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Early Hominids
Learning Outcomes
At the end of this lecture you should be able to:
Introduction
This lecture is about early hominids. This title needs to be treated as a colloquialism. It is
what everyone calls the group of animals that will be discussed but it really does not fit with
any rational classification. Early Hominins would be a better title but it is such a mouthful
that no one actually uses it. The group contains the direct human ancestors from the point of
divergence with the chimpanzee lineage which molecular (and morphological) evidence
suggests occurred approximately 6 million years ago.
The period covered is known as the Plio-pleistocene since it covers both the Pliocene and the
Pleistocene. That is the last 5 million years up to about 10,000 years ago by which time most
species on Earth are pretty much as they are today (certainly in terms of morphology). In this
lecture I will introduce the species in approximately chronological order and investigate how
they might all be related. I will also look at the evolutionary pressures that may have led to
the two major features of modern humans: habitual bipedalism and huge brains.
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Background
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Earliest Hominids
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Figure 4 shows some of the major finds and their locations. There are several general features
to note.
Firstly, the fossils are all named with a site number (or occasionally a popular name for the
really famous fossils although they have accession numbers too). This is to remove ambiguity. Fossils get assigned to a variety of different species as people alter classifications but the
site number will never change. If I talk about WT-17,000 (for West Turkana fossil number
17,000) then I can only be talking about a certain fossil whereas if I say the Paranthropus
aethipicus cranium it assumes that we all agree that species designation which is certainly not
the case.
Secondly, the sites. There are two main areas where almost all the early hominid material is
found. The first site is the East African rift valley system (1,200 miles long) associated with
mountain building, faulting and vulcanism over last few million years. Earth movement mean
sediments get exposed (Plio-Pleistocene 4 to 1 mya), and volcanic activity causes layers of
volcanic ash (tuffs) which can be dated (potassium argon, or fission tracks) accurately. Here
is a brief description of some of the localities within the Rift Valley:
Hadar
Very many fossils and artefacts. Fossils from 3.9 to 3 mya. Most famous is Lucy (Afar
Locality (AL) 288-1, Australopitheucs afarensis), a 40% complete skeleton (one of only 2
hominid skeletons earlier than 100,000 ya). AL 333 is a group of over 13 individuals including 4 infants: a catastrophic assemblage perhaps.
Omo
Very thick continuous sequence (0.5 mile thick). 2.9 to 1 mya. Very rich fauna, so useful for
biostratigraphic dating, but fossil hominids restricted to teeth and bone fragments.
East Turkana
Possibly the richest site. Approx. 1.8 mya, though there are some much older beds (3.3 mya).
Complete skulls, jaws and postcrania.
West Turkana
West side of Lake Turcana. 2 very famous finds: an almost complete, 1.6 mya Homo erectus
adolescent (see later) and the so called black skull (Paranthropus aethiopicus), a 2.4 mya
robust australopithecine skull that is still causing problems with classification!
Olduvai Gorge
Mini Grand Canyon. 2 mya to present, providing an excellent sequence of fossils and artefacts, including the original robust australopithecine cranium (Paranthropus bosei).
Laetoli
3.75 to 3.5 mya. Most famous for the Laetoli footprints: thousands of footprints of over 20
different species including hominids. One set of hominid tracks are of 2-3 individuals in a
trail more than 25 m long - bipedal walking such as this is considered to be characteristic of
hominids
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The other main area is South Africa. Unlike the East African deposits, the South African ones
are all cave deposits - piles of mineralised sediment that has fallen into caves. They are not
currently datable to any great accuracy and the ages are usually inferred by looking at the
faunal context of the fossil and comparing it to faunal remains from better dated sites. However, they have produced many fossils including some good postcranial material (including
pelvises and foot bones: important for investigating bipedalism)
Taung
Limestone mine which produced the Taung Child in the 1920s (Australopithecus africanus). Before this, it was always thought that the earliest humans would be discovered in
Europe, or perhaps the Far East. Consequently, it took quite a while before this discovery was
accepted for what it was.
Makapansgat
Another cave
Ardipithecus
Ardipithecus ramadus
This is the oldest of the group and is considered to be close to the split point between chimpanzees and humans. We do not have a great deal of fossil material to work on but due to the
presence of certain ape-like features (thin enamel, narrow molars) some people consider it to
be an early chimp ancestor (i.e. on the chimp side of the split). However, it does appear to be
upright due to the forward placement of the foramen magnum.
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morphology is thought to be diet related so this may have allowed sufficient niche separation
to prevent exclusion.
Australopithecus afarensis
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but sexual dimorphism seems the most likely. We have an extremely good postcranial skeleton (Lucy) and recently a decent cranium has also been found. This animal is assumed to be
the source of the hominid bipedal trackways found at Laetoli.
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There is some dispute about the nature of A. afarensis bipedality. The trackway shows a very
humanlike gait with the big toes in line with the other digits. Analysis of A. afarensis foot
bones has suggested that the pollux may have been divergent and that the digits are long and
curved. This has led to speculation that A. afarensis, whilst able to walk upright, still spent a
great deal of time arboreally.
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Australopithecus africanus
Australopithecus anamensis
This is a relative recent addition to the Australopithecus genus. It has been found in several
sites in East Africa and is between 3.9 and 4.2 mya. There is a very small amount of postcranial material but what there is has been described as showing evidence of bipedality and
sexual dimorphism.
Australopithecus bahrelghazali
What can I say? A few bits of upper and lower jaw and teeth from Chad. They are dated at 3
to 3.5 mya and show that there were australopithecines further West than just the East African rift valley.
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Paranthropus
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all are more recent than the earliest graciles and this suggest that they are a now extinct
offshoot from the direct human lineage.
Paranthropus bosei
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Paranthropus aethiopicus
This is the oldest robust australopithecine. It has been described as hyper robust, but also as
intermediate between A. afarensis and the other robusts. It dates from 2.3 to 2.7 mya.
Paranthropus robustus
Bipedalism
The main evolutionary question of this period is the one of bipedalism. Australopithecines do
have quite big brains but not really any bigger than a modern chimpanzee and the real push to
increase brain size seems to have occurred later on. However, the evidence suggest that
australopithecines were fully capable bipeds (even if some species still spent some time in
trees). The question is therefore what caused this change to habitual bipedalism.
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Phylogeny
So how do these groups link up to modern humans? We will come back to this when we talk
about modern humans but as you might expect there are plenty of alternatives even when just
considering the australopithecines.
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This one however does and obviously favours the A. africanus sideshoot. This is a good tree
since it attempts to go all the way through to modern humans so we will come back to it
again.
Stop Press
I wrote this lecture in 2000 and of course Science does not stand still. There have been
several major fossil finds since then. These do not effect the basic conclusions but they have
had a great impact on the details especially in terms of timing the chimp/human common
ancestor and the numbers of species involved.
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Bibliography
Aiello, L.C., Collard, M. Our newest oldest ancestor? Nature 410: 526-527.
Behrensmeyer, A.K. 1992. Fossil deposits and their investigation. In: The Cambridge Encyclopedia of Human Evolution, eds. Jones, S., Martin, R., Pilbeam, D. pp. 187-190. Cambridge University Press: Cambridge.
Britten, R.J. 2002. Divergence between samples of chimpanzee and human DNA sequences
is 5%, counting indels. Proc. Natl. Acad. Sci. Online at http://www.pnas.org/
Brunet, M., Guy, F., Pilbeam, D., Mackaye, H.T., Likius, A., Ahounta, D., Beauvilain, A.,
Blondel, C., Bocherens, H., Boisserie, J.R., De Bonis, L., Coppens, Y., Dejax, J., Denys, C.,
Duringer, P., Eisenmann, V.R., Fanone, G., Fronty, P., Geraads, D,. Lehmann, T., Lihoreau,
F., Louchart, A., Mahamat, A., Merceron, G., Mouchelin, G., Otero, O., Campomanes, P.P.,
De Leon, M.P., Rage, J.C., Sapanet, M., Schuster, M., Sudre, J., Tassy, P., Valentin, X.,
Vignaud, P., Viriot, L., Zazzo, A., Zollikofer, C. A new hominid from the Upper Miocene of
Chad, central Africa. Nature 418 (6894): 145-151.
Conroy, G. C. 1990. Primate Evolution. London: Norton.
Fleagle, J.G. 1999. Primate Adaptation and Evolution. London: Academic Press.
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