Written by
C. David Kreger
Introduction
Homo habilis is a well-known, but poorly defined species.
The specimen that led to the naming of this species (OH 7) was
discovered in 1960, by the Leakey team in Olduvai Gorge, Tanzania.
This specimen and its designation was the subject of much controversies
up through the 1970s. The material was found in the same region
where A. boisei had previously been found, and many researchers
of the time did not fully accept that the material was sufficiently
different from that material (or maybe A. africanus)
to denote a new species. Louis Leakey was convinced that this
was the Olduvai toolmaker he had spent his life looking for,
and placed this as a direct human ancestor, with H. erectus
a dead-end side-branch.
The
specimen was subjected to intense study by the multidisciplinary
team of Louis Leakey, John Napier, and Phillip Tobias. They
placed the material as different from penecontemporary australopithecines
due to the teeth, which fell outside the known range of A.
africanus, with very large incisors. Also, the large brain
size and shape of the hand suggested a closer affinity with
Homo. In January 1964, the team announced the new species
Homo habilis. The name was suggested by Raymond Dart,
and means "handy man," in reference to this hominids
supposed tool making prowess.
Leakey
believed that habilis was a direct human ancestor,
with erectus out of the picture. While H. habilis
is a generally accepted species, they opinion that it was
a direct human ancestor seems to be in question. There are
now at least two species of early Homo (whether habilis
and rudolfensis or an undescribed species) living prior
to 2.0 myr. In addition, H. erectus (which is almost
universally accepted as a direct human ancestor) continues
to be pushed further back into the paleontological record,
making it possible that it is the first Homo ancestor
of modern humans.
Other
problems include that some people see KNM-ER
1813 as a near perfect erectus, except for its
small brain and size. It could be an erectus that was
at the small scale of a wide variation of traits, or it may
belong to ergaster, which some believe to be the ancestor
of erectus. The questions are far from solved, and
new specimens are needed. Homo habilis may be a direct
human ancestor, a dead-end side-branch that leads nowhere,
an invalid species whose designated examples belong in other
species, or Wolpoff may be right, and all these species are
basically part of one highly variable widespread species.
Diagnostic Features
It is particularity hard to list the features of Homo habilis,
because the specimens attributed to habilis (and the
reasons the material was placed there) vary widely. The species
is a mishmash of traits and specimens, whose composition depends
upon what researcher one asks. The simplest way to describe
the general features is to describe specimens that are generally
considered habilis by most people, and list their relevant
traits.
OH
7 is the type specimen of habilis, and the first material
attributed to the species. The specimen consists of a nearly
complete left parietal, a fragmented right parietal, most
of the mandibular body (including thirteen teeth), an upper
molar, and twenty-one finger, hand, and wrist bones. The remains
belonged to a 12 or 13 year old male. The brain size attributed
to this specimen varies, ranging from 590-710 cc. P. Tobias
and G. von Koenigswald used three traits to set habilis
apart, as a transitional species between A. africanus
and H. erectus:
- Expanded
cranial capacity (relative to africanus).
- Reduced
postcanine tooth size.
- The
presence of a precision grip (determined from the hand bones
present in OH 7), which provides the anatomical basis for
tool-making.
General
features of the specimen seems to support these three traits
(whether or not it is transitional from africanus to
erectus):
- Larger
cranial capacity (though very problematic). Tobias gives
an estimate of 647 cc, Holloway gives an estimate of 710
cc, and Wolpoff has estimated it at 590 cc.
- Molar
megadontia is gone, with molars longer than they are wide.
- The
P3 is smaller and more asymmetric.
- The
P4 is much more similar to the P3.
- Metacarpal
1 and trapezium is much less interlocked, which allowed
more movement.
- The
distal phalanges have apical tuffs.
OH
8 - a fairly complete foot - was found nearby OH 7, and was
initially determined to be from another individual. This was
due to the fact that OH 7 was known to be from an adolescent
around age 12, and the foot seemed to be of a more advanced
age, due to the presence of arthritis in the specimen. However,
the partially gnawed remains have arthritis due to a sustained
injury, and the actual age runs close to OH 7, making it likely
they are from the same individual (many researchers consider
this part of OH 7 now, rather than OH 8). The remains show
clear signs that this was an obligate biped, including:
- Presence
of digital shortening.
- Enlargement
of the hallux, as well as being fully abducted.
- Alignment
of digits 2-5.
- Thickened
metatarsal shaft with a humanlike cross-sectional shape.
- A
fully developed double arch to the lower surface.
- Mechanically
set up for efficient weight transmission at the ankle.
While
it shows definite obligate bipedalism, the specimen also has
a marked tubercule for the tibialis posterior muscle, an invertor
of the foot that could be useful for climbing. So it is possible
that while this individual was an obligate biped, it still
spent some time in the trees (which goes well with paleoecological
evidence that suggests that various hominid species spent
most of their time in marginal woodland environments). From
the talus, H. McHenry calculated an estimated weight of 31.7
kg. Using the various estimated brain size, one gets brain/body
weight ratios of:
Wolpoff:
590 cc brain = 1.86%
Tobias: 647 cc brain = 2.04%
Holloway: 710 cc brain = 2.24%
Even
using the smaller brain estimate, this is one of the largest
relative brain size for any male hominid up to the time period
this individual lived (1.75 myr). When compared to primate
allometry, the OH 7 brain size is at the top of the allometric
expectations within non-human primates. This is a large brained
specimen relative to its body size.
Another
relatively complete habiline specimen is OH 13 - "Cinderella".
This is a poorly preserved and fragmentary specimen of a 15-16
year old female habilis, dating to a little younger
than 1.66 myr. This makes it one of the most (if not the most)
recent habilis specimens known. The material consists of the
mandible and the maxilla, several teeth, pieces of the cranial
vault, and some postcranial elements, including a small piece
of proximal ulna. This specimen (along with OH 16) were the
object of much inaccurate brain size estimations, which originally
lead to the two being classified as H. erectus. More
recent estimates put the brain size at around 500 cc, and
along with an estimated body size near that of AL 288-1, gives
this specimen a relative brain/body weight ratio similar to
OH 7.
The
case of OH 16 is a tragic one. The specimen was discovered
nearly complete near the end of a field day, so the position
of the find was marked and roped off. The next morning the
researchers were horrified to discover that a herd of cattle
had charged through the area, and completely crushed it. Some
of the specimen's features include:
- Very
large teeth (close to australopithecus in size).
- An
uncertain brain size, but probably larger than OH 7.
- The
individual was age 15-16 when they died.
- The
individual had very bad caries on one side of its jaws (very
unusual in ancient specimens), which lead to differential
chewing on the other side, causing it to develop a huge
temporalis muscle on that side.
- Cranial
bone markedly thinner that erectus.
- Dramatic
differences in the supraorbital torus and the nuchal torus
that distinguish it from erectus.
These
features (the last two shared with OH 13) seem to indicate
that the specimen is a habiline, and not an erectus
specimen, as was attributed by J. Robinson.
OH
24 ("Twiggy") is the most australopithecine-like
of specimens attributed to H. habilis, and may be more
highly correlated with A. africanus. The specimen was
found completely fractured, and cemented together in a coating
of limestone. R. Clarke was the researcher who went through
the long and painful process of reconstruction, but over 100
small fragments could not be placed in the reconstruction.
Hence, the specimen is extremely distorted, making an accurate
estimate of its brain size very difficult, though Holloway
has given an estimate of 590 cc (many researchers believe
that number is too high). Several features caused this specimen
to be placed in habilis, including:
- Increased
cranial capacity over australopithecines (though some doubt
this estimate).
- Less
postorbital constriction.
- Elongated
molars.
- Absence
of postcanine megadontia.
- Large
front teeth relative to the postcanines.
- A
broad and short cranial base.
- Anteriorly
positioned foramen magnum.
- Less
convex and bulging zygomatics, and more vertically oriented.
- A
distinct maxillary notch.
While
these features seem to support the notion that it is not an
australopithecine, several other features do not support the
habilis distinction. For example:
- Lacks
a salient anterior nasal spine.
- Lacks
broad nasal bones.
- Lacks
nasal bone peaking caused by the internasal angle.
- Lacks
the projection of the middle and top of the nose away from
the face, shown by expanded and outward projecting maxillary
bones to its side (i.e., maxillary pillar eversion).
While
this specimen does not seem to be an australopithecine, it
also does not seem to fit perfectly into the classic habilis
mold. Perhaps it fits more closely with rudolfensis,
or an undefined penecontemporary species. Another option may
be that the distorted reconstruction is blurring a clear species
designation. For now, a clear designation is up in the air.
The
last to discuss is OH 62. Publicized widely as "Lucy's
Child" by Johanson, it is a very scrappy collection of
302 bone fragments. Portions of the maxilla (which permitted
identification as habilis), parts of the femur, and
upper limb bones. The entire specimen is problematic, and
raises many questions as to sexual variation and behavior.
The most controversial aspect of the specimen was the Johanson
et al. calculation of a humero-femoral index of 95%. The material
was far too sparse to calculate such an index, but even using
their own estimated range of possible lengths for the incomplete
femur, they should have computed an index quite close to the
A.L. 288-1 value of 83.8%. It is unclear why Johanson et al.
calculated the index in the manner they did, and it is generally
not accepted in any form at all.
Conclusions
Homo habilis is a very complicated species to describe.
No two researchers attribute all the same specimens as habilis,
and few can agree on what traits define habilis, if
it is a valid species at all, and even whether or not it belongs
in the genus Homo or Australopithecus. Hopefully,
future discoveries and future cladistic analyses of the specimens
involved may clear up these issues, or at least better define
what belongs in the species.
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