Fossil Primates and the Earliest Hominins: The Oligocene and Miocene Apes from Parapithecus and Aegyptopithecus to Dryopithecus, Sivapithecus, Ramapithecus and Gigantopithecus, the Pre-Hominins Sahelanthropus, Orrorin and Ardipithecus, the Australopithecines and Paranthropus, and Homo habilis
2. The pre-hominins: Sahelanthropus, Orrorin and Ardipithecus
Molecular clocks place the separation of the human and chimpanzee lineages at roughly six to eight million years ago, and three fossils fall in or near that window. Sahelanthropus tchadensis, nicknamed Toumaï ("hope of life" in the Goran language), is a nearly complete but crushed cranium found in 2001 at Toros-Menalla in the Djurab desert of Chad by Michel Brunet's team and published in 2002; dated to about seven (six to seven) million years, it combines a small, ape-sized brain (about 360 cc) with a flat face, small canines, thick enamel and a foramen magnum placed forward enough to suggest an upright posture — though a 2020 study of its femur questioned bipedalism. It is the earliest candidate hominin, and the first from west of the Rift Valley, which overturned the "East Side Story" that tied hominin origins to the Rift. Orrorin tugenensis ("original man" in Tugen), found in the Tugen Hills of Kenya in 2000 by Brigitte Senut and Martin Pickford and called "Millennium Man", is about six million years old and known from teeth, jaw fragments and three femora whose neck and head suggest bipedal walking, together with curved finger bones for climbing. Ardipithecus ramidus, from Aramis in the Middle Awash of Ethiopia, was announced by Tim White, Gen Suwa and Berhane Asfaw in 1994 as Australopithecus ramidus, moved to the new genus Ardipithecus in 1995, and described in full in 2009 from the 4.4-million-year-old partial skeleton "Ardi" (ARA-VP-6/500): a woodland-living form about 1.2 m tall, with a small brain (300–350 cc), an opposable big toe for climbing, a pelvis adapted to upright walking on the ground, and small canines without honing in males — read as evidence that bipedalism arose in woodland rather than savanna and that the last common ancestor was not chimpanzee-like. The older Ardipithecus kadabba (5.8–5.2 million years) precedes it.
3. The gracile australopithecines: afarensis, africanus and bahrelghazali
Australopithecus ("southern ape") was named by Raymond Dart in 1925 for the Taung child, a juvenile skull with a natural brain endocast found in a limestone quarry in South Africa in 1924: a small brain, but a forward foramen magnum and human-like teeth. Dart's claim of an African ape-man was ridiculed for twenty years until Robert Broom's finds at Sterkfontein (the adult "Mrs Ples", Sts 5, 1947) and elsewhere confirmed Australopithecus africanus, now dated about 3.3–2.1 million years, with a brain of 430–520 cc, a projecting face and a bipedal pelvis. In East Africa, Donald Johanson's team found "Lucy" (AL 288-1), a 40%-complete female skeleton about 1.1 m tall, at Hadar in the Afar region of Ethiopia in 1974, and the "First Family" (AL 333) in 1975; with Mary Leakey's finds at Laetoli in Tanzania — where the 3.66-million-year-old footprint trails discovered in 1978 record fully bipedal walking — these were named Australopithecus afarensis by Johanson, White and Coppens in 1978. Dated about 3.9–2.9 million years, afarensis has a brain of 380–430 cc, a prognathic face, canines larger than ours but reduced, a dental arcade between U and parabolic, long arms and curved fingers alongside a bipedal pelvis and knee, and marked sexual dimorphism; "Selam", the Dikika child (2000), and the 2016 study of Lucy's fall from a tree belong to it. Australopithecus bahrelghazali ("Abel"), a mandible found by Brunet at Koro Toro in Chad in 1995 and dated about 3.5 million years, is the only australopithecine from outside eastern and southern Africa; many regard it as a western population of afarensis. Australopithecus anamensis (Kenya, 4.2–3.9 million years; Meave Leakey, 1995) precedes afarensis; A. garhi (2.5 million years, Ethiopia) and A. sediba (about 2 million years, Malapa, South Africa, 2008) follow.
| Species | Type find | Date | Brain (cc) | Key traits |
|---|---|---|---|---|
| A. afarensis | Lucy, Hadar 1974 (Johanson); Laetoli footprints 1978 (M. Leakey); named 1978 | 3.9–2.9 mya | 380–430 | Bipedal pelvis and knee; long arms, curved fingers; strong dimorphism; ~1.1 m (Lucy) |
| A. africanus | Taung child 1924 (Dart, 1925); Mrs Ples, Sterkfontein 1947 (Broom) | 3.3–2.1 mya | 430–520 | Rounder skull, less prognathic than afarensis; South Africa only |
| A. bahrelghazali | Abel, Koro Toro, Chad 1995 (Brunet) | ~3.5 mya | — (mandible only) | First australopithecine west of the Rift; perhaps a regional afarensis |
| Paranthropus robustus | Kromdraai 1938, Swartkrans (Broom) | ~2.0–1.2 mya | ~500–530 | Sagittal crest, huge molars, flat face; South Africa |
| Paranthropus boisei | OH 5 "Zinjanthropus", Olduvai 1959 (M. Leakey) | ~2.3–1.2 mya | ~500–550 | "Nutcracker man"; the most massive chewing apparatus; East Africa |
| Homo habilis | OH 7, Olduvai 1960 (J. Leakey); named 1964 (L. Leakey, Tobias, Napier) | ~2.4–1.6 mya | 510–700 | Larger brain, smaller teeth; Oldowan tools; "handy man" |
4. The robust forms: Paranthropus robustus and boisei
Between about 2.7 and 1.2 million years ago a second australopithecine lineage specialised in heavy chewing. Its skulls carry a sagittal crest for the temporalis muscles, flaring cheekbones, a dished face, small incisors and canines, and enormous, thick-enamelled premolars and molars; the brain stayed at 500–550 cc. Robert Broom found the first, at Kromdraai in South Africa in 1938, and named it Paranthropus robustus ("beside man"); more came from Swartkrans. Mary Leakey found the East African form at Olduvai Gorge in 1959: the skull OH 5, called Zinjanthropus boisei by Louis Leakey (after Charles Boise, who funded the work) and "Nutcracker Man" and "Dear Boy" by the press, dated by the new potassium-argon method to 1.75 million years — the date that first showed how old the hominin record was. The still more primitive "Black Skull" (KNM-WT 17000, Paranthropus aethiopicus, 2.5 million years, west of Lake Turkana, 1985) links the robust line to afarensis. Whether the robust forms are a genus of their own (Paranthropus, the usage the syllabus follows in parentheses) or robust species of Australopithecus is a matter of taxonomic taste; what is agreed is that they are a side branch that went extinct about 1.2 million years ago, overlapping with early Homo and probably eating hard or fibrous plant foods, though isotopes show boisei ate mostly C4 grasses and sedges. The syllabus's spelling "Paranthropous" is a misprint of Paranthropus.
5. Homo habilis, the "early transitional human"
In 1960 Jonathan Leakey found a juvenile mandible, parietals and hand bones (OH 7) in Bed I at Olduvai Gorge, close to the Oldowan pebble tools that Louis Leakey had long attributed to a toolmaker more advanced than Zinjanthropus. In 1964 Louis Leakey, Phillip Tobias and John Napier named Homo habilis, "handy man" (Dart suggested the name), on a brain estimated at about 640–680 cc — below the 750 cc "cerebral Rubicon" that Arthur Keith had set for Homo, so they lowered the threshold to about 600 cc — together with smaller cheek teeth, a more parabolic arcade, a hand capable of the precision grip and a foot (OH 8) with arches. Later finds include the 1.9-million-year-old skull KNM-ER 1470 from Koobi Fora (Richard Leakey's team, 1972; about 750 cc), which many separate as Homo rudolfensis, and OH 62 (1986), whose ape-like limb proportions surprised those who expected a more human body. Dated about 2.4–1.6 million years and known only from Africa (Olduvai, Koobi Fora, Omo, Hadar, Sterkfontein), habilis is the syllabus's "early transitional human": it retains australopithecine body size and limb proportions while showing the enlarged brain, reduced dentition and (by association) stone tool-making that define Homo. The 2015 description of a 2.8-million-year-old Homo mandible from Ledi-Geraru in Ethiopia pushes the genus back further, and the debate whether habilis is a true Homo or should be returned to Australopithecus (Wood and Collard, 1999) is still open. Its successor, Homo erectus, is the subject of the next chapter.
Key takeaways
- Aegyptopithecus (Fayum, ~30 mya) is the stem catarrhine; Dryopithecus (Europe, ~12–9 mya) names the Y-5 pattern; Sivapithecus and Ramapithecus (Siwaliks, ~12–8 mya) are now one orangutan-line ape, though Simons and Pilbeam once made Ramapithecus the first hominid; Gigantopithecus is the largest primate, known from teeth and jaws.
- Pre-hominins: Sahelanthropus tchadensis (Toumaï, Chad, 2001, ~7 mya, Brunet), Orrorin tugenensis (Kenya, 2000, ~6 mya, Senut and Pickford) and Ardipithecus ramidus (Aramis, Ethiopia, 4.4 mya, White; named Australopithecus ramidus in 1994 and moved to Ardipithecus in 1995).
- A. afarensis (Lucy, Hadar 1974; Laetoli footprints 3.66 mya; 3.9–2.9 mya) and A. africanus (Taung 1924, Dart 1925; Sterkfontein; 3.3–2.1 mya) are the gracile forms; A. bahrelghazali (Chad, 1995) is the only one west of the Rift.
- The robust forms, Paranthropus (syllabus "Paranthropous") robustus (Kromdraai 1938, Broom) and boisei (Zinjanthropus, Olduvai 1959, Mary Leakey, K-Ar dated 1.75 mya), have sagittal crests and huge cheek teeth on a 500–550 cc brain and died out about 1.2 mya.
- Homo habilis (OH 7, Olduvai 1960; named 1964 by Leakey, Tobias and Napier; 2.4–1.6 mya; 510–700 cc) is the transitional form: australopithecine body, enlarged brain, reduced teeth, Oldowan tools; KNM-ER 1470 is often separated as H. rudolfensis.
Practice questions (10)
Attempt each one before opening the answer. Every explanation names the tempting wrong option as well as the right one, because that is where marks are lost.
"Lucy" (AL 288-1), the partial skeleton found at Hadar, Ethiopia, in 1974, belongs to
Show answer
Answer: B — Australopithecus afarensis
Donald Johanson's team found Lucy at Hadar in the Afar region; with the Laetoli material she became the type of A. afarensis, named by Johanson and White in 1978 and dated about 3.2 million years.The Taung child, on which Raymond Dart founded the genus Australopithecus in 1925, was found in
Show answer
Answer: B — South Africa
The juvenile skull came from a limestone quarry at Taung in South Africa in 1924. Ethiopia has Hadar and Aramis, Tanzania Olduvai and Laetoli, Chad Toumaï and Abel.In which year was Zinjanthropus boisei (OH 5), now Paranthropus boisei, discovered by Mary Leakey at Olduvai Gorge? Give the year.
Numerical answer — type the value.
Show answer
Answer: 1959
Mary Leakey found the "Nutcracker Man" skull in Bed I at Olduvai on 17 July 1959; its potassium-argon date of 1.75 million years, published in 1961, was the first radiometric date for an early hominin.Match the fossil with its site.
(a) Sahelanthropus tchadensis
(b) Orrorin tugenensis
(c) Ardipithecus ramidus
(d) Australopithecus bahrelghazali. Sites:
(1) Aramis, Middle Awash
(2) Koro Toro
(3) Toros-Menalla
(4) Tugen HillsShow answer
Answer: D — a-3, b-4, c-1, d-2
Toumaï came from Toros-Menalla in Chad's Djurab desert (2001), Orrorin from Kenya's Tugen Hills (2000), Ardipithecus from Aramis in Ethiopia's Middle Awash (1994), and Abel from Koro Toro in Chad (1995).Ramapithecus, found in the Siwalik deposits at Haritalyangar and once proposed as the earliest hominid, is now generally regarded as
Show answer
Answer: A — a smaller form of Sivapithecus, an ape on the orangutan lineage
Simons and Pilbeam's 1960s hominid reading rested on small canines and thick enamel; the complete Sivapithecus faces from Pakistan (1979–80) and molecular dating of the human–ape split to 6–8 million years placed Ramapithecus within Sivapithecus on the orangutan line.Which features distinguish the robust australopithecines (Paranthropus) from the gracile ones? Select all that apply.
Show answer
Answer: A — A sagittal crest for enlarged chewing muscles; B — Very large, thick-enamelled premolars and molars with small incisors and canines
The robust forms are defined by the chewing apparatus — crest, flaring cheekbones, huge cheek teeth — while their brains stayed at 500–550 cc and they remained bipedal like all australopithecines.Homo habilis was formally named in 1964 by
Show answer
Answer: D — Louis Leakey, Phillip Tobias and John Napier
The 1964 Nature paper by Leakey, Tobias and Napier named "handy man" on OH 7 and related Olduvai material, arguing that a brain near 650 cc, smaller teeth and a precision-grip hand set it in Homo. Johanson and White named A. afarensis in 1978.Which statements about Ardipithecus ramidus are correct? Select all that apply.
Show answer
Answer: A — The 4.4-million-year-old partial skeleton "Ardi" combines a grasping big toe with a pelvis adapted to upright walking; B — It was first published in 1994 as Australopithecus ramidus and transferred to a new genus in 1995; D — Its males lacked the large honing canines of chimpanzees
The Aramis fauna and flora show a woodland habitat, which is why Ardi is cited against the savanna hypothesis; the naming history, the mosaic foot and pelvis, and the small non-honing canines are correct.Assertion (A): The discovery of Sahelanthropus tchadensis in Chad in 2001 confirmed the "East Side Story" of hominin origins. Reason (R): Toumaï, dated to about seven million years, was found some 2,500 km west of the East African Rift Valley, to which the hypothesis had tied the emergence of bipedal hominins.
Show answer
Answer: D — A is false, but R is true
A is false: Coppens's hypothesis had hominins arising in the dry country east of the Rift while apes stayed in the forests to the west, and a seven-million-year-old candidate hominin in Chad contradicts that geography, so the find weakened the hypothesis rather than confirming it. R is true: Michel Brunet's team found Toumaï at Toros-Menalla in the Djurab desert, some 2,500 km west of the Rift.Arrange these in chronological order from oldest to youngest:
(a) Homo habilis
(b) Sahelanthropus tchadensis
(c) Australopithecus afarensis
(d) Ardipithecus ramidus
(e) Australopithecus anamensisShow answer
Answer: D — b, d, e, c, a
Sahelanthropus (~7 mya) precedes Ardipithecus ramidus (4.4 mya), then A. anamensis (4.2–3.9 mya), then A. afarensis (3.9–2.9 mya), which anamensis is thought to have given rise to; Homo habilis appears about 2.4 mya.