Early Human Phylogeny

The phylogenetic tree below shows one reconstruction of the relationships among
early human species, as we best know them today.

Early Human Species:

Ardipithecus ramidus Homo habilis
Australopithecus anamensis Homo rudolfensis
Australopithecus afarensis Homo ergaster
Australopithecus africanus Homo erectus
Australopithecus garhi Homo heidelbergensis
Paranthropus aethiopicus Homo neanderthalensis
Paranthropus boisei Homo sapiens
Paranthropus robustus
 The "Robust" Forms

When the fossil record was still poorly understood, it was thought that there existed two distinct
branches of early human. These were the "gracile", meaning small, and "robust", meaning large,
forms of the genus Australopithecus. The gracile forms were represented by the species A.
afarensis and A. africanus; the robust forms by the species A. robustus and A. boisei.
You may be saying "Hey, those last two species were called "Paranthropus robustus" and
"Paranthropus boisei" in the phylogenetic tree!" And you're right. That's the debate.
The word "robust" originally referred to the larger body that the members of these two species were
supposed, by paleoanthropologists, to have had. Few postcranial bones existed at the time for
either, and inferences were based on the size of their teeth. Now that there are more fossils from
which to draw our conclusions, it is apparent that the average body size was not significantly
different between any of the species. But the term does accurately reflect the difference in the size
of the cheek teeth (molars and premolars) of these species. Compare the size of the cheek teeth of
the "robust" P. boisei on the left and the "gracile" A. africanus on the right. These photos are to
scale.

These robust forms were "megadont" -- they had huge, broad cheek teeth with thick enamel. The
incisor teeth were by contrast small. The emphasis was on the rear teeth, and they were designed
to support the stresses of heavy chewing. Combined with the morphology of the other parts of the
skull -- large zygomatic arches to allow the passage of large chewing muscles and a large sagittal
crest to provide a large area to anchor these muscles to the skull -- these early humans showed
adaptations to chewing tough, fibrous foods, giving us clues as to their diet. Microscopic studies of
the teeth show wear indicative of a vegetarian diet for both P. robustus and P. boisei. Similar
expansion of chewing structure can be seen in the evolution of many groups of animals, for
example wild pigs.

Now the debate . . .

All of these species had been assigned to the genus Australopithecus. Yet, because of the extreme
heavy chewing adaptations of the "robust" forms, it is thought that they represent a group of species
(or clade) separate from that which led to modern humans, and eventually this clade became
extinct. As such, many researchers believe that they should be placed in their own genus
Paranthropus, separate from the australopiths that gave rise to later humans. This is not universally
accepted by researchers, although we use that designation for our presentation of the human
lineage. This division would place all of the robust forms together, with the early P. aethiopicus
species as the probable ancestor to both later robust species, P. robustus in southern Africa and P.
boisei in eastern Africa.

However, some researchers suggest, based on subtle facial features, that the eastern African forms
P. aethiopicus and P. boisei form a lineage separate from the South African species P. robustus. In
this case, P. robustus would descend from the southern gracile A. africanus, and the genus name
Paranthropus would be invalid. This implies that the evolution of similar heavy chewing complexes
occurred twice independently. (See a graphic representation of the two implied phylogenies.)

There is still much debate as to the exact ancestor/descendent relationship among the robust
species. As such, some species are referred to in the literature by more than one name. As more
fossils are found, our understanding of this branch of the human family tree will hopefully become
clearer.

Alternative Phylogenies
Robust Forms

Here are two alternate phylogenies implied by recent studies. Both are being supported in the
paleoanthropological community by various researchers.
This first phylogeny represents an ancestor/descendent relationship where the robust forms
descend from A. afarensis through P. aethiopicus, forming their own genus.

In this case A. africanus is distinct from all of the "robust" forms, and the genus designation
Paranthropus is a valid genus name because all of its members are evolutionarily related to one
another..

However, an alternative hypothesis supported by some research on the facial features of the South
African fossils, implies that there were two distinct lineages of early humans that were regionally
distinct: a southern African (A. africanus and P. robustus) and an eastern African lineage (A.
afarensis and P. aethiopicus and P. boisei).

For this hypothesis, each lineage evolved similar robust forms in parallel as the environment
demanded that some populations of early humans needed to subsist on tough and fibrous foods.
Thus, the similar chewing morphologies seen in P. robustus and P. boisei are not the result of a
common ancestor, but of adaptations to similar environmental factors. Here, the genus designation
"Paranthropus" would be invalid because it would include members of two totally separate
evolutionary lineages..

Australopithecus afarensis

Inhabiting eastern Africa between

four and three million years ago,
Australopithecus afarensis was a
long-lived species that may have
given rise to the several lineages
of early human that appeared in
both eastern and southern Africa
between two and three million
years ago. For its antiquity, A.
afarensis is one of the better
known species of early human,
with specimens collected from
over 300 individuals. It is a Composite Reconstruction
species that exhibits many cranial
features which are reminiscent of
our ape ancestry, such as a
forward protruding (prognathic)
face, a "U-shaped" palate (with
the cheek teeth parallel in rows to
each other similar to an ape) and
not the parabolic shape of a
modern human, and a small
neurocranium (brain case) that
averages only 430cc in size (not
significantly larger than a modern
chimpanzee).

The specimens recovered have

given representative examples of
almost all of the bones of the A.
afarensis skeleton. From this, it is
clear that there are many
significant difference between A.
afarensis and its ape
predecessors, one of which is
crucial to later human evolution,
bipedality.

The position of A. afarensis in the AL 200-1

phylogeny of early humans is
under debate. Many feel that it is
ancestral to the east African
"robust" early humans, and
possibly to all robust forms.
Additionally, A. afarensis is
proposed as the ancestor to later
Homo. Yet, research now
suggests that A. africanus might
be ancestral to later Homo.

Pictured to the right are several

of the more important specimens
of A. afarensis in the fossil
record, including Tim White and
Bill Kimbel's composite
reconstruction based on several
specimens, the famous Laetoli
footprints, and the AL 129 knee Laetoli Footprint ***
discovered by Donald Johanson,
below.
 AL 129-1

*** The photograph of the Laetoli footprint has been provided to the Smithsonian
Institution by John Reader, and is used here with his consent. Please note that this image
is the copyrighted material of Mr. Reader, and and cannot be used or reproduced without
his consent.

Australopithecus africanus

The Transvaal region of

South Africa was the home
to the species
Australopithecus africanus,
which lived 3.3 to 2.5
million years ago. This
species was the first of the
australopiths to be
described; Raymond Dart
named the genus and
Taung 1 species in 1925 after his
discovery of the famous
Taung child.

Many features of the

cranium of A. africanus are
more evolved than that of
earlier A. afarensis. These
features include a more

Makapansgat Composite Skull
STS 5
globular cranium and
slightly higher ratio of brain
size to body size. Also the
teeth and face appear less
primitive. For years
researchers considered the
evolution of early humans
to pass from A. afarensis
to A. africanus and lead to
early Homo.
However, some
researchers now believe
that facial features link A.
africanus to the "robust"
early human species of
southern Africa,
Paranthropus robustus.
Known as anterior pillars,
which are located on either
side of the nose, these
features are found in A.
africanus and P. robustus,
and not in the eastern
African species. This
implies that the designation
of the genus Paranthropus
may be incorrect.
As if to confuse the issue
even more, recent
comparative studies of the
postcranial fossils of A.
afarensis and A. africanus
have placed a whole new
unknown into the question.
Evidently, the proportion of
arm to leg lengths was
more ape-like in A.
africanus than in A.
afarensis. This confuses
the phylogeny of early
humans because of the
discovery of the OH 62
fossil, and the post-cranial
paradox it has posed. As a
result, some researchers
are once again pointing to
A. africanus as a possible
ancestor of early Homo.
In all, A. africanus is an
 enigma to
paleoanthropology.
Researchers are still
unsure about where A.
africanus came from and
which species, if any, it led
to. It can safely be said
that to figure out A.
africanus would lead to a
great clarification of our
early evolutionary history.

To the left are four

specimens. The first is the
type specimen of the
species, the Taung Child.
In the middle is a
composite reconstruction
of an adult male A.
africanus composed of
three individuals from the
STS 71 Makapansgat Limeworks in
South Africa. Next is the
famous "Mrs. Ples" of
Sterkfontein. At the bottom
is the STS 71 partial
cranium.

Homo habilis
 OH 7

Until 1964,
Australopithecus
remains had been
found in Africa, but
remains of the
oldest
representative of
the genus Homo
had been
recognized only in
Asia. In that year, KNM ER 1813
however, Louis
Leakey, Phillip
Tobias, and John
Napier announced
the new species
Homo habilis, or
"handy man". They
had to redefine the
genus to
accommodate this
oldest form.

The type specimen

was a mandible,
with associated KNM ER 1805
postcranial bones,
and a fragmentary
cranial vault;
Olduvai Hominid 7
(OH 7). They based
their placement of
OH 7 in Homo
primarily on brain
expansion. Until
then, an arbitrary
lower limit had
been set between
700cc and 800cc
as the cutoff for the
genus Homo. With
an estimated
cranial capacity of
680cc, Leakey and
his colleagues OH 24
chose to lower this
number to 600cc.
While calling
attention to
anatomical
differences
between OH 7 and
Australopithecus,
they chose a
behavior- the ability
to make stone
tools-to help place
OH 7 in Homo. This
point relied on
stone tools found in
the same geologic
horizon as the
fossils.

The OH 7 mandible
is shown at the top
right. In the 1960s,
many researchers
argued that Homo
habilis was not a
valid species, and
that the fossils
attributed to H.
habilis were really
members of other
species. But with
the discovery of
KNM ER 1470,
acceptance of
Homo habilis
became universal.
In hindsight, this
seems strange
since ER 1470 is
now considered to
belong to a species
distinct from H.
habilis. There is
much debate as to
the number of
species that existed
in Homo 2 million
years ago, and
KNM ER 1470 is
now assigned to
the species Homo
rudolfensis. The
name Homo habilis
is reserved
primarily for the
Olduvai material
and several other
specimens. The OH
62 partial skeleton
of a female H.
habilis provides
another interesting
twist in the debate
about early
members of the
genus Homo.

Homo habilis was

originally thought to
be the ancestor to
all later Homo. In a
neat, linear
progression, later
species emerged
resulting in what we
call modern
humans. This is
now known not to
be the case.

Also shown are the

KNM ER 1813
skull, OH 24, and
part of the
fragmentary KNM
ER 1805 cranium.

Homo rudolfensis
 The species Homo
rudolfensis was
originally proposed
in 1986 by V. P.
Alexeev for the
specimen to the
left, KNM ER 1470.
Originally thought
to be a member of
the species Homo
habilis, much
debate surrounded
the fossil and its
species
assignment. It was
thought that 2
million years ago
there existed a
single species in
the genus Homo,
and this species
evolved in a linear
fashion into modern
humans.

But the differences

in this skull, when
compared to other
habilines, are too
pronounced,
leading to the
formulation of the
KNM ER 1470 species Homo
rudolfensis,
contemporary with
Homo habilis.

It is not yet certain

if H. rudolfensis
was ancestral to
the later species in
Homo, or if H.
habilis was.

Homo ergaster
By 1.9 million years
ago, another
lineage of the genus
Homo emerged in
Africa. This species
was Homo ergaster.
Traditionally,
scientists have
referred to this
species as Homo
erectus and linked
this species name KNM ER 3733
with a proliferation
of populations
across Africa,
Europe, and Asia.
Yet, since the first
discoveries of
Homo erectus, it
had been noted that
there were
differences between
the early
populations of
"Homo erectus" in KNM ER 992
Africa, and the later
populations of
Europe, Africa and
Asia. Many
researchers now
separate the two
into distinct species
Homo ergaster for
early African "Homo
erectus", and Homo
erectus for later
populations mainly
in Asia. Since
modern humans
share the same
differences as H.
KNM ER 3883
ergaster with the
Asian H. erectus,
scientist consider H.
ergaster as the
probable ancestor
of later Homo
populations.

H. ergaster had a
rounded cranium
and a prominent
browridge. Its teeth
were much reduced
in size, especially
when compared to
Australopithecus.
Several features KNM WT 15000
that distinguish H.
ergaster from H.
erectus are thinner
bones of the skull
and the lack of an
obvious sulcus, or
depression, just
behind the
browridge.

By 1.6 million years

ago, an advance in
stone tool
technology is
identified with H.
ergaster. Known as
the Achulean stone
tool industry, it
consisted of large
cutting tools,
primarily hand axes
and cleavers.
Originally thought to
be responsible for
the spread of early
humans beyond
Africa, it is now
known that the
migration out of
Africa predates this
tool industry.

At the top-left is the

amazingly well
preserved KNM ER
3733 cranium.
Second from the top
is the type
specimen of the
Homo ergaster
species, the KNM
ER 992 mandible.
At the bottom is the
famous Turkana
Boy KNM WT
15000, a nearly
complete skeleton
dating back to 1.6
million years.

Homo erectus

To understand what
we mean today by
"Homo erectus",
some history of
paleoanthropological
thought is needed.
The first early
human fossil found
outside of Europe
was the Trinil 2 fossil
skullcap from the
Solo River in Java,
pictured to the right. Trinil 2
The fossil was
placed in the species
Pithecanthropus
erectus by its
discoverer Eugene
Dubois. Almost 40
individuals have
been recovered from
Java to this day,
roughly equivalent to
the number of fossils
found at the caves of
Choukoutien in
China. The
Choukoutien fossils
found were originally
 NG 6
assigned the species
name Sinanthropus
pekinensis. It was
not until the 1950's
that Ernst Mayr
proposed that all of
the specimens from
these two roughly
contemporaneous
locales, along with
others localities from
Europe and Africa,
represented a single
species, Homo
erectus. Since the
1950's, however, the
early African
populations of what
Mayr termed Homo Weidenreich Reconstruction
erectus have once
again been split into
a separate species
Homo ergaster.

Homo erectus
exhibits many
features particular to
the species,
including a long skull
shaped with thick
cranial walls. The
back of the skull is
marked with a
protruberance
known as a
transverse torus.
Over the eyes is a
large and prominent Sangiran 2
browridge, or
supraorbital torus,
which joins the rest
of the frontal bone at
a depression called
the sulcus. Cranial
capacities of Homo
erectus average
around 1000cc,
which is far greater
than earlier
australopiths and
even early Homo.
The dentition of
Homo erectus is
nearly identical to
modern humans,
although the cheek
teeth do remain
larger, and the
mandible is
generally more
robust.

The species Homo

erectus is thought to
have diverged from
Homo ergaster
populations roughly
1.6 million years
ago, and then
spread into Asia. It
was believed that
Homo erectus
disappeared as
other populations of
archaic Homo
evolved roughly
400,000 years ago.
Evidently, this is not
the case. Recent
studies into the
complicated
stratigraphy of the
Java Homo erectus
sites have revealed
some surprising
information.
Researchers have
dated the deposits
thought to contain
the fossils of H.
erectus near the
Solo River in Java to
only 50,000 years
ago. This would
mean that at least
one population of
Homo erectus in
Java was a
contemporary of
modern humans
(Homo sapiens).

Homo heidelbergensis
Homo heidelbergensis is the
species name now given to a
range of specimens from
about 800,000 years ago to
the appearance of
anatomically modern Homo
sapiens (the species to
which we belong). The
species name was originally
proposed for the fossil
mandible discovered at
Mauer, a town near
Heidelberg, Germany. It is a
nearly complete early human
mandible that is very robustly
built, but lacks a chin.
Additional finds of early Mauer 1
humans with morphological
attributes of both modern
humans and Homo erectus
have shown that the
transition from early and
middle Pleistocene forms
and the morphology of
modern humankind was not
a neat transition that could
be easily explained.

For many years, scientists

placed any problematic
specimens displaying
mixtures of "erectus-like" and
"modern" traits into a
confusing category:
"Archaic" Homo sapiens Kabwe Cranium
(basically meaning any
Homo sapiens that didn't
look quite modern). Recently,
it has been proposed to
separate these individuals
into a distinct species. For
this purpose, the Mauer
mandible, and the species
name Homo heidelbergensis
has seniority.

Homo sapiens
The species to which you and all
other living human beings on this
planet belong is Homo sapiens.
Anatomically, modern humans can
generally be characterized by the
lighter build of their skeletons
compared to earlier humans.
Modern humans also have very
large brains, which vary in size
from population to population and
between males and females, but
the average is around 1300 cc.
Housing this enlarged brain has
involved the reorganization of the
skull into what is thought of as the
"modern" appearance -- a high
vaulted cranium with a flat and
near vertical forehead. The
supraorbital torus is lost in most
modern humans, and ridging
above the orbits in general is very
reduced. The widest part of the
skull is high on the skull, as
opposed to earlier Homo erectus
and H. ergaster. The back of the Cro-Magnon 1
skull lacks the transverse torus of
H. erectus and the occipital bun of
H. neanderthalensis (Compare the
crania of H. neanderthalensis and
H. sapiens).

The origin of modern Homo

sapiens is not yet resolved. Two
extreme scenarios have been
proposed. According to the first,
the distribution of anatomical traits
in modern human populations in
different regions was inherited
from local populations of Homo
erectus and intermediate "archaic"
forms. This "Multiregional
Hypothesis" states that all modern
humans evolved in parallel from
earlier populations in Africa,
Europe and Asia, with some
genetic intermixing among these
regions. Support for this comes
from the similarity of certain minor
anatomical structures in modern
human populations and preceding Wadjak 1
populations of Homo erectus in the
same regions.

A different model proposes that a

small, relatively isolated population
of early humans evolved into
modern Homo sapiens, and that
this population succeeded in
spreading across Africa, Europe,
and Asia -- displacing and
eventually replacing all other early
human populations as they
spread. In this scenario the
variation among modern
populations is a recent
phenomenon. Part of the evidence
to support this theory comes from
molecular biology, especially
studies of the diversity and
mutation rate of nuclear DNA and
mitochondrial DNA in living human
cells.From these studies an
approximate time of divergence
from the common ancestor of all
modern human populations can be
calculated. This research has
typically yielded dates around
200,000 years ago, too young for
the "Multiregional Hypothesis."
Molecular methods have also
tended to point to an African origin
for all modern humans, implying Skhul V
that the ancestral population of all
living people migrated from Africa
to other parts of the world -- thus
the name of this interpretation: the
"Out of Africa Hypothesis."

Whichever model (if either) is

correct, the oldest fossil evidence
for anatomically modern humans is
about 130,000 years old in Africa,
and there is evidence for modern
humans in the Near East
sometime before 90,000 years
ago.

Homo neanderthalensis
The discovery in 1856 of a
skullcap and partial skeleton
in a cave in the Neander
valley near Dusseldorf,
Germany, signaled the first
recognized fossil human
form. While it was later
realized that several
Neanderthal sites had La Ferrassie 1
previously been discovered,
their remains were not
recognized as those of an
archaic form of human until
the discovery of
"Neanderthal Man." In 1864
a new species was
recognized: Homo
neanderthalensis.

Neanderthals inhabited
Europe and western Asia
during the latter part of the
Pleistocene. The climate in
these regions was much
colder than it is today, and
several glaciations, or Ice
Ages, are known to have
occurred during the time of La Chappelle-aux-Saints
Neanderthal occupation.
Neanderthal localities are
known today from Spain to
Uzbekistan (near
Afghanistan). Several
important sites in the vicinity
of Qafzeh Cave, Israel,
suggest that Neanderthals
arrived in the region after
modern Homo sapiens. This
would indicate that the
population of modern
humans in this area was not
descended form
Neanderthals, and that there
was some period of
coexistence, or an
alternating series of
migrations into this region by
the two species.
Neanderthals are known
from Europe and western
Asia from about 200,000
years to about 30,000 years
ago, when they disappeared
from the fossil record and
were replaced in Europe by
anatomically modern forms.

The original interpretation of

Neanderthal anatomy was
one of a primitive early
human based on a flawed
reconstruction of the nearly
complete skeleton of an
elderly Neanderthal male
found at La Chapelle-aux-
Saints, France (second
photograph from the top).
However, Neanderthals and Shanidar 1
modern humans (Homo
sapiens) are very similar
anatomically -- so similar, in
fact, that in 1964, it was
proposed that Neanderthals
are not even a separate
species from modern
humans, but that the two
forms represent two
subspecies: Homo sapiens
neanderthalensis and Homo
sapiens sapiens. This
classification was popular
through the 1970's and 80's,
although many authors today
have returned to the previous
two-species hypothesis.
Either way, Neanderthals
represent a very close Le Moustier
evolutionary relative of
modern humans.

Several features of the

skeleton unique to
Neanderthals appear to be
related to cold climate
adaptations. These features
include limb-bone
proportions and muscle
attachments indicative of a
 broad, slightly short, and
strong body; a large,
rounded nasal opening; and
a suite of anatomical traits of
the skull (compare the crania
of H. neanderthalensis and
H. sapiens).

In all, the fossil record for

Neanderthals is significantly
better than for earlier human
species. One reason for this
is that Neanderthal fossils
are relatively young
compared to other early
human species, and fossils
decay over time. But another
very important factor is the
purposeful burial of their
dead. Many Neanderthal
sites include the remains of
individuals who were
deliberately placed in graves
dug into the earth. Some of
these burials show evidence
that may indicate that these
graves were adorned with
offerings (such as flowers).
This cultural advance, which
represents an awareness
and recognition of life and
death, may have first been
practiced by the
Neanderthals.

Paranthropus robustus
From around 2 million to 1.2
million years ago, southern
Africa was inhabited by a robust
species of early human. From
the original finds in 1938 and
later finds in 1948, it was evident
that they represented very
different early human
morphologies than were seen in
the known Australopithecus
specimens.
SK 48
Adaptations of the cranium were
associated with a "heavy-
chewing comnplex." This
complex is thought to have
made it possible for these early
humans to eat large amounts of
tough, fibrous foods. The
zygomatic arches - cheek
bones - are large and positioned
forward on face, creating a
characteristic dish-shaped face.
They flare very wide, creating a
large space between the arch
and the skull, and opening
known as the temporal fossa.
Two sets of muscles are TM 1517
associated with the grinding
action necessary to process
these types of food, the
masseter complex, which
attaches on the bottom of the
zygomatic arch, and the
temporalis muscles which pass
under the arch and attach to the
top of the skull. The foreward
migration of the cheekbones
creates more space for
temporalis muscles to pass
under the zygomatic arch, and
the increased size of the arches
provides more room to
accommodate a larger masseter
muscle. Another feature of the
robust skull is the presence, at
least in males, of a prominent
sagittal crest, a bony ridge that SK 46
runs along the length of the top
of the skull. This bony ridge
provides an anchoring point for
the large temporalis muscles.

These unique adaptations lead

Robert Broom to place the
robust early humans from
southern Africa into their own
genus Paranthropus. Several
species names have been
proposed, including P. robustus
or P. crassidens. In the 1960's
paleoanthropologists began to
note similarities between all of
the early human species before
the appearance of Homo. As
such, many researchers began
to place all early human species
into a single genus
(Australopithecus) and described
each species as either a
"gracile" or "robust" Australopith
(see the robust debate). The
robust speciemns from southern
Africa were then placed in the
species "Australopithecus
robustus." One resercher,
Milford Wolpoff, even went so far
as to propose that all robust
forms were really just the males
and the gracile forms the
females of a single early human
species (see description of
Paranthropus boisei for the
resolution of this debate).

In recent years, many

researchers have sought to
emphasize the uniqueness of
the heavy-chewing adaptations
seen in at least three separate
species of ealry human. Many
favor the separation of these
species into a robust genus of
early human, for which the name
Paranthropus was the first used,
and therfore has seniority over
all other names. Itis this
classification that we favor here,
but it should be noted that there
is, as yet, no consensus among
paleoanthropologists on this
issue.
Paranthropus boisei

In the time period dating from 2.3 and 1.2

million years ago eastern Africa was
populated by the early human species
Paranthropus boisei. This species
represents the most extreme version of the
"robust" early humans in eastern Africa.
Paranthropus boisei had a skull highly
specialized for heavy chewing. They
flourished in the drier savanna areas that
existed in eastern Africa at the time. But by
around 1.2 million years ago, Paranthropus
boisei disappears from the fossil record.
With a major change in Earth's climate,
which involved larger, irregular fluctuation,
adaptability may have been critical to
survival as old resources dwindled or
disappeared. The highly specialized
Paranthropus boisei might not have been
able to effect such adaptability in the face of OH 5
change.

The original specimen of this species, the

"type specimen," is the OH 5 skull found in
1959 by Mary Leakey at Olduvai Gorge in
Tanzania. The photograph on the top-right is
a reconstruction of OH 5 with the mandible
of another individual scaled to fit (the
original find did not include the lower jaw).

Below OH 5 are two very important P. boisei

fossils; KNM ER 406 and KNM ER 732,
thought to be a male and female
respectively.

There was some earlier debate in the

paleoanthropological community as to the
number of early human species in southern
Africa between 3 and 1 million years ago.
Conventional wisdom had it that two species KNM ER 406
existed, Australopithecus africanus and
Paranthropus robustus. In contradiction to
this view, Milford Wolpoff, of the University
of Michigan, advocated the"single species
hypothesis". It claimed that the differences
between the southern forms were caused by
age differences and sexual dimorphism of
the specimens. Many researchers had
problems with this hypothesis. For example,
why in southern Africa were the supposed
males dying at a different place than the
supposed females? And why were they
dated to almost a half a million years later?
It was clear that a larger fossil record would
be needed to prove or disprove this
hypothesis.

Interestingly, the answer to the question of

the southern African early humans would KNM ER 732
come from hundreds of kilometers away in
East Africa. The discovery of two fossils,
KNM ER 406 and KNM ER 732, at Koobi
Fora in eastern Africa would provide the
necessary expansion of the record needed
to disprove the "single species hypothesis".
Upon discovery its in 1969, ER 406 showed
enough similar morphology to be assigned
to the same species as OH 5; with the
addition of ER 732, comparisons could be
drawn between the two that could shed light
on the nature of dimorphism in early
humans. As these two specimens were
examined, researchers found that the early
humans of this period followed what is
called the great ape model of sexual
dimorphism. Male crania were larger than
females, and more heavily constructed.
While differences existed between the two
skulls, these differences were exactly what Peninj Mandible
would be expected between the sexes in
other great apes.

The two southern African forms, however,

did not fit this model of the distinction
between the sexes. The differences were
too great to be the result of sexual
dimorphism. This observation favored the
idea of two distinct species in southern
Africa.

The final blow to the "single species

hypothesis" was the 1975 discovery of the
cranium KNM ER 3733, assigned now to
Homo ergaster, in the same layer as ER
406, the "robust" form Paranthropus boisei.
Scientists finally knew for sure that more
than one species of early human coexisted
in the same geographical area. The old
single line of progressive evolution was,
once and for all, split into branches. And the
human family tree has never looked the
same since.

The "Peninj mandible" is a nearly complete

mandible of Paranthropus boisei. It provided
researchers with their first understanding of
the complete adult dentition and the
structure of the lower jaw of this species.

KNM ER 23000 KNM WT 17400

Paranthropus aethiopicus
 By 2.7 million years
ago, a new lineage
of early humans had
evolved in East
Africa. Paranthropus
aethiopicus was
originally proposed
in 1967 by a team of
French
paleontologists to
describe a partial
mandible (Omo 18)
that was thought to
differ enough from
the mandibles of the
early human species
known at that time.

This naming of a
new species was
KNM WT 17000 generally dismissed;
many
paleoanthropologists
thought it premature
to name a new
species on the basis
of a single
incomplete
mandible. In 1985,
when Alan Walker
and Richard Leakey
discovered the
famous "black skull"
west of Lake
Turkana in Kenya,
the classification
reemerged. With its
mixture of derived
and primitive traits,
KNM WT 17000
validated, in the
eyes of many
researchers, the
recognition of a new
"robust" species
dating to at least 2.5
million years ago in
eastern Africa. The
issue was debated,
but today there is
general consensus
that there was a
distinct robust
species,
Paranthropus
aethiopicus, living in
eastern Africa 2.5
million years ago.

The exact phylogeny

of P. aethiopicus is
still not fully
understood. It is
thought to have
descended from the
earlier A. afarensis,
and to be ancestral
to P. boisei.
However, go to the
robust forms page to
learn more about the
current debate
concerning this
interesting branch of
the human tree.