Research TOC
THE NATURAL HISTORY
AND EVOLUTIONARY FATE
OF HOMO ERECTUS
Andrew Kramer
A
lthough scientists who study human evolution are popularly perceived as constantly being involved in rancorous
debates, these paleoanthropologists would unanimously
agree that we have evolved from more primitive predecessors.
Homo erectus, a hominid species that flourished for over one million
years in Africa and Asia, is generally regarded as humanity’s
immediate ancestor. However, there is still plenty of controversy
surrounding the question of how (and even if) Homo erectus
evolved into Homo sapiens.
In this chapter I present the remarkable story of the discoveries
of and the ideas surrounding the original Homo erectus fossils. This
leads into a discussion of how these early interpretations have
influenced the present-day debates concerning the evolution of
Homo erectus. Finally, I detail my own perspectives and contributions to the resolutions of these questions, and I conclude with
some suggestions for future research.
DUBOIS’S APE-MAN1
In the late 1800s the famous German naturalist Ernst Haeckel
posited that a form of prehuman that bridged the evolutionary gap
between apes and humans existed sometime in the distant past.
Haeckel was even bold enough to name this hypothetical creature
“Pithecanthropus alalus,” literally, “speechless ape-man.” This idea
fired the popular imagination and was the original source of the
term missing link, a phrase coined by an American journalist at the
time. In the early 1880s a young Dutch physician named Eugène
Dubois, an anatomy assistant under Haeckel at the University of
Jena, fell under the thrall of his mentor’s evolutionism. In fact,
Haeckel’s ideas captivated Dubois so completely that he gave up
his promising medical career to devote himself full time to searching for the missing link!
Although Dubois’s chances amounted to what was in all likelihood a million-to-one shot, he defied the odds and, amazingly
enough, actually discovered what he had set out to find. Dubois
decided to search for his fossils in what is today the island country
of Indonesia, in southeast Asia. His decision was based on a combination of theoretical and pragmatic considerations. First, Haeckel
thought that the gibbon was the closest living relative to humans,
not the African apes as Darwin had proposed (correctly, it turned
out). Therefore, because gibbons occupy the forests of mainland
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and island southeast Asia, Dubois logically concluded that the
common ancestor of these apes and humans may have lived in this
same region. The young Dutchman’s decision to direct his energies
to the area’s islands, not to the mainland, was purely practical:
What is today known as the Republic of Indonesia was the Dutch
East Indies then.
Dubois enlisted in the Dutch colonial army as a health officer
and arrived on the island of Sumatra in 1887. His paleontological
research was eventually supported by the colonial government
because much of the geological information he recovered was of
economic value, such as sites that could produce potentially profitable mines. By 1890 Dubois had himself transferred to Java, and
two years later he made the discovery that would rock the anthropological world. In September of 1891, near the village of Trinil
along the banks of the Solo River, his team of convict excavators
uncovered a long and low skull cap with protruding, ape-like brow
ridges. The following August, only forty feet from the original find,
a humanlike femur (thigh bone) was discovered that belonged,
Dubois believed, to the same individual as the skull. Dubois published his interpretations of these finds in 1894 and concluded that
because the skull was intermediate in size between apes and
humans he had truly found the missing link. Honoring Haeckel,
and acknowledging the upright, bipedal gait suggested by the
femur, Dubois named the fossil “Pithecanthropus erectus” (“upright
ape-man”).2
Upon his return to Europe in 1895, Dubois was plunged into
the maelstrom of scientific controversy swirling around his finds.
Although there were those who agreed with his missing link interpretation, including (not surprisingly) Haeckel, there were others,
led by the renowned German pathologist and anthropologist
Rudolf Virchow, who dismissed Dubois’s fossils as nothing more
than an extinct, giant gibbon. The acrimony of these exchanges
may have been more than Dubois could take, for he became less
and less involved in the public debates over the following years.
Legend has it that he ultimately signaled his complete withdrawal
by burying the fossils beneath the floorboards of his dining room
and leaving them there unstudied for the next twenty years!
Although Dubois’s final thoughts concerning the status of
“Pithecanthropus” currently remain a contentious topic, 3 it is
acknowledged by all that his discoveries set the stage for those that
followed during the succeeding decades.
HOMO ERECTUS
5
SPECIES, SPECIES,
AND MORE SPECIES
Beginning in the 1920s and continuing through the 1930s, fossils
found in China demonstrated that Dubois’s find was not unique.
The remains of ancient hominids that were remarkably similar to
“Pithecanthropus” of southeast Asia were recovered in a cave now
known as Zhoukoudian, approximately thirty miles from Beijing.
Davidson Black, an anatomist at the Peking Union Medical School,
placed those specimens into a new genus and species,
“Sinanthropus pekinensis” (“Chinese man of Peking”). Black created
the new name in 1927 on the basis of a single molar tooth, and
although the skulls that were found later in the same cave were
much like “Pithecanthropus,” there was no effort at the time to consolidate the Javan and Chinese fossils into the same genus, much
less the same species (see Figure 1).
This lack of consolidation reflected the “splitting” mentality
that was pervasive throughout paleoanthropology until the 1960s.
Almost every new hominid fossil that was discovered, despite its
resemblance to the Javan and Chinese forms, was given at least a
new species, and often a new genus, name reflecting less the realities of biology and more the egos of the discoverers and namers.
This trend is well illustrated by the treatment of the fossils discovered in Africa during the middle decades of the twentieth century.
The genus names proposed for these specimens, such as
“Telanthropus,” “Atlanthropus,” and “Tchadanthropus,” were biologically meaningless labels that served only to clutter the taxonomic
landscape.4
The lone voice in the wilderness during this “age of splitters”
belonged to the famed German paleoanthropologist Franz
Weidenreich. A refugee from Hitler’s holocaust, Weidenreich published minutely detailed monographs describing and interpreting
the finds from Zhoukoudian. He also reproduced extremely accurate casts and molds of these fossils. The scientific world owes a
debt of everlasting gratitude to Weidenreich for these scholarly
activities because the original Chinese fossils were mysteriously
lost during World War II. In 1940 Weidenreich was the first to suggest that the proliferation of names given to the Javan and Chinese
fossils was hindering the understanding of human evolution:5
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Figure 1
“Pithecanthropus” from Java and “Sinanthropus” from China
Today both are considered to be representatives of Homo erectus.
Note how they share relatively large brain cases that are long and low,
projecting brow ridges, jutting lower faces, and the lack of a chin.
Those terms which are generally used to designate different
human types involve the idea that each one represents a more
or less divergent genus without generic connections. In order
to avoid this incorrect interpretation, the time has come…to
eliminate all those names which may lead to some misunderstanding in this regard.6
Weidenreich observed that “Pithecanthropus” and
“Sinanthropus” were not different enough from modern humans
(Homo sapiens) to be placed into separate genera, and they were not
different enough from one another to be considered separate
HOMO ERECTUS
7
species. Therefore, Weidenreich concluded that both could be
accommodated by the name Homo erectus, recognizing that the
Javan and Peking hominids belonged to the same early species of
humanity, and their differences were no greater than those seen
between modern humans living in southeast Asia and China
today. Weidenreich truly was a visionary. He was decades ahead
of his time in his recognition of the ubiquity of normal, intraspecific (within species) variation among fossil hominids and its
importance in any reconstruction of human evolution.
Weidenreich’s perspective became increasingly popular in
anthropology over the following decades. Particularly persuasive
arguments that Homo erectus was the appropriate taxonomic
assignment for the Chinese and Javan fossil hominids were made
by the American evolutionary biologist Ernst Mayr in 1950, and
by the British paleoanthropologist W. E. LeGros Clark in 1955.7
The 1960s and 1970s signalled the ascendancy of the “lumpers,”
who, unlike the “splitters” of the first half of the century, tried to
bring paleoanthropology into line with the rest of modern biology.
These researchers were much less inclined to name new fossil
hominid genera and species and believed that Homo erectus could
accommodate the variation present not only in the Javan and
Chinese forms but also in similar fossils discovered later in Africa.
Most anthropologists at this time viewed Homo erectus as humanity’s immediate predecessor, a fossil species occupying an intermediate position on the unbroken evolutionary continuum
between modern humans and the earliest African hominids, the
australopithecines.
These ideas held sway throughout the 1970s and 1980s, as
exemplified by the writings of such scholars as F. Clark Howell
and William Howells, and continue to be championed today by
Philip Rightmire.8 However, in the mid-1980s an alternative viewpoint was presented concerning the taxonomic and evolutionary
affinities of Homo erectus. Researchers such as Peter Andrews, Chris
Stringer, and Bernard Wood suggested that the differences
between the Asian and African forms of Homo erectus were significant enough to separate them into different species.9 The Asian fossils would remain in Homo erectus while the African hominids
would be placed into a new species, alternately called Homo
ergaster,10 or Homo leakeyi.11 These paleoanthropologists employed
cladistics, an analytical approach that determines evolutionary
relationships based on the presence or absence of certain traits
shared between groups of fossils. Their analyses isolated what they
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perceived to be unique features shared by the Javan and Chinese
fossils that were absent not only in the East African representatives
of Homo erectus, but also in modern humans. They concluded that
Asian Homo erectus went extinct and that the African form was the
sole ancestor of modern humans.
WAS HOMO ERECTUS A DEAD END?
Is this “neo-splitting” position justified? Was Asian Homo erectus
simply a doomed twig on the bush of human evolution, leaving no
descendants among modern people? Or is it more likely that these
archaic hominids were an integral part of the human evolutionary
tree that significantly contributed to the origins of modern Asians?
Before the latter two questions can be addressed, the first must be
resolved, because the “resplitting” of Homo erectus bears directly on
the issue of this fossil species’ influence upon the evolution of
modern Homo sapiens.
After the first wave of publications claiming that Homo erectus
in Africa was a different species from its Chinese and Indonesian
cousins, other scientists turned a critical eye on the features that
were supposedly unique to the Asian hominids. For example,
Günter Bräuer and Emma Mbua studied African and Asian Homo
erectus, and later hominids from these continents and from Europe
as well. They demonstrated that the characteristics supposedly
unique to Asia were not confined to Asian Homo erectus, but were
also present in varying frequencies in the fossils from both Africa
and Europe.12
I became interested in the problem of “lumping” versus “splitting” for a variety of reasons. As a graduate student at the
University of Michigan I was trained by professors who were
decidedly “lumpers.” These scholars emphasized the importance
of factors that could produce significant variation within species,
such as sexual dimorphism (size and shape differences between
adult males and females) and polytypism (geographic variation).
My professors argued reasonably that these intraspecific factors
had to be eliminated first as potential sources of variation in fossil
samples before the presence of multiple species could be claimed.
During the latter half of my graduate studies and the beginning of
my professional career, from the mid-1980s through the early
1990s, it seemed to me that paleoanthropology was slipping “back
HOMO ERECTUS
9
to the future”—more and more publications were trumpeting the
reality of new species that were split from other well-established
taxa. I decided to investigate whether or not this trend was warranted with respect to the splitting of African from Asian Homo
erectus. However, unlike other researchers, I chose to examine this
question quantitatively by comparing skull measurements instead
of the osteological (bony) features that were being argued over by
the cladists.
My research compared Homo erectus with a sample of modern
humans (made up of over 2,500 individual skeletons from all over
the world) and another sample composed of various fossils representing 2 to 3 different hominid species (called the mixed hominid
sample).13 The modern human sample was chosen because it exemplified the nature and degree of variation to be expected in a single
but geographically variable hominid species. The mixed hominid
sample was used to depict the variation present in a sample composed of multiple fossil hominid species. If my comparisons
showed that Homo erectus was most similar in its magnitude and
pattern of variation to the mixed hominids, then this would support the cladists’ conclusion that Homo erectus included more than
one species. However, if Homo erectus proved to be no more variable than modern humans, then this would support the “lumpers’”
position that it was a single species.
The results of these comparisons would be quite important to
the reconstruction of later human evolution. If there were multiple
species of Homo erectus, this would support a very “bushy” view of
human evolution, with many origins and extinctions of regionally
isolated hominid species. This perspective would indicate that
modern humans probably arose relatively late in the Pleistocene
epoch from a geographically restricted area, such as Sub-Saharan
Africa. In this scenario, the descendants of east African Homo erectus (Homo ergaster) were the sole ancestors of modern humanity. In
contrast, if the results indicated that Homo erectus was indeed a single species, this would support a human evolutionary “tree,”
whose “trunk” was composed of Homo erectus, gradually growing
into Homo sapiens around the world.
Most people think that science is objective. This objectivity
may sometimes be illusory, however. People bring biases and preconceived notions to everything they do, and scientists are no different. It is important to acknowledge these biases whenever possible. As one can easily tell by the foregoing discussion, I was
predisposed to reconfirming that Homo erectus was a single species.
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But I did take precautions in my study not to make this a foregone
conclusion. For example, the Homo erectus sample that I used was
chronologically as broad as possible, dating from 1.8 million to less
than 500,000 years old. Because of their great time-depth, these fossils were potentially much more variable than the modern humans,
who were sampled from a single point in geological time. This
comparison, then, predisposed my study to conclude that Homo
erectus was significantly different from Homo sapiens and therefore
was composed of more than one species. In effect, I bent over backwards to prevent my conclusions from being predetermined by my
biases.
In my analyses I used eight skull measurements that were
common to all three samples (Homo erectus, the mixed hominid
group, and the modern humans). I examined each of the variables
individually and in combination, but the results of these preliminary investigations were not sufficient to resolve the Homo erectus
species problem. Whether or not the fossils attributed to Homo erectus belonged to one or more species was ultimately a statistical
question: What is the probability that the variation present in a fossil sample (such as Homo erectus) could be found in a sample taken
from a single species (such as Homo sapiens)?
To answer this question I used a randomization procedure that
is ideal for solving problems such as these. This procedure uses a
computer to generate many random samples of a particular reference species (in this case, modern humans). The variation in these
samples is then compared to that present in a fossil sample (in this
case, Homo erectus) to determine whether or not the variation in the
fossils is greater than the variation in equivalently-sized samples
randomly drawn from a known single species, such as Homo sapiens. If the variation in the fossil sample exceeds the variation in 95
percent or more of the randomized modern human samples, then it
is likely that the fossil sample is composed of two or more species.
On the other hand, if a significant percentage (defined as greater
than 5 percent) of the randomized modern human samples are
more variable than the fossils, this provides evidence that the fossil
sample represents a single species.
Let me illustrate this technique with the following example
(see Figure 2). One of the eight measurements that I recorded was
the greatest width of the brain case, formally known as maximum
cranial breadth and abbreviated as XCB. This measurement could be
taken on all 8 of the fossils in the mixed hominid sample, on 16 of
19 Homo erectus skulls, and on all 2,533 of the modern humans.
HOMO ERECTUS
11
How variable XCB was in the two fossil samples was determined
by calculating the coefficient of variation, known more simply as
the CV. To begin with, was the CV for XCB in the mixed hominid
fossil sample greater than expected for a single species, such as
modern Homo sapiens? This was answered as follows: First, eight
skulls were randomly drawn by the computer from the modern
human sample and their CV for XCB was calculated. Only eight
human skulls were taken because that was the number of fossil
specimens present in the mixed hominid sample. After the CV was
calculated, the eight skulls were returned to the human pool and
the computer randomly drew eight more and calculated this new
sample’s CV. The computer repeated this process one thousand
times, generating one thousand CVs for one thousand different
eight-skull samples. Finally, the CV for XCB in the mixed hominid
sample was compared to the modern human CVs generated by
randomization. The extremely high CV of the mixed hominid sample exceeded 99.8 percent (998 of the 1,000) of the randomized
modern human CVs, a result confirming the presence of multiple
species in the mixed hominid sample.
In contrast, the randomization procedure told a different story
regarding variation in Homo erectus (see Figure 3). This time the
computer drew one thousand samples of sixteen skulls each from
the modern human pool—sixteen skulls each because the Homo
erectus sample included sixteen individual fossil specimens. When
the CVs of the randomized samples were compared to the Homo
erectus CV it was revealed that most (81 percent) of the randomized modern human CVs for XCB were greater than or equal to
the relatively low Homo erectus CV for XCB. This result strongly
supports the retention of Homo erectus as a single species because
variation in these fossils is usually exceeded by the variation present in samples drawn from the known single species, modern
Homo sapiens. The other results from my study corroborate this
conclusion.
The fact that the Homo erectus sample, whose members were
separated from each other by thousands of miles and by over one
million years, was generally less variable than randomly drawn
modern human samples directly counters the claims of the “neosplitters.” Those who think that African Homo ergaster is a species
distinct from Asian Homo erectus would expect that the reproductive isolation necessary to produce two species would result in
the development of significant differences between the two
forms. Instead, I showed that African and Asian Homo erectus
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Figure 2
The randomization procedure randomly draws, with replacement,
one thousand eight-skull samples from the modern human sample
and determines the coefficient of variation (CV) for maximum cranial
length (XCB) for each sample. The CV for XCB in the eight mixed
hominid skulls is then compared to the one thousand CVs of the one
thousand random modern human samples. Because 99.8 percent of
the modern human CVs are smaller than the mixed hominid fossil
CV, it is likely that this fossil sample includes at least two species.
shared a level and pattern of variation most similar to modern
humans, and that their differences could be best explained by
polytypism: geographic variation within a species. Thus, this evidence suggests that Homo erectus was like us, a far-flung species
that developed regionally distinct forms that were never reproductively isolated enough from each other to evolve into separate
species.
The larger question remains: What became of Homo erectus?
Did this species gradually transform into modern humans around
the world, or did only one geographically constrained population
of Homo erectus provide humanity’s ancestry while the rest were
doomed to extinction?
HOMO ERECTUS
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Figure 3
The randomization procedure randomly draws, with replacement,
one thousand sixteen-skull samples from the modern human sample
and determines the coefficient of variation (CV) for maximum cranial
length (XCB) for each sample. The CV for XCB in the sixteen Homo
erectus skulls is then compared to the one thousand CVs of the one
thousand random modern human samples. Because 81 percent of
the modern human CVs are larger than the Homo erectus CV, it is
likely that the Homo erectus sample includes only one species.
HOMO ERECTUS AND THE ORIGINS
OF MODERN HUMANS
The debate about modern human origins has been raging in paleoanthropology for the last decade. In its most basic form, the controversy can be stated as two polar opposites: replacement versus
multiregionalism. The replacement view, supported by Stringer,
Andrews, and their colleagues, states that modern humans evolved
relatively recently (one hundred thousand to two hundred thousand years ago) in Sub-Saharan Africa and subsequently spread
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around the Old World, replacing all of the indigenous, archaic
human populations that they encountered.14 In contrast, multiregionalism, espoused by Milford Wolpoff, Alan Thorne, David
Frayer, and their co-workers, contends that archaic hominids from
throughout the Old World (not just in Africa) contributed to the
evolution of their local modern successors.15 Both of these hypotheses produce very different predictions concerning the evolutionary
fate of Asian Homo erectus: Replacement suggests that the Javan
and Chinese hominids had nothing to do with the origins of modern humans in those regions while multiregionalists maintain that
these fossil hominids are part of an unbroken genetic continuum
linking Asia’s ancient past to its present.
I decided to test these predictions with data I collected during
my dissertation research. The fossils I studied are a group of
hominid mandibles (lower jaws) from the site of Sangiran in central Java that date to approximately one million years ago. Initially
I was intrigued by these jaws because of their variability: Some
were as huge as the biggest australopithecine mandibles while others were quite a bit smaller, as is typical for Homo erectus. My studies of the anatomy and measurements of these fossils led me to
conclude that they all represented Homo erectus and that their great
size variation could be best explained by sexual dimorphism.16
Later I realized that the morphology of these mandibles could provide some important additional information bearing on the origins
of modern humans.
The Sangiran mandibles are among the oldest Homo erectus fossils found outside of Africa and as such provide an excellent test
case of the polarized predictions generated by the replacement and
multiregional models. I compared the Javan jaws to samples of
modern human mandibles from Kenya and Australia to explicitly
test the following expectations.17 According to the replacement
hypothesis, the two modern human samples should display much
more in common with one another, when compared to the fossils,
because the Kenyans and Australians presumably share a relatively recent African ancestor that had nothing to do with one million year old Javan Homo erectus. In contrast, multiregionalism suggests that the Sangiran hominids’ ancestry to modern humans in
the region, such as Australian Aborigines, would be reflected in
the number of features shared between the Sangiran and
Australian jaws that are not found in the mandibles of modern
Kenyans.
Most previous studies of modern Asian origins have generally
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presented varying lists of morphological features to promote either
the replacement or the multiregionalism models. For example,
Wolpoff and his colleagues support the multiregional position by
noting numerous skull features that are shared between Middle
Pleistocene (≈two hundred thousand to five hundred thousand
years ago) Javan hominids and recent Australians that are absent
in early modern humans from Africa.18 Critics of this interpretation
contend that these morphological similarities are due either to cultural practices of cranial deformation or parallelism (independent
evolution of the same features by ancient Javans and recent
Australians).19
Little has been done to actually quantify these similarities and
differences, particularly with respect to the earliest Homo erectus
specimens from Java. Therefore, I used a statistical test to determine which of the two predictions would be borne out by the data.
If the statistics showed that the modern humans from Kenya and
Australia shared more features in common when compared to the
fossils, this would support the replacement model. If the Sangiran
and Australian jaws were demonstrably more similar to each other
than either were to the Kenyans, then multiregionalism was a more
likely explanation. Fisher’s Exact Test of Independence is an analytical tool that can be used to answer these types of questions.
Fisher’s test determines whether trait frequencies between two
samples are significantly different (i.e., whether the differences are
unlikely to have occurred by chance). For example, suppose that
“Feature A” is present in all (100 percent) of the fossil jaws, in 90
percent of the Australian mandibles, but in only 20 percent of the
Kenyans. Fisher’s Exact Test of Independence would show that the
frequency of “Feature A” is statistically indistinguishable when the
Sangiran and modern Australian mandibles are compared, but that
the frequencies are significantly different when the Australians are
compared to the Kenyans. This was the predominant finding in my
study. Of the sixteen mandibular features analyzed by Fisher’s test,
thirteen displayed insignificant differences in the Sangiran and
modern Australian comparisons. In contrast, nine of sixteen trait
frequencies were statistically different in the comparisons between
modern Australians and Kenyans.
If the replacement model is closer to the historical truth, then
the similarities between the ancient Javans and modern Australians
could not be simply explained by an ancestral-descendant relationship. A more tortuous explanation involves the independent evolution of these similarities twice: once in the Homo erectus fossils and
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a second time among the modern Australians. Although the small
size of the fossil sample tempered my conclusions with caution,
my results clearly provide more support for the multiregional
hypothesis—namely, that early Homo erectus from Java did contribute to the evolution of modern Australian Aborigines.
WHERE DO WE GO FROM HERE?
The evidence discussed in this chapter establishes that Homo erectus
was a single, hominid species that evolved into Homo sapiens
throughout the Old World over the past million years. Ironically,
the very fact that modern Homo sapiens is the product of this gradual transformation has spurred calls for the “sinking” of Homo erectus! Wolpoff, Thorne, Jan Jelinek, and Zhang Yinyun have recently
published a paper arguing that because there is no obvious morphological gap in the direct evolution of Homo erectus into Homo
sapiens the former species should be formally “sunk” into the
latter. 20 The authors maintain that Homo erectus could only be
retained as a separate species if the fossil record indicates that the
origin of Homo sapiens was the result of a branching speciation
event, whereby the ancestral species (i.e., Homo erectus) ceased to
exist by dividing and producing two descendant species. I agree
with Wolpoff and his colleagues who do not see any evidence of
such branching evolution in modern human origins. Homo sapiens
appears to have a much greater time-depth and much more morphological variability than traditional views would assume.
Is this interpretation justified? Given the evolutionary species
concept that Wolpoff et al. employ in conjunction with the evidence from the fossil record, their logic is unassailable.21 But does
their argument illuminate or obfuscate evolutionary relationships
in labeling both big-browed, smaller-brained ancient hominids and
high-browed, bigger-brained modern humans as Homo sapiens?
This point will be debated for years to come. Most scholars would
agree, however, that the label is not important in and of itself. The
reconstruction of evolutionary patterns and relationships should
remain our primary focus.
To that end I hope that future research on Homo erectus will
concentrate on the regional patterns of evolutionary change leading to the emergence of Homo sapiens. Was the tempo of this evolution gradual throughout or was there a period of acceleration that
could be used to mark the transition, thereby rendering the bound-
HOMO ERECTUS
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ary between these two species less arbitrary? Although questions
such as these may be answered by applying new quantitative
methods to old data, the age old battle cry of paleoanthropology
will certainly continue to ring out across our ancestral homelands:
more fossils, more fossils!
NOTES
1. Much of the information in this section comes from C. Loring
Brace, “Tales of the Phylogenetic Woods: The Evolution and
Significance of Evolutionary Trees,” American Journal of Physical
Anthropology 56 (1981): 411; C. Loring Brace, The Stages of Human
Evolution, 4th ed. (Englewood Cliffs, NJ: Prentice Hall, 1991); F.
Clark Howell, “Thoughts on Eugene Dubois and the
‘Pithecanthropus’ Saga,” Courier Forschungsinstitut Senckenberg 171
(1994): 11.
2. Eugène Dubois, Pithecanthropus Erectus: Eine Menschenähnliche
Uebergangsform Aus Java (Batavia: Landersdruckerei, 1894).
3. Compare, for example, Brace, “Tales of the Phylogenetic Woods,”
with Stephen J. Gould, “Men of the Thirty-Third Division,”
Natural History (April 1990): 12.
4. The following are the references for original works in which these
genera were named: “Sinanthropus”—Davidson Black, “On a
Lower Molar Hominid Tooth from Chou-kou-tien Deposit,”
Palaeontologica Sinica Series D7 (1927): 1; “Telanthropus”—Robert
Broom and John T. Robinson, “New Type of Fossil Man,” Nature
164 (1949): 322; “Atlanthropus”—Camille Arambourg, “L’hominien
fossile de Ternifine (Algérie),” Comptes Rendus de l’Academie des
Sciences (Paris) 239 (1954): 893; “Tchadanthropus”—Yves Coppens,
“L’hominien du Tchad,” Comptes Rendus de l’Academie des Sciences
(Paris) 260D (1965): 2869.
5. Franz Weidenreich, “Some Problems Dealing with Ancient Man,”
American Anthropologist 42 (1940): 375.
6. Ibid., p. 383.
7. Ernst Mayr, “Taxonomic Categories in Fossil Hominids,” Cold
Spring Harbor Symposium on Quantitative Biology 15 (1950): 109; W.
E. LeGros Clark, The Fossil Evidence for Human Evolution (Chicago:
University of Chicago Press, 1955).
8. F. Clark Howell, “Hominidae,” in Vincent J. Maglio and H. B. S.
Cooke, eds., Evolution of African Mammals (Cambridge, MA:
Harvard University Press, 1978), pp. 154–248; William W.
Howells, “Homo erectus—Who, When, and Where: A Survey,”
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9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
RESEARCH FRONTIERS
Yearbook of Physical Anthropology 23 (1980): 1; G. Philip Rightmire,
The Evolution of Homo erectus: Comparative Anatomical Studies of an
Extinct Human Species (New York: Cambridge University Press,
1990).
Peter Andrews, “An Alternative Interpretation of the Characters
Used to Define Homo erectus,” Courier Forschungsinstitut
Senckenberg 69 (1984): 167; Christopher B. Stringer, “The Definition
of Homo erectus and the Existence of the Species in Africa and
Europe,” Courier Forschungsinstitut Senckenberg 69 (1984): 131;
Bernard A. Wood, “The Origin of Homo erectus,” Courier
Forschungsinstitut Senckenberg 69 (1984): 99.
Originally named by Colin P. Groves and V. Mazak, “An
Approach to the Taxonomy of the Hominidae: Gracile
Villafranchian Hominids of Africa,” Casopis pro Mineralogii a
Geologii 20 (1975): 225.
Originally named by G. Heberer, “Über einen neuen archanthropinen Typus aus der Oldoway-Schlucht,” Zeitschrift für
Morphologie und Anthropologie 53 (1963): 171.
Günter Bräuer and Emma Mbua, “Homo erectus Features Used in
Cladistics and Their Variability in Asian and African Hominids,”
Journal of Human Evolution 22 (1992): 79.
Andrew Kramer, “Human Taxonomic Diversity in the Pleistocene:
Does Homo erectus Represent Multiple Hominid Species?,”
American Journal of Physical Anthropology 91 (1993): 161.
Christopher B. Stringer, “The Emergence of Modern Humans,”
Scientific American 263 (1990): 98; Christopher B. Stringer and Peter
Andrews, “Genetic and Fossil Evidence for the Origin of Modern
Humans,” Science 239 (1988): 1263.
Alan G. Thorne and Milford H. Wolpoff, “The Multiregional
Evolution of Humans,” Scientific American 266 (1992): 76; David W.
Frayer, “Testing Theories and Hypotheses about Modern Human
Origins,” in Carol R. Ember, Melvin Ember, and Peter N.
Peregrine, eds., Research Frontiers in Anthropology: Advances in
Archaeology and Physical Anthropology (Englewood Cliffs, NJ:
Prentice Hall, 1995).
Andrew Kramer, “A Critical Analysis of Claims for the Existence
of Southeast Asian Australopithecines,” Journal of Human Evolution
26 (1994): 3; Andrew Kramer and Lyle W. Konigsberg, “The
Phyletic Position of Sangiran 6 as Determined by Multivariate
Analyses,” Courier Forschungsinstitut Senckenberg 171 (1994): 105.
Andrew Kramer, “Modern Human Origins in Australasia:
Replacement or Evolution?” American Journal of Physical
Anthropology 86 (1991): 455.
Milford H. Wolpoff, Wu Xinzhi, and Alan G. Thorne, “Modern
HOMO ERECTUS
19
Homo sapiens Origins: A General Theory of Hominid Evolution
Involving the Fossil Evidence from East Asia,” in Fred H. Smith
and Frank Spencer, eds., The Origins of Modern Humans: A World
Survey of the Fossil Evidence (New York: Alan R. Liss, 1984), pp.
411–483; Milford H. Wolpoff, “Multiregional Evolution: The Fossil
Alternative to Eden,” in Paul Mellars and Chris Stringer, eds., The
Human Revolution: Behavioural and Biological Perspectives on the
Origins of Modern Humans (Edinburgh: Edinburgh University
Press, 1989), pp. 62–108; Milford H. Wolpoff, “Theories of Modern
Human Origins,” in Günter Bräuer and Fred H. Smith, eds.,
Continuity or Replacement: Controversies in Homo sapiens Evolution
(Rotterdam: Balkema, 1992), pp. 25–63.
19. Erik Delson, “One Source Not Many,” Nature 332 (1988): 206;
Stringer and Andrews, “Genetic and Fossil Evidence for the
Origin of Modern Humans,” p. 1267; Christopher B. Stringer,
“Replacement, Continuity, and the Origin of Homo sapiens,” in
Günter Bräuer and Fred H. Smith, eds., Continuity or Replacement:
Controversies in Homo sapiens Evolution (Rotterdam: Balkema,
1992), pp. 9–24.
20. Milford H. Wolpoff, Alan G. Thorne, Jan Jelinek, and Zhang
Yinyun, “The Case for Sinking Homo erectus: 100 Years of
Pithecanthropus Is Enough!” Courier Forschungsinstitut Senckenberg
171 (1994): 341.
21. An “evolutionary species” can span a considerable amount of geological time and is formally defined as a single lineage of ancestordescendant populations that maintains its identity from other
such lineages and has its own evolutionary tendencies and historical fate. In contrast, the traditional “biological species” has no time
depth and is defined as a group of actually or potentially interbreeding individuals who are reproductively isolated from other
such groups.
SUGGESTED READINGS
Howells, William W. “Homo erectus—Who, When, and Where: A
Survey.” Yearbook of Physical Anthropology 23 (1980): 1–23. State of
the art in 1980 on the distribution, taxonomy, and evolution of
Homo erectus; today a bit dated.
Rightmire, G. Philip. The Evolution of Homo erectus: Comparative
Anatomical Studies of an Extinct Human Species. New York:
Cambridge University Press, 1990. State of the art today in book
length, by the acknowledged leading expert on the subject.
20
RESEARCH FRONTIERS
. “Homo erectus: Ancestor or Evolutionary Side-branch?”
Evolutionary Anthropology 1 (1992): 43–49. State of the art today in
easily digestible article length.
Wu, Rukang, and Lin Shenglong. “Peking Man.” Scientific American
248 (1983): 86–94. A fascinating and lively look at the excavations
and interpretations of the most famous Chinese Homo erectus site.
Research TOC