During the Early and Middle Miocene eras, dating from 20 to 10 million years ago,
Africa
had a much higher annual rainfall level than today. A single super-rainforest covered most of the continent
from shore to shore. A hominoid (primate) ancestor common to all living
apes and humans lived in
the trees of this super-rainforest.
Africa first became
connected to Eurasia around 18 million years ago, causing major geological
upheavals, and transforming formerly flat landscapes by pushing up some of
the world's
youngest mountain ranges: the Pyrenees, the Alps, and the Zagros
mountains, as well as the mountains and rift valleys of East Africa.
Geographical transformations were accompanied by gradual climatic and
environmental changes which may even have triggered the emergence of the
first specialised hominoids.
The very earliest known representatives of the hominoids have been found only
in Africa and were very different from living apes and humans. The oldest
finds belong to a group of species in the genus Proconsul. Fossils
of Proconsul have been found in Early Miocene deposits 22 million
years old in Kenya, and along the rift valley, although it probably
originated rather earlier, between 25 and 28 million years ago.
Middle Miocene coastline
Proconsul's skeletons are described as generalised in that it shows none of the
particular features such as thickened tooth enamel or adaptation for
knuckle-walking which characterised the later apes. It was adapted to
living in trees and was about the size of a gibbon.
Proconsul africanus
Proconsul suffered somewhat during Africa's collision with Eurasia.
Volcanic activity reached a peak 18 million years ago, especially in East
Africa. The Kisingiri volcano erupted, apparently with a pyroclastic flow,
killing whole populations of Proconsul which lived in an
environment that was semi-arid, covered mostly in dry, deciduous,
single-canopy woodland, with some evergreen forest in restricted,
low-lying areas.
Despite repeated volcanic activity around this period, the species persisted
into the Middle Miocene (15 to 10 million years) when it overlapped with
Kenyapithecus.
The appearance of Kenyapithecus between 15 to 11 million years ago marked the
point at which some specialisations began to appear. This hominoid also
lived exclusively in the rainforests of the rift valley, and showed modifications in the teeth and limb bones,
making them more like those of the living great apes.
Kenyapithecus africanus
can be regarded as being part of the combined great ape and human group but cannot be
linked directly to any one of the living hominoids.
However, evidence has emerged which strongly suggests that
Kenyapithecus is not one genus but two. Kenyapithecus wickeri
appears to have migrated out of Africa at the same time as many other
species and lain the roots of the later European apes. Kenyapithecus
africanus has been rechristened by some scholars as Equatorius
africanus, because the Equator is where all its remains have been
discovered.
Kenyapithecus africanus
By around 14 million years ago there were other hominoid variations in
existence, and these had migrated out of Africa, with some representatives
spreading east across into the Asian continent. By 14 to 11 million years
ago Ramapithecus and Sivapithecus were in Anatolia and
Pakistan (and had reached South Asia by 8 million years ago where they
lived alongside a sister-species, the enormous
Gigantopithecus).
They are
closely related to the orang-utan, and differed from each other only in
size, suggesting that they were also closely related to each other and may
perhaps be males and females of the same species.
Both also share some of
the characteristics of Kenyapithecus, as well as particular
features which show they must be related exclusively to the orang-utan and
not to humans. This evidence indicates that the orang-utan was the first
to split away from the general hominoid pool in Africa.
Ramapithecus & Sivapithecus
Dryopithecus emerged in Africa and Europe at around the same time (and
persisted in Central and North Western Europe into the Late Miocene).
The connection between the two continents was more direct than it is
today (see map, above). Much
of the bulk of southern Europe was still a relatively thin corridor of
land no more than twice the width of modern Italy in most places, while
the Tethys Sea was twice as big as the Mediterranean Sea it would later
become.
Dryopithecus remains have only been discovered in the past
thirty years or so, and the creature, which was more like Proconsul,
is highly unlikely to be related to any of the living hominoids, but there
is a suspicion that it could have a place in early hominid ancestry.
Dryopithecus
10-7 million
At around 10 million years ago a large number of the first apes had
blossomed into a huge radiation of species that reached across Europe and
much of Asia. Over subsequent time the majority of them became
extinct, mostly due to climate changes and extensive glaciation.
Further geological upheavals now occurred as the
sub-continent of India moved north to collide with Asia. This created the Himalayas, causing weather patterns to change.
Massively increased rainfall in India (the Monsoon) stripped the air of
moisture so that the air currents that reached Africa were no longer wet,
but dry.
In East Africa this caused drier conditions during the
Late Miocene era (10 to 5 million years ago). Temperatures began to
increase, resulting in a spread of grasslands as the
super-rainforest began to die back over the next 5 million years. Animals had to adapt in response to
these changes.
Between 7.0 to 5.8 million years ago, hominid transitional species
appeared as the ancestors of man and
chimpanzee divided. Hominid refers to the family of primates that includes
all species on the "human" side of the evolutionary tree after that split. The chimpanzee's ancestors remained living in the
remnants of the great forests, while early hominids placed an increasing
reliance on surviving outside the shrinking forests.
Unfortunately, conditions in Africa between around 11 million to 5
million years ago were very detrimental to the preservation of fossils.
There are very few finds made from this period, and very little evidence
from which to build up an accurate picture of evolutionary events.
However, some finds have been made more recently...
Primitive hominoids
Earliest Hominids
6.7 million
Sahelanthropus tchadensis flourished between 6.7 to 6.3 million
years ago.
This very recent discovery in the central African state of Chad, in
the southern Sahara desert (Brunet et al. 2002, Wood 2002), is
poised to upset the human family tree. The fossil skull
that was found, nicknamed Toumaļ, is as old as any hominid fossil found to date, yet
its features appear much more human-like than those of other contenders
for the title of human ancestor.
It was discovered by Ahounta Djimdoumalbaye in 2001. Based on faunal
studies, it is estimated to be between 6 and 7 million years old, and more
likely in the older part of that range. This is a mostly complete cranium
with a small brain (between 320 and 380 cc) comparable in size to that of
a chimpanzee.
No bones below the skull have yet been discovered, so it is not known
whether Toumaļ was bipedal or not. Brunet concurs that it was likely to be a habitual biped because it shares
characteristics with other hominids known to be bipedal. Other scientists
have pointed out the foramen magnum (the hole through which the spinal
cord exits the skull) of Toumaļ is positioned towards the back of the
skull as in apes, indicating that the skull was held forward and not
balanced on top of an erect body.
Brunet's camp considers Toumaļ to be a hominid, that is, on the human side of
the chimpanzee-human divide and therefore more closely related to us than to
chimpanzees. This is not at all certain. Some scientists think it probable;
others have suggested that it may come from before the point at which
hominids separated from chimpanzees, while Brigitte Senut (one of the
discoverers of Orrorin tugenensis) has suggested that it may be an early gorilla.
It
seems to be impossible to know how Toumaļ is related to hominids until other fossils
can be found from the same time period. Whatever it is, all scientists
agree with its finders that Toumaļ is of major significance, and
subsequent discoveries tend to support the view that Toumaļ is a hominid.
6.2 million
Orrorin tugenensis appeared between 6.2 to 6.0 million years
ago.
This species was named in July 2001 from fossils discovered in western
Kenya (Senut et al. 2001). The fossils include a left femur, pieces of jaw
with teeth, isolated upper and lower teeth, arm bones, and a finger bone.
Preliminary analyses suggests the hominid was an agile climber and that it
walked on two legs when on the ground.
The tentative date of six million years, taken from the age of the
deposits in which the fragments were discovered, indicate a date very
close to the common ancestor of humans and chimpanzees, although this date
may now need to be pushed back. The limb bones are about 1.5 times larger
than those of Lucy, and suggest that it was about the size of a female
chimpanzee.
As no complete skull can yet be formed from the finds, an artist's
impression of Orrorin tugenensis is not yet possible.
Orrorin tugenensis
5.8 million
Ardipithecus ramidus proved to be the second longest surviving
hominid species (known to date), flourishing between 5.8 to 4.4 million years ago
as Middle Miocene became Late Miocene.
While Ardipithecus ramidus is not the sought-after "Missing
Link" - the so-far undiscovered creature that lived at the cusp of the
evolutionary division between man and chimpanzee - Yohannes Haile-Selassie,
a doctoral candidate at the University of California at Berkeley, said the
hominid certainly is very close to the branching point.
Seventeen Ardipithecus ramidus fossils had been located by the end of 1993 from a
cluster of localities West of the Awash River, within the Afar Depression
in Aramis, Ethiopia. The physical attributes of this hominid show a range of primitive
traits, which are most likely character retentions from the last
hominid/chimpanzee ancestor. At the same time, some hominid innovations
are apparent.
The currently known traits of Ardipithecus ramidus
can in general be placed within two categories: ape-like traits and
Australopithecine-like traits. The creature's teeth share more
characteristics with all later-discovered hominids than with the teeth of all
fossil and modern apes. The relatively large back teeth and narrow front
teeth indicate that Ardipithecus ramidus ate less fruit and more
soft leaves and fibrous food than his chimpanzee contemporaries, who were
specialised frugivores. Vertebral differences also suggest that
if ramidus was not bipedal in the modern sense, it at least had key
adaptations toward a similar end, almost certainly walking on two legs
when on the ground.
Haile-Selassie believes that ramidus was about the size of a
modern-day chimpanzee and about 20 percent larger than the 'Lucy'
specimen. Because neither the skull nor intact limb bones of ramidus
have been found an artist's impression of the creature is impossible at
this time.
At the time of it's existence, ramidus lived in a forested,
flood-plain environment - a far cry from Ethiopia's present day
environment of harsh desert surroundings. The area where the hominid
dwelled was as much as 1,500 feet higher in elevation than today and it
was much cooler and wetter. But the hominid lived at a time when Africa
was in the throes of continental change. The area was peppered with active
volcanoes and intense earthquakes related to the formation of the rift
valley (caused by the continent's collision with
Europe and Asia, and its continued northerly movement).
The Awash region then was showered with pulses of thick, hot volcanic
ash from nearby volcanoes. "It's hard to imagine that life would go on
under such hostile environmental conditions," Giday WoldeGabriel, a
geologist with Los Alamos National Laboratory, said. "Ardipithecus and the
other animals inhabiting the region were real survivors." The researchers
found that numerous animals lived during the time of ramidus. The
research team found more than 1,900 fossil specimens comprising the
remains of more than 60 identified mammal species. The fossils included
primitive elephants, horses, rhinos, rats and monkeys. Researchers found
the remains of more than 20 primitive elephants together at one site.
During the Pliocene era (5 million to 1.8 million years ago), hominids were walking upright on a permanent basis, allowing them to fully
leave behind their former arboreal habitats and survive on the ground in
an increasingly treeless environment in Africa. This also freed their
hands for important new tasks, such as food-gathering.
4.2 million
Australopithecus anamensis appeared in Pliocene Africa,
surviving until
3.8 million years ago. It was probably descended from
Ardipithecus
ramidus.
First discovered in the Kanapoi region of East Lake Turkana, Kenya, in 1965,
anamensis comes from 'anam', meaning lake in the local Turkana
language. This species was named in August 1995 (Leakey et al. 1995). The
material consists of nine fossils, mostly found in 1994, from Kanapoi, and
twelve fossils, mostly teeth found in 1988, from Allia Bay in Kenya
(Leakey et al. 1995).
A general similarity to other Australopithecus species seems to exist
in anamensis. It had a mixture of primitive features
in the skull, and advanced features in the body. The teeth and jaws are
very similar to those of older fossil apes, as is its ape-like crania,
although this clearly marks it out as a bipedal hominid.
A partial tibia (the larger of the two lower leg bones) is strong
evidence of bipedal behaviour, and a lower humerus (the upper arm bone) is
extremely humanlike. Note that although the skull and skeletal bones are
thought to be from the same species, this is not confirmed.
Curiously, the tibia and humerus of anamensis may be more
similar to those from members of the genus Homo than they are to
Australopithecus afarensis. This has not been decisively shown, but,
if true, would bring up a very interesting possibility. It may be
that Homo sapiens is more closely related to this 4 million
year-old hominid than to the widely successful later hominid -
Australopithecus afarensis. For the moment this is pure speculation,
but it exists as a possibility.