman redated StW 573 to 3.67 million years ago, making it the oldestAustralopithecus specimen from South Africa. They considered its antiquity further evidence of species distinction, drawing parallels withA. anamensis andA. afarensis from Middle Pliocene East Africa.[15] Little foot is the most complete early hominin skeleton ever recovered, with about 90% preserved.[16]
In addition to Taung, Sterkfontein, and Makapansgat,A. africanus was in 1992 discovered inGladysvale Cave. The latter three are in theCradle of Humankind.[17] Many hominin specimens traditionally assigned toA. africanus have been recovered from Sterkfontein Member 4 (including Mrs. Ples and 2 partial skeletons), previously dated to 2.8 to 2.15 million years ago. But in 2022 a team including Clarke and Kuman used cosmogenic nuclide techniques to date Member 4 at 3.4 million years, which it says discredits the assumption thatA. africanus descended fromA. afarensis.[18] However, given the wide range of variation exhibited by these specimens, it is debated if all these elements can be confidently assigned to onlyA. africanus.[19][20]
At present, the classification of australopithecines is in disarray.Australopithecus is considered agrade taxon, whose members are united by their similar physiology rather than close relations with each other over other hominin genera. It is unclear howA. africanus relates to other hominins.[21] The discovery ofEarly PleistoceneHomo in Africa during the latter half of the 20th century placed humanity's origins on the continent andA. africanus as ancestral toHomo. The discovery ofA. afarensis in 1978, at the time the oldest known hominin, prompted a hypothesis thatA. africanus was ancestral toP. robustus, andA. afarensis was the last common ancestor betweenHomo andA. africanus/P. robustus.[22] It is also suggested thatA. africanus is closely related toP. robustus but not to the otherParanthropus species in East Africa,[23] or thatA. africanus is ancestral to allParanthropus.[24]A. africanus has also been postulated to have been ancestral toA. sediba which also inhabited the Cradle of Humankind, perhaps contemporaneously.A. sediba is also postulated to have been ancestral toHomo, which if correct would indeed putA. africanus in an ancestral position toHomo.[25]
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Based on 4 specimens, theA. africanus brain volume averaged about 420–510 cc (26–31 cu in). Based on this, neonatal brain size was estimated to have been 165.5–190 cc (10.10–11.59 cu in) using trends seen in adult and neonate brain size in modern primates. If correct, this would indicate thatA. africanus was born with about 38% of its total brain size, which is more similar to non-human great apes at 40% than humans at 30%.[26] Theinner ear has widesemicircular canals like non-human apes, as well as loose turns at the terminal end of thecochlea like humans. Such a mix may reflect habitual locomotion both in the trees and walking while upright because inner ear anatomy affects thevestibular system (sense of balance).[27]
A. africanus had aprognathic jaw (it jutted out), a somewhat dished face (the cheek were inflated, causing the nose to be at the bottom of a dip), and a defined brow ridge. Thetemporal lines running across either side of the braincase are raised as small crests. Thecanines are reduced in size compared to non-human apes, though still notably bigger than those of modern humans. Like other early hominins, thecheek teeth are large and feature thickenamel. In the upper jaw the thirdmolar is the largest molar, and in the lower jaw it is the second molar.A. africanus had a fast, apelike dental development rate.[2]: 293–297 According to Clarke, the older "A. prometheus" is distinguished by larger and more bulbous cheek teeth, larger incisors and canines, more projecting cheeks, more widely spacedeye sockets, and asagittal crest.[13]A. africanus has a wide range of variation for skull features, which is typically attributed to moderate to high levels ofsexual dimorphism in that males were more robust than females.[28]
In 1992, American anthropologistHenry McHenry estimated an average weight (when assuming humanlike or apelike body proportions, respectively) of 40.8 or 52.8 kg (90 or 116 lb) for males based on five partial leg specimens, and 30.2 or 36.8 kg (67 or 81 lb) for females based on seven specimens.[29] In 2015, American anthropologistWilliam L. Jungers and colleagues similarly reported an average weight (without attempting to distinguish males from females) of 30.7 kg (68 lb) with a range of 22.8–43.3 kg (50–95 lb) for weight based on 19 specimens.[30] Based on seven specimens, McHenry estimated that males, on average, grew to 138 cm (4 ft 6 in) tall and females 125 cm (4 ft 1 in).[31] In 2017, based on 24 specimens, anthropologist Manuel Will and colleagues estimated a height of 124.4 cm (4 ft 1 in) with a range of 110–142 cm (3 ft 7 in – 4 ft 8 in).[32] The elderly, probably female StW 573 was estimated to have stood about 130 cm (4 ft 3 in).[33]: 7
Based on theA. afarensis skeletonDIK-1-1, australopiths are thought to have had a humanlike spine, with 7neck vertebrae, 12thoracic vertebrae, and (based on other early australopith skeletons) 5 flexiblelumbar vertebrae.[34] In StW 573, theatlas bone in the neck, important for swiveling and stabilising the head, is more similar to non-human apes and indicates greater mobility to swivel up and down than in humans. Such motion is important forarboreal species to locate and focus on climbable surfaces. The StW 573 atlas shows similarmechanical advantages for the muscles which move theshoulder girdle as chimps andgorillas, which may indicate lesslordosis (normal curvature of the spine) inA. africanus neck vertebrae. However, the later StW 679 has some similarities to human atlases, which could potentially indicate gradual evolution away from the ape condition.[35] StW 573 has a narrowthoracic inlet unlikeA. afarensis and humans. Theclavicle is proportionally quite long, with a similar absolute length to that of modern humans.[33]: 12
Like in modern women, L3–L5 curve outwards in specimen StS 14, whereas these are straighter in StW 431 as in modern men. This probably reflects reinforcement of the female spine to aid in walking upright while pregnant.[36] The StS 14 partial skeleton preserves a rather completepelvis. Like in the restored pelvis of theLucy specimen (A. afarensis), thesacrum was relatively flat and orientated more towards the back than in humans, and thepelvic cavity had an overallplatypelloid shape. This could indicate a broadbirth canal compared to neonate head size, and thus a non-rotational birth (unlike humans), though this is debated. When standing, the angle between the sacrum and thelumbar vertebrae was reconstructed to have been about 148.7°, which is much more similar to that of chimps (154.6°) than humans (118.3°). This would indicateA. africanus standing posture was not as erect as in humans.[37]
TheA. africanus hand and arm exhibit a mosaic anatomy, with some aspects more similar to humans and others to non-human apes. It is unclear if this means australopiths were still arboreal to a degree, or if these traits were simply inherited from thehuman–chimpanzee last common ancestor. Nonetheless,A. africanus exhibits a more ape-like upper limb anatomy thanA. afarensis, and is typically interpreted as having been, to some extent, arboreal. Like in arboreal primates, the fingers are curved, the arms relatively long and the shoulders are in a shrugging position. TheA. africanus shoulder is most like that oforangutans, and well suited for maintaining stability and bearing weight while raised and placed overhead.[38] However, the right clavicle of StW 573 has a distinctly S-shaped (sigmoid) curve like humans, which indicates a humanlikemoment arm for stabilising theshoulder girdle against thehumerus.[33]: 12 TheA. africanus arm bones are consistent with powerful muscles useful in climbing. Nonetheless, the brachial index (the forearm tohumerus ratio) is 82.8–86.2 (midway between chimpanzees and humans), which indicates a reduction in forearm length from the more ancient homininArdipithecus ramidus.[16] The thumb and wrist indicate humanlike functionality with aprecision grip and forceful opposition between the thumb and fingers. The adoption of such a grip is typically interpreted as an adaptation for tool making at the expense of efficient climbing and arboreal habitation.[39]
The leg bones clearly show thatA. africanus habitually engaged in bipedal locomotion, though some aspects of thetibiae are apelike, which could indicate that the leg musculature had not been fully reorganised into the human condition. If correct, its functional implications are unclear.[16] Thetrabecular bone at thehip joint is distinctly humanlike, which would be inconsistent with the great degrees of hip loading required in prolonged arboreal activity.[40] The tibia met the foot at a similar angle as it does in humans, which is necessary for habitual bipedalism. Consequently, the ankle was not as adept for climbing activities as it is in non-human apes.[41] However, the modern CongoTwahunter–gatherers can achieve a chimp-like angle with the ankle while climbing trees due to the longer fibres in thegastrocnemius (calf) muscle instead of specific skeletal adaptations.[42] Some aspects of theankle bone were apelike which may have affected walking efficiency. The foot elements ofA. africanus are largely known from remains from Sterkfontein Member 4. The foot is humanlike with a stiff midfoot and lack of a midtarsal break (which allows non-human apes to lift the heel independently from the rest of the foot). ThoughA. africanus had anadducted big toe (it was not dextrous) like humans,A. africanus likely did not push off with the big toe, using the side of the foot instead. StW 573 is the oldest hominin specimen with an adducted big toe. The specimen StW 355 is the most curved proximal footphalanx bone of any known hominin, more similar to that of orangutans andsiamangs.[43]
The arms of StW 573 were about 53.4 cm (1 ft 9 in), and her legs 61.5 cm (2 ft 0 in). This means the arm was 86.9% the length of the leg. She is the first and only early hominin specimen to definitively show that the arms were almost all long as the legs. Nonetheless, these proportion are more similar to humans than non-human apes, withhumans at 64.5–78%,chimpanzees about 100%,gorillas 100–125%, andorangutans 135–150.9%.[33]: 17–18
In 1954, Robinson proposed thatA. africanus was ageneralistomnivore whereasP. robustus was a specialised herbivore; and in 1981, American palaeoanthropologistFrederick E. Grine suggested thatP. robustus specialised on hard foods such as nuts whereasA. africanus on softer foods such as fruits and leaves. Based oncarbon isotope analyses,A. africanus had a highly variable diet which included a notable amount ofC4 savanna plants such as grasses, seeds,rhizomes, undergroundstorage organs, or perhaps grass-eating invertebrates (such aslocusts ortermites),grazing mammals, orinsectivores or carnivores. Most primates do not eat C4 plants.[44][45] More recent carbon and nitrogen isotope analysis ofA. africanus specimens from Sterkfontein has suggested that mammal meat was not a significant component of their diet.[46]A. africanus facial anatomy seems to suggest adaptations for producing high stress on the premolars, useful for eating small, hard objects such as seeds and nuts that need to be cracked open by the teeth, or for processing a large quantity of food at one time. However, like forP. robustus, microwear analysis on the cheek teeth indicate small, hard foods were infrequently eaten, probably as fall back foods during leaner times.[47] Still,A. africanus, like chimps, may have requiredhammerstones to crack open nuts (such asmarula nuts), thoughA. africanus is not associated with any tools.[44]
A. africanus conspicuously lacks evidence of dentalcavities, whereasP. robustus seems to have had a modern humanlike cavity rate;[48] this could possibly indicate thatA. africanus either did not often consume high-sugar cavity-causing foods—such as fruit, honey, and some nuts and seeds—or frequently consumed gritty foods which decrease cavity incidence rate.[49] However, the 2nd right permanent incisor (STW 270) and right canine (STW 213) from the same individual show lesions consistent withacid erosion, which indicates this individual was regularly biting into acidic foods such as citrus. Tubers could have caused the same damage if some chewing was done by the front teeth.[50]
Barium continually deposits ontoA. africanus teeth until about 6–9 months of development, and then decreases until about 12 months. Because the barium was most likely sourced from breast milk, this probably reflects the weaning age. This is comparable to the human weaning age. Following this initial period, barium deposits stall and then restart cyclically every year for several years. In the first molar specimen StS 28 (from Sterkfontein), this occurred every 6–9 months, and in the lower canine specimen StS 51 every 4–6 months, and this carried on until 4–5 years of development.Lithium andstrontium also deposit cyclically. Cyclical barium, lithium, and strontium bands occur in modern primates—for example, wild orangutans up to 9 years of age—which is caused by seasonal famine when a child has to rely on nursing to sustain themselves and less desirable fallback foods. However, it is unclear if this can be extended toA. africanus.[51]
The group dynamics of australopithecines is difficult to predict with any degree of accuracy. A 2011strontium isotope study ofA. africanus teeth from thedolomite Sterkfontein Valley found that, assuming that especially small teeth represented female specimens and especially large teeth males, females were more likely to leave their place of birth (patrilocal). This is similar to the dispersal patterns of modern-day hominins which have a multi-male kinship-based society, as opposed to theharem society of gorillas and other primates. However, the small canines of males compared to those of females would seem to suggest a much lower degree of male–male aggression than non-human hominins. Males did not seem to have ventured very far from the valley, which could either indicate small home ranges, or that they preferred dolomitic landscapes due to perhaps cave abundance or factors related to vegetation growth.[52]
In a sample of tenA. africanus specimens, seven exhibited mild to moderatealveolar bone loss resulting from periodontal disease (the wearing away of the bone which supports the teeth due to gum disease).[53] The juvenile specimen STS 24a was diagnosed with an extreme case of periodontal disease on the right side of the mouth, which caused pathological bone growth around the affected site, and movement of the first two right molars during cyclical periods of bacterial infection and resultant inflammation. Similarly, the individual appears to have preferred to chew using the left side of the jaw. The periodontal disease would have severely hindered chewing, particularly in the last year of life, and the individual potentially may have relied on group members to survive for as long as it did.[54]
In 1992, anthropologists Geoffrey Raymond Fisk and Gabriele Macho interpreted the left ankle bone Stw 363 as bearing evidence of a healedcalcaneal fracture on the heel bone (which was not preserved), which they believed resulted from a fall from a tree. If correct, then the individual was able to survive for a long time despite losing a great deal of function in the left leg. However, they also noted that similar damage could potentially have also been inflicted bycalcite deposition and crystallisation during the fossilisation process. Calcaneal fractures have been recorded in humans, and are present quite often in arboreal primates.[55]
South African australopithecines appear to have lived in an area with a wide range of habitats. At Sterkfontein,fossil wood belonging to thelianaDichapetalum cf. mombuttense was recovered. The only living member of this tree genus in South Africa isDichapetalum cymosum, which grows in dense, humidgallery forests. In modern day,D. mombuttense only grows in theCongolian rainforests, so its presence could potentially mean the area was an extension of this rainforest. The wildlife assemblages indicate a mix of habitats such asbush savanna, openwoodland, or grassland. TheshrubAnastrabe integerrima was also found, which today only grows on the wetter South African coastline. This could indicate the Cradle of Humankind received more rainfall in the Plio-Pleistocene. In total, the Cradle of Humankind may have featured gallery forests surrounded by grasslands.[56] Taung also appears to have featured a wet, closed environment.[57]Australopithecines and earlyHomo likely preferred cooler conditions than laterHomo, as there are no australopithecine sites that were below 1,000 m (3,300 ft) in elevation at the time of deposition. This would mean that, like chimps, they often inhabited areas with an average diurnal temperature of 25 °C (77 °F), dropping to 10 or 5 °C (50 or 41 °F) at night.[58]
In 1983, studyingP. robustus remains, South African palaeontologistCharles Kimberlin Brain hypothesised that australopithecine bones accumulated in caves due to large carnivore activity, dragging in carcasses. He was unsure if these predators actively sought them out and brought them back to the cave den to eat, or inhabited deeper recesses of caves and ambushed them when they entered.Baboons in this region modern day often shelter insinkholes especially on cold winter nights, though Brain proposed that australopithecines seasonally migrated out of theHighveld and into the warmerBushveld, only taking up cave shelters in spring and autumn.[59] TheA. africanus fossils from Sterkfontein Member 4 were likely accumulated bybig cats, thoughhunting hyenas andjackals may have also played a role.[60] Scratches, gouges, and puncture marks on the Taung child similar to those inflicted by moderncrowned eagles indicate this individual was killed by abird of prey.[61][62]
Around 2.07 million years ago, just before the arrival ofP. robustus andH. erectus,A. africanus became extinct in the Cradle of Humankind. It is possible that South Africa was arefuge forAustralopithecus until the beginning of major climatic variability and volatility, and, perhaps, competition withHomo andParanthropus.[63]
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