Homo erectus or Homo ergaster?

Homo erectus appears in the fossil record about 1.9 million years ago and lasted until
250,000 years ago, with some populations in southeast Asia that may have persisted
until as recently as 40,000 years ago. Homo erectus was the first hominin to leave the
African continent, and fossils have been recovered from sites in Africa, Asia, and parts of
Europe. Homo erectus is a highly variable species which is not surprising, since it lived in a
broad geographic range for over 1.5 million years. Many scientists suggest that the
differences between the physical characteristics of Asian and African specimens are
significant enough to justify separation at the species level (Tattersall 2015). The
term Homo erectus is sometimes used to describe east Asian fossils exclusively,
while Homo ergaster is the species designation for the African specimens that appear in
the fossil record around 1.9 million years ago. Whether the variation in this widely
dispersed group represents two species, or one highly variable species continues to be
debated. It is still common practice for paleoanthropologists to use Homo erectus as a
broad, all-encompassing term that includes African and Eurasian hominins that exhibit a
shared suite of physical characteristics. While some physical differences between Asian
and African Homo erectus are acknowledged, the remainder of the module will follow
that approach and consider them all together as Homo erectus.
Homo erectus Physical Characteristics
Homo erectus exhibit a number of unique physical characteristics. The skull of Homo
erectus has a distinctive shape. The skull is low and long with a sloping forehead. The
cranium is widest at the base, and when viewed from the back, it has a somewhat
pentagonal shape. The Homo erectus skull has a postorbital constriction, which is a
narrowing of the skull in the region behind the eye orbits (see Figure 5-7).
Many Homo erectus fossils (particularly the east Asian specimens) possess a sagittal
keel – a small ridge of bone that runs from front to back along the top of the skull. The
back of the skull has an occipital torus, which is a bulging appearance in the occipital
region with a large area for muscle attachment.
Homo erectus has defined brow ridges above the eyes. This horizontal ridge on the
frontal bone above the eye sockets is called the supraorbital torus.
The face of Homo erectus is large. It is less prognathic than earlier hominins, but more
prognathic than later members of the genus Homo. Compared to modern humans, Homo
erectus has large teeth and jaws. The lower jaw does not have a chin.
Homo erectus exhibits significant increase in the size of the brain. Most Homo
erectus have a cranial capacity between 800cm and 1200cm with an average about
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1000cm . This is about two-thirds the size of an anatomically modern human (Homo
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sapiens) (see Figure 5-1, which compares the cranial capacity of Homo erectus to that of
the genera and species discussed in previous modules, as well as to Homo
floresiensis and Homo naledi – two more species which you will learn about later in the
module).
Homo erectus has a larger body size than earlier hominins and modern limb proportions.
It is the first hominin that was fully committed to a terrestrial life, having lost their
arboreal adaptations. There is variation in body size, but Homo erectus are typically larger
than earlier hominins with an average height of 1.7 metres and average weight of 58 kg
(Walker 1993). The Homo erectus post cranial skeleton is similar to a modern human in
size and body structure.
Figure 5-2 illustrates the physical characteristics of Homo erectus and demonstrates
some differences in skull shape between Indonesian (Homo erectus) and
African (Homo ergaster) fossils.

Homo erectus - A Bigger, Smarter, Faster Hominin Lineage

About two million years ago, a new set of fossils began to appear in the
human fossil record. Designated as Homo erectus, they show evidence of
increases in both body size and brain size.
Homo erectus is arguably the earliest species in the human lineage to have so many human-like qualities.
Earlier hominins had important similarities with living humans, like bipedality, and H. erectus still had a
long evolutionary path to become like you and me, but the fossils assigned to H. erectus display a number
of new and distinctly modern human traits.
Homo erectus is often referred to as the first cosmopolitan hominin lineage, meaning the first hominin
species whose geographic range had expanded beyond a single continental region. While fossil remains
from H. erectus are found in Africa, like those of earlier hominins, they have also been identified at fossil
sites widely dispersed across Eurasia.

There are a number of fascinating evolutionary questions that can be asked of H. erectus. The species was not only
geographically widespread, it also had a long temporal span in the hominin fossil record (Antón 2003). With its earliest
appearance in the fossil record from localities in the Lake Turkana Basin, Kenya, sometime around two million years ago, H.
erectus populations persisted until near the end of the Pleistocene, as evidenced by fossils from Southeast Asia. Homo
erectus thus presents paleoanthropologists with the challenge of trying to interpret fossil variation in the context of both
widespread geographic and temporal distribution.
Furthermore, the expansion of H. erectus across a large range of environments suggests a change in the ecology of this
lineage relative to early hominins, a change that certainly has significance for how evolutionary forces acted to shape the
pattern of variation we observe in the fossil lineage.
These are some of the questions that researchers ask of H. erectus fossils: How did the ecology of Homo erectus differ from
that of preceding hominins? What are the characteristics of H. erectus that allowed it to expand across different habitats
throughout portions of Eurasia and Africa? What limitations constrained the expansion and evolution of H. erectus in the
Pleistocene? What role did behavioral and technological innovation play in establishing the complex and geographically
widespread evolutionary pattern of H. erectus? How might we describe and explain the evolutionary pattern of H. erectus?

History and Geography

Eugene Dubois first identified and described a new human-like set of Indonesian fossils at the end of the 19 th century,
naming the specimens Pithecanthropus erectus (upright, ape-man) because of their combination of bipedality and a brain
size much smaller than living humans. Dubois had specifically been looking for the missing link between apes and humans,
and for him the combination of a human-like body and ape-like brain represented just that (Shipman 2002). Subsequent
discoveries in the 1920s and 1930s from the site of Zhoukoudian, China, of fossils with similar characteristics-originally
designated Sinanthropus pekinensis-raised the question of a possible evolutionary relationship between these regional
samples. Today, these two samples, along with a much larger collection of fossils from Asia, Africa, and Europe, are most
commonly referred to simply as Homo erectus.
What is the evolutionary relationship among fossils that share many similarities, but also subtle differences, separated
across time and space? This question, prompted by the early Chinese and Javan fossils, remains an active research
question today for the much larger sample of fossils assigned to H. erectus. Whether or not a sample from one region, for
example Africa, part of a polytypic, geographically widespread lineage (Homo erectus), or whether it is part of a related, but
different species, is a debated topic and reveals much about the evolutionary pattern of the species (Rightmire 1998). For
example, some researchers argue that H. erectus is restricted largely to Eastern and Southeast Asia, consistent with the
original fossils attributed to the taxon. In that case, the bulk of its representatives lived from the end of the Lower Pleistocene
through the Middle Pleistocene (~1.4-0.2 mya). From this perspective, earlier fossils from Western Asia (e.g., Dmanisi,
Georgia; Figure 2) and Africa (e.g., Koobi Fora, Kenya) that are similar to the classic Asian H. erectus, but also have
more ancestral traits, might be considered a separate lineage (often called Homo ergaster). Middle Pleistocene remains
from Europe might be a second or third separate lineage (Homo heidelbergensis). In this view, the ecological
niche occupied by these species is more limited, leading to the isolation, and ultimately speciation, among different regional
populations.

Humans are widespread and variable today, but much of the variation observed across contemporary populations is the
result of relatively recent events in the past 100,000 years of our evolutionary history. Patterns of variation in H.
erectus occurred on a time scale as long as a million years, and may have been different from those we observe today. This
presents a challenge for researchers in terms of how we explain the pattern of variation seen in H. erectus, but also presents
an opportunity to study how evolutionary forces operate across such scales.

Who was Homo erectus?

As with any fossil lineage, identifying the earliest appearance of the species is difficult. Nevertheless, a set of
shared, derived features can be assigned to all of the fossils assigned to H. erectus, regardless of where they fall amid the
geographic and temporal range of the lineage. The following sections summarize some of these characteristics.

Increased body size

One of the traits most commonly associated with Homo erectus is an increase in body size. The Nariokotome specimen, an
adolescent male individual, was over five feet tall at the time of his death (Walker & Leakey 1993; Figures 3 & 4). Estimates
of his adult stature, had he continued to live to adulthood, differ depending on how researchers estimate his age at death
based on his teeth and bones (for more see Smith & Alemseged NKP article LINK HERE) and the amount of growth he had
left. Living humans generally experience a marked increase in growth during early adolescence (i.e., a ‘growth spurt'), a
growth pattern that some researchers say distinguishes us from other apes. If early H. erectus had a human-like growth
spurt, Nariokotome likely had a lot of growing left to do. Even if H. erectus did not have such a modern human-like pattern of
growth, the specimen was clearly a tall individual relative to earlier hominins. Not all H. erectus were tall, however, as earlier
fossils remains from sites like Olduvai Gorge in Tanzania and Dmanisi, Georgia, attest. It is important to note that variations
in size, not just an increase in size over that of earlier hominins, is characteristic of H. erectus, much like living humans.

Increased brain size/encephalization

As the body size of hominins increases, the brain size increases as well (Ruff et al. 1997). While the smallest-bodied
early H. erectus fossils have brain sizes only slightly larger than earlier hominins (australopiths), early large-bodied
specimens, such as the Nariokotome individual, have a brain volume greater than 800 cm 3, more than 50% larger than
earlier australopiths (and about 60% of the typical brain size of someone living today). However, in addition to the absolute
increase in brain volume that accompanies an increase in body size, there is also a proportional increase. This is referred to
as encephalization, and is an important characteristic of H. erectus. Throughout the evolutionary history of H. erectus there
is substantial evidence for selection leading towards increased encephalization, so that while early members of the lineage
have a cranial capacity of 600-800 cm3, the cranial capacities of most later specimens are well in excess of 1000 cm 3, which
is within the lower range of contemporary humans, without appearing considerably larger in body size than early H. erectus.

Increased technological/ecological intensification

The large body and large brain of H. erectus needed more energy, and thus food, than previous hominins. Larger biological
structures, particularly energy-intensive ones like muscles and brains, require greater energy inputs to maintain. Thus, H.
erectus is often reconstructed as occupying an intensified ecological niche (Leonard & Robertson 1997).
The intensified niche goes hand in hand with the expansion in brain and body size. Larger bodies, and longer limbs in
particular, increase locomotor efficiency (Pontzer et al. 2010). Homo erectus could cover more ground on a day-to-day
basis, through walking or running, than smaller hominins and with lower energy cost. In addition, the larger brain gave these
hominins better capabilities for processing complex ecological information across the more expansive terrain containing
higher quality food items. For example, there is clear evidence of H. erectus accessing medium- and large-sized animal
carcasses for meat, through hunting and/or scavenging, in the form of fossil remains of animals with cut marks left by
butchery. This behavior, regularly accessing animal carcasses, is an ecological change from earlier hominins (Link to
Pobiner's NKP article). While the earliest H. erectus specimens are found in association with very basic stone tools, typically
referred to as the Oldowan stone tool industry, by 1.5 million years ago populations of H. erectus were creating a more
complex and typologically codified set of tools that we refer to as the Acheulean industry (Bar-Yosef & Belfer-Cohen 2001).

Reduced post-canine dentition size

The change in ecology associated with H. erectus coincides with a corresponding change in the dentition and jaws of this
species. Relative to earlier australopiths and contemporary robust australopiths (Paranthropus), the size of the post-canine
dentition (premolars and molars) and the molarization of the premolars are dramatically reduced in H. erectus. The corpus of
the mandible (i.e., the toothless part that connects to the cranium) also displays increased gracility (i.e., slenderness), with a
characteristic reduction in the relative breadth of the structure and supportive masticatory structures. Toothwear analyses
suggest that across their range, H. erectus engaged in a diverse and broad diet (Ungar et al. 2006). The food an organism
ingests can also produce subtle changes in the chemistry of body tissues (you actually are what you eat), including the
dentin and enamel that make up the crown of a tooth. Using this information, investigations of the stable isotope chemistry
of H. erectus teeth also support the idea of a flexible and diverse diet (Ungar & Sponheimer 2011). Whatever the flora and
fauna H. erectus ate at the varied geographic localities where H. erectus fossils are found, their tooth and jaw anatomy
reveal that their diet did not require the same robust masticatory adaptations seen in earlier hominins.

Unanswered Questions About Homo erectus

Two of the more significant, yet elusive, questions about H. erectus concern the levels of sexual dimorphism within the
lineage and the capacity for language. Sexual dimorphism, the physical differences between males and females, is an
important source of variation within species, and in primates can be an indicator of reproductive strategy and group
dynamics. Sexual dimorphism, given its role in intraspecific variation, can also be a confounding factor in proper taxonomic
identification. The large amount of size variation observed within H. erectus, taken primarily from fragmentary fossil remains,
makes it difficult to estimate average levels of dimorphism. The H. erectus fossil record provides clear evidence of a large
range of skeletal size variation, at least equivalent to that observed in living human populations, but it does not provide
conclusive evidence that males were systematically larger than females to a greater extent than they are today. If H.
erectus did have more sexual dimorphism than H. sapiens, we would infer that male competition for mates was more
dependent on body size than it is today.
Language is perhaps the hallmark human trait, but can be difficult to assess directly from the fossil record. Attempts to
identify language ability in the fossilized skeletal remains of H. erectus have focused on aspects of the nervous system,
including the size of the vertebral canal (a proxy for the size of the spinal cord), and external features of endocasts (natural
fossils of endocranial space and a proxy for brain size and shape). Thus far, there have been no definitive anatomical
findings to cause researchers to reject the idea that H. erectus was capable of some kind of human-like proto-language.
More recently, arguments about the origins of language have focused on the reconstructed histories of genes associated
with language production in humans. The recovery of ancient genetic sequences from Neandertals and other archaic human
specimens (e.g., a specimen from Denisova Cave in Siberia, Russia) have provided new insight into the genetic history of
language production. The human FOXP2 gene exists in a derived form in humans today and appears to play a critical role in
language development. The identification of the human form of FOXP2 in both Neandertal and Denisovan genomes
suggests this gene likely goes back at least to the Middle Pleistocene, with H. erectus a possible source lineage (although
there is no H. erectus ancient DNA to test this hypothesis). This does not suggest H. erectus had well-developed language
capabilities but, like the anatomical evidence, does not provide any evidence to reject the idea.
Summary
Homo erectus represents a significant transformation from previous hominins, like the australopiths, to a species much more
similar to modern humans. Relative to their australopith forebears, Homo erectus was bigger, smarter, and more able to
occupy and survive in differing landscapes in a changing world. The movement towards a more ecologically intense,
cognitively reliant, and behaviorally malleable adaptive pattern set the stage for the evolutionary change that followed in the
Pleistocene, up to and including the present. In many ways, modern humans are just an updated version of our H.
erectus ancestors.

Glossary
Acheulean - Lower and Middle Pleistocene hominin stone tool industry. The Acheulean tool complex is often characterized
by a high percentage of bifacially flaked stone cores and the presence of tear-drop shaped tools referred to as ‘hand axes'.
ancestral - A trait that is present in the common ancestor of a species. The large body size of contemporary humans is
ancestral, as evidenced by the presence of this feature in Homo erectus.
derived - A trait that is not present in the common ancestor of a species, but is newly arisen. The marked encephalization of
Homo erectus is a derived characteristic relative to earlier, small-brained hominins.
ecological niche - The overall set of relations that defines the place of a species within its environment. This includes the
other organisms a species interacts with, such as prey or predators, as well as the physical habitats a species utilizes in its
existence.
endocasts - A natural fossil cast formed within the endocranial space of a skull. When present, endocasts provide some
resolution on the size, shape, and surface structures of the brain in fossil taxa.
encephalization - Expansion of the brain relative to body size. Encephalization represents an increase in proportional
resources dedicated to the growth, development, and maintenance of brain activities.
masticatory - Of, or relating to, the chewing structures of an organism.
Oldowan - The earliest well-characterized hominin stone tool industry, present in the terminal Pliocene and Lower
Pleistocene. This tool complex is characterized by a simple core-flake set of tools.
polytypic - The presence of multiple forms of a lineage across a species' range.
sexual dimorphism - The characteristic differences between males and females within a species. Often this refers
specifically to size sexual dimorphism, the average difference in body mass or skeletal size between males and females of a
species.
Homo erectus in Africa
The oldest Homo erectus fossils come from Africa. Several Homo erectus fossils have been
identified at Koobi Fora, Kenya including KNM-ER 3733. This specimen is a nearly
complete skull with a cranial capacity of 850 cm3 . This is the oldest
African Homo erectus cranium, dating to 1.7 million years ago (Lepre and Kent 2010).
Note the brow ridges and low sloping forehead of this specimen (see Figure 5-3). This
specimen is identified as a female, because it is less robust than known male specimens
like the Turkana boy. It is identified as an adult based on closed cranial sutures, the
presence of all the permanent teeth and the degree of wear on those teeth (Johanson
and Edgar 2006). This fossil, like other African Homo erectus fossils, is often identified
as Homo ergaster
The Nariokotome Boy (KNM-WT 15000) skeleton, also called the Turkana Boy, is a remarkable
discovery. Discovered in Nariokotome, West Turkana, Kenya, this is the most complete early
hominin that has ever been discovered. Much of the postcranial skeleton and a nearly complete
cranium were recovered (See Figure 5-4). This fossil is remarkable for its completeness which
allowed paleoanthropologists to determine the stature and limb proportions of Homo erectus.
This skeleton demonstrates that Homo erectus was fully bipedal and had lost its climbing
adaptation. Ar/ Ar dating reveals he lived 1.47 million years ago (McDougall et al. 2012). He is
40 39

estimated to be about 12 years old and was already 1.68 m tall, and unfused growth plates
suggest that this hominin was not yet full grown (Brown et al. 1985). Some physical
anthropologists suggest that Homo erectus may have developed more quickly than modern
humans, therefore, an age estimate of 8 years old may be more accurate (Dean and Smith 2009).
Recent skeletal analysis suggests that at full growth he would have had a stature of 176-180 cm
and weighed 80-83 kg (Ruff and Burgess 2015). This is the same range as Homo sapiens.
Nariokotome boy had a cranial capacity of 880 cm , and it is suggested that would have increased
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to 909 cm at maturity (Johanson and Edgar 2006). This fossil, like other African Homo
3

erectus fossils, is often classified as Homo ergaster.

Homo erectus fossils have also been recovered from Olduvai Gorge, Tanzania which, as mentioned
in previous modules, was also the location of australopithecine and Homo habilis fossils. An
interesting fossil from Olduvai is OH-9. This specimen has a large cranial capacity of about 1000
cm with a low, sloping forehead and heavy brow ridges (see Figure 5-5). Similarities between the
3

size and structure of OH-9 and specimens from Indonesia and China are noted by Rightmire
(1990) who suggests that OH-9 with its large brow ridges and large cranium may represent a
male Homo erectus.

Hominin Migration beyond Africa

Homo erectus is recognized as the first hominin to migrate beyond the African continent.
Figure 5-6 identifies the location and dates of some early Homo sites outside Africa. The
age of the oldest sites outside of Africa indicates that very soon after Homo
erectus appears on the scene it begins to move into Eurasia. Why this migration took
place is a matter of debate, but it seems that Homo erectus was able to adapt to a wide
variety of environments.
Homo erectus in Georgia
Homo erectus is the first hominin to leave the African continent. Dmanisi, Georgia, a
Eurasian country east of the Black Sea has the earliest well-dated evidence for the
migration of Homo erectus beyond Africa. Stone stools indicate a hominin presence at
the site as early as 1.85 million years ago (Ferring et al. 2011). Excavations have also
yielded numerous hominin fossils that date to about 1.75 million years ago (Vekua et al.
2002). A total of five crania and many post-cranial elements have been recovered from
the site. The hominin-bearing levels also contained stone tools similar to Oldowan tools
manufactured by both Homo habilis and early Homo erectus. Although the fossils share
some similarities in structure to Homo erectus, the bodies and crania from Dmanisi are
smaller than is typical for Homo erectus. A compete skull with a large prognathic face and
small braincase of only 546 cm3 exhibits similarities to earliest Homo from Africa
(Lordkipanidze et al. 2013). The size of the other skulls is also similar to Homo
habilis ranging between 600 cm3 to 775 cm3, but characteristics of the skulls most closely
link them to African Homo erectus (Gabunia 2000, Rightmire et al. 2006, Lordkipanidze et
al. 2013). Paleoanthropologists suggest that the Dmanisi hominins appear to represent
the initial dispersal of Homo ergaster from Africa, and they may represent the branch
from which Asian Homo erectus was derived (Rightmire et al. 2006). The discovery
of Homo erectus at Dmanisi, Georgia demonstrates that the initial migration took place
prior to a significant increase in body and brain size.

Video

We are the paradoxical ape. Bipedal, naked large-brained. Long the

master of fire, tools and language, but still trying to understand ourselves.
Aware that death is inevitable, yet filled with optimism. We grow up slowly. We hand
down knowledge. We empathize and deceive. We shape the future from our shared
understanding of the past. CARTA brings together experts from diverse disciplines
to exchange insights on who we are and how we got here. An exploration made
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- [Philip] We've heard a lot at this point about the evolution of hominids in Africa.
It's a complicated history for sure. Let me at this point just cut to the chase and
say that humans moved out of Africa probably just after 2 million years ago
and it will be that part of the record that I want to emphasize this afternoon.
The site of Dmanisi in the Georgian Caucasus is very important, records the
oldest known, at this point, the oldest known occupations of Eurasia beginning
before 1.85 million years ago. We don't actually have human remains that are that
old, but certainly there are stone tools approaching that date. The good part about
Dmanisi is that in fact we have not just scraps of headlamp and bumper and so
forth, but virtually whole skeletons and a number of them. Now we have five skulls in
various states of repair or disrepair. Along with them, there are post cranial
bones associated with one juvenile individual particularly perhaps but not
clearly associated with one of the adults. Also the material is extremely
well-preserved. We're very fortunate in that respect. Along with the humans, of
course there are animal bones and many, many of them and there is a very complete
lithic record to go along with this material which is well-preserved, as I
say, in a very carefully studied stratigraphic context. Several of the
specimens from Dmanisi have been in the past likened to African Homo erectus but
the skeletons are quite primitive. One of them in particular is strikingly so, Skull
5 which I will talk about. At the moment, I think it's fair to say that the
taxidermic identity and the paleobiological significance of the
Dmanisi materials remain controversial. Certainly there have been plenty of
suggestions and I'm afraid I've been responsible for some of them but we'll see
where things go. Dmanisi is situated in Georgia. Georgia is stuck there between
the Black Sea and the Caspian with Azerbaijan off to the east. From Tbilisi,
the capital, it's about an hour and a half, an hour and 45 minutes ride. Roads
are pretty good these days. Roads were terrible 15 years ago. Things have
improved. Down to the site, Dmanisi is just a few kilometers from the Armenian
border. This is the obligatory excavations in progress slide. There are about four
meters of sedimentary deposits at Dmanisi. Much of this stuff is volcanic in origin,
it's very ashy. There are some other sediments and silts, but ash is always a
primary component which is a good thing. The stratigraphy is complex, all of the
sediments are piled atop the Mashavera basalt which is about 1.85 million years
old, that's the bottom of the site, the earliest record, 1.85 million years is the
date obtained from radiometric methodology. It is secure. The
stratigraphy at the site is complex partially because there are a number of
piping features. Water was present near the site during periods of heavy rain and
so on. Pipes formed underground and then progressed through breaching to collapse
towards the surface filled with sediments then got buried again. So it's been a
mess, it's been very hard to sort it out. Our geologist, Reid Ferring, has done a
huge amount of work in this respect, huge in the Trumpian sense. But there have been
problems. The first traces of human material were found at Dmanisi in 1991.
Excavations in fact have been underway at the site for quite a period of time before
that. The site is underneath an old medieval town that was on the Silk Road.
The archeologists were busy at Dmanisi for some time poking around the foundations
of the old buildings and eventually they began to dig up stuff that didn't seem to
belong there, not just the goats and fish bones from medieval suppers, but things
that looked quite antique indeed. The paleontologists came in and ascertain that
yes, the material was ancient, deeper excavations got underway and in 1991, the
folks at Dmanisis were rewarded with this jaw, the D211 mandible. It's remarkably
complete, not all of it is there. But what there is remarkably well-preserved. It's a
small jaw and in a number of respects, it does look like Homo erectus. You've seen
one reference to this specimen already. The teeth are about right for Homo erectus
as are the proportions of the mandible itself. The cranium which turns out to be
the match to the little mandible was found later in 1999, D2282 is a small cranium,
very small capacity, surprisingly so for Homo erectus. Only a bit more than 650 CCs
in this case. Despite the small brain, the thing does share a number of
characteristics with particularly early African erectus. This hulk turned up at
the site in 2005. It's way down at the bottom of the site and within a few days
after the fossil had been uncovered and cleaning was underway prior to trying to
lift it out, there was a very heavy rain. Things were very nearly washed out. Of
course we have a cover over the site, there is protection, but it rained so much
and so long that water began to trickle in around the sides of the excavations and
things were dicey for a while. Fortunately, D4500 survived. There it is
all cleaned up. It turns out that the cranium found in 2005 is a perfect match,
once again, to a mandible, D2600, which had been found earlier in the year 2000.
The upper and the lower, the cranium and the mandible simply clicked together once
the stuff have been cleaned off, there was no doubt at all. There is some pathology
on the mandible that matches, comparable pathology in the region of the ear of the
cranium so there is no doubt about the match. More than other Dmanisi hominins,
Skull 5, as this one is known, exhibits very robust morphology. It's pretty
clearly a male individual. Determining sex in the case of these fossil hominins is
often tricky, often can't be done very accurately. But in this case, we think we
have a match, the skull says male all over. Such a pattern given the fact that
it has the smallest brain of all the Dmanisi hominins because it is unexpected.
Normally for other primates, humans too of course, but for primates, higher primates
generally, males tend to exceed the females in brain size by something like 8
to 10 to 15%. So, having the tiniest brain attached to the most robust cranium and
jaw is a bit surprising. You can see that there is a good deal of variation within
the Dmanisi assemblage. The little jaw goes with a smallish brain case which is
quite gracile in its construction. We peg that one Skull 2 as likely a female. Skull
5, on the other hand is much more robust, clearly distinctive in a number of
respects. Number 1 is like Homo erectus, number 4 is a small individual. That one
seems to have lived to a ripe old age since it had lost almost its entire
dentition, maybe one tooth was still in place at the time the individual died.
That one may or may not be male, we're not sure. Anyway, there is a great deal of
diversity at Dmanisi. The crania do look different. This raises the question of
"How many species might be documented at the site?" This is a question that's been
plaguing us for some time. I think myself on the basis of the shared anatomy among
the Dmanisi individuals, they have a common bow plan extending not just to the
cranial vault but to the face insofar as we have it represented. And also to the
details of the cranial base suggest, the common bow plan suggests that all of the
individuals are drawn from just one group. We've done extensive resampling analysis
as well which cause us to come to the same conclusion that really, in fact, the
skulls, the post-cranial remains that go with them, are drawn from just one
population. Now there is stratigraphic evidence relating to this question. It
doesn't solve the question of course, but it's important information. It's good to
know that the material was all washed into these deposits or arrived in the site by
one means or another at about the same time that is the duration here can't be
more than a few hundred or perhaps a thousand years or so according to the best
analyses conducted by the geological side of the team. We'll say, then, that it's
very likely that the Dmanisi assemblage samples a population belonging to a single
species. I know there may be objections to this. I'm sure there will be, there have
been in the past. If it's true, then such a situation is quite rare. Of course, at
most localities where hominins are discovered, you've heard a lot about East
Africa at this point, Koobi Fora, Olduvai. Also at Sangiran in Java where there are a
number of fossils. The material is scattered through a very long sequence of
deposits covering a long period in time. Time as a contributor variation just
cannot be discounted. If the Dmanisi fossils document what we can call a
population in the past extending over quite a number of years of course, then
the next question, the next important question is how the Dmanisi sample may
relate to the hominin taxa that have previously been recognized. Skill 5 of
course has a very small brain case, a very large projecting face in the vault. Also
in the basal cranium, there are some resemblances, not a lot, but some
resemblances to Homo erectus. Skull 3 which I have not shown you a picture of
before is the sub-adult from Dmanisi. Skull 3 is pictured here down below. Skull
3 is similar to Homo habilis. This is true for the bow ridge, the extended brow ridge
development. It's true particularly for the shape of the vault, the rounding at
the back and for the mid-facial profile. Skull 3, I must point out, is sub-adult so
we must allow for some extra growth to have occurred if the individual had grown
up. It might have looked, had it grown up, a bit more robust and a bit like the skull
to the left there, Homo habilis KNM-ER 1813. Skulls 2 and 4 also have their
peculiar aspects, of course. They have a number of primitive characters that they
also share some features with Homo habilis. So, which species? There is, as
I've pointed out, much variation within the Dmanisi paleodeme. This is not an easy
question, the question as to which species may be represented. Skull 5, the very
small brained and very robust and very primitive looking individual does indeed
share some characters with Australopithecus as well as Homo. So
perhaps in this case, the line, the division between Australopithecus on the
one hand and earlier Homo on the other is not so clear cut after all. Many of these
shared similarities are primitive characters and unfortunately they don't
help us much in answering key questions about phylogenetic affinities. Other
characters expressed in the Dmanisi materials are Homo erectus-like and pretty
clearly they are specialized characters, characters that have changed during the
course of evolution, characters that are said to be derived. These characters
include the form of the brow ridge for example which is larger and bar-like, a
little bit of midline, keeling on the vault, details of temporal bone
construction, things of that sort on the underside of the vault. Indeed, when
Skulls 1 and 2 were first described back in the year 2000, they were grouped with
early Homo erectus from the Turkana basin. If the fossils are included with Homo
erectus, clearly that's one way to deal with the material is simply to lump it
with Homo erectus. If that is the course we take, then it must be recognized that
the boundaries between Homo erectus on the one hand and other early Homo taxa will
become less distinct. It will be particularly difficult to distinguish
early Homo erectus, African Homo erectus from specimens attributed to earlier Homo
to Homo habilis in particular. Homo habilis is considered apart from Homo
rudolfensis. So, to sum up at this point, here is again a speciose view of Hominin
phylogeny done by Bernard Wood with Meave Leakey several years ago. You can read
the
caution sign. To sum up then, there is apparently no simple answer to the
question as to which species may be represented at our site. Indeed this
question is often a tricky one. It's been a hard one for paleoanthropologists to
deal with for a long time. In one view, this view expressed on the slide, Dr. Wood
showed you another version of this very spaciose hominin phylogeny. In that view
hominin evolution has produced a veritable flowering of lineages over more than 6
million years. Such bushiness, as it were, is particular evident for the 2.5 to 1
million year ago interval. This interval in which Paranthropus on the one hand,
Australopithecus and Homo are represented by multiple species for each group. At
Dmanisi the fossils seem clearly to be Homo. Now there are some points of overlap
with Australopithecus as I pointed out, but I would say unbalanced the evidence
favors grouping all of our fossils with the genus Homo, very little doubt about
that. At the same time, the assemblage at Dmanisi does not fall neatly into one of
the taxonomic packages that have been proposed: Homo habilis, Homo rudolfensis,
Homo ergaster, Homo erectus and so on. If I were pressed and I do feel pressed at
this point, given the morphological resemblances of Dmanisi to both Homo
habilis in a very strict sense, just those fossils allocated to Homo habilis, not to
Homo rudolfensis. Given the resemblances of our material to Homo habilis, and to
early Homo erectus, particularly African Homo erectus, I would probably argue, I
will argue, that it is most reasonable to place all of these fossils within a single
evolutionary species. I would say that the Dmanisi fossils constitute just one
population within this unbranched lineage. Now, this is not to raise the specter of
just one species at a time, or to suggest that there isn't a great deal of diversity
in the hominin record, clearly there was particularly in that interval after, about
2.5 million years. But as far as our evidence is concerned, it seems to me the
best way to go, simply to place all the fossils within one evolutionary species.
Then of course we'd have to argue about what to call it, but this is not the place
for that. So, with that, thanks for listening. Thanks very much.
♪ [music] ♪
English

Homo erectus in Indonesia

The first Homo erectus fossils were discovered in 1891 by Eugene Dubois at Trinil on the
Indonesian Island of Java. A hominin calvaria (top of a skull), femur, and teeth (see Figure
5-7) were identified by Dubois (1896:241) as representing “a great man-like
mammal…considered an intermediate between man and the apes and named
accordingly Pithecanthropus erectus.” The name he assigned to the fossils means ‘erect
ape man’. The skullcap had a low, receding forehead and estimated cranial capacity of
1000 cm3. Dubois (1896) recognized this hominin as a premodern human and
placed Pithecanthropus “directly in the stem of the human race.” This initial discovery has
subsequently been reclassified as Homo erectus, and since the initial discovery,
many Homo erectus fossils have been discovered on the Indonesian island of Java and
throughout the world.
The time frame that Homo erectus occupied Java is remarkable. Some of the fossils are
ancient, indicating that Homo erectus had rapidly migrated to southeast Asia. Dates as old
as 1.66 million years ago have been reported for the site of Sangiran (Swisher et al.
1994), and Ar/ Ar ages indicate that Homo erectus occupied the Solo Basin of Java at
40 39

least 1.5 million years ago (Larick et al. 2001). While these dates are ancient, there is also
late fossil evidence that suggests Homo erectus populations survived until relatively
recently on Java. Homo erectus fossils from the sites of Ngandong and Sambungmacan
are reported to date as recently as 40,000 to 70,000 years ago (Yokoyama et al. 2008).
Homo erectus in China
Zhoukoudian is the most well-known Homo erectus site in China. Excavations in the
1920s and 1930s uncovered many hominin fossils. Canadian anatomist Davidson Black
first identified the hominin fossils at Zhoukoudian as Sinanthropus pekinensis, but they
were later reclassified as Homo erectus. These fossils are often called the “Peking Man”
fossils. Most of the specimens recovered from Zhoukoudian date between 300,000 to
600,000 years ago (Johanson and Edgar 2006). Figure 5-8 illustrates a composite Homo
erectus skull from Zhoukoudian. Note the low sloping forehead and large supraorbital
torus which are typical Homo erectus traits. The skulls recovered from the site have an
average cranial capacity of 1,043 cm (Johanson and Edgar 2006). Davidson Black
3

suggested Homo erectus used fire at Zhoukoudian. This has been debated for decades,
but recent investigations at the site along with laboratory analysis of sediments have
confirmed that levels dating to about 300,000 years ago contain clear evidence for the
controlled use of fire by Homo erectus (Gao et al. 2017)
Mystery surrounds the whereabouts of the original Zhoukoudian fossils. Work at the site
was halted because of World War II. When the Japanese invaded China, it was feared
that the fossils would be lost or damaged in the midst of conflict. A plan to ship the
fossils to the United States for safe keeping was put in place. The fossils were crated and
placed aboard a train headed to the coast while guarded by U.S. marines. The fossils
never made it to the United States. Unfortunately, en route to the coast, the marines
were taken prisoner and the crates of fossils were lost. Although numerous monetary
rewards have been offered for their return, they have never been located. Luckily,
scientists created casts of the fossils prior to their loss, and the casts are still available for
study.
Video 3
Dian this is lanu mountain in the
village of Jan Southwest of Beijing lonu
means animal fossils the s's long been
important because fossils are used in
Chinese
medicine tests on some of them in 1926
however revealed they weren't of animals
but of human teeth estimated to be
between 4 and 500,000 years
old the ancestor who had them became
known as peking
man the discovery of traces of an
unknown human ancestor shook the world
of anthropology and led to large scale
excavations an almost perfect skull of
peing man was discovered on the 2nd of
December
1929 the forehead was much lower and the
eyebrows jutted out more than modern
man's its less rounded head is believed
to have contained a brain about 2/3 the
size of the human brain
today this is a reconstructed bust model
of a peaking man the discovery of
peaking man's remains has been
indispensable in learning about
Evolution this photo was taken the day
after the
discovery the skull was wrapped in
layers of newspaper and kept in
plaster P when Jong made the find he is
often referred to as the father of
Chinese
archaeology subsequent excavations led
to caves where peaking men lived various
kinds of stone tools were
found peaking man used different sizes
and degrees of hardness according to the
task in
hand some remains indicated the use of
Fire
this is fossilized Ash peaking man used
fire to cook their prey the discovery
meant the oldest historical evidence of
human use of fire had to be Rewritten a
few hundred, years earlier than
previously
thought other skull fragments were later
found making a total of five skulls but
they were lost during the second world
war excavations resumed afterwards
significant peaking man's skull
fragments were found in
1966 a perfectly reconstructed model of
a peaking man skull was shown to the
public for the first time in China in
2003

Zhoukoudian is the richest early hominin site in China, but it is not the oldest. An earlier hominin
presence has been confirmed in the Yuanmou Basin of southwest China. Here, hominin fossils
with affinities to Homo erectus have been discovered along with stone tools that are typical of the
Oldowan tool industry. These materials have been dated to 1.7 million years ago. The incisors are
identical to the incisors of Homo erectus from Zhoukoudian (Zhu et al. 2008). The Nihewan Basin
in northern China has also revealed evidence of an early hominin occupation dating back 1.66
million years ago (Zhu et al. 2004). There is evidence of stone tool processing of animal tissues in
the form of stone tools and modified animal bones. Both of these sites demonstrate a very early
hominin presence in China.

Homo erectus in Western Europe

There is debate about whether early hominin fossils recovered at sites in western Europe
should be identified as Homo erectus. The Atapuerca region of Spain is an area with many
limestone caves that have yielded the earliest hominin fossils from western Europe.
Deeply buried layers of deposits were first exposed in the 1890s when trenches were
cut through the hills for the construction of the railroad. Archaeologists have been
excavating at Atapuerca for decades and have discovered many remarkable sites
including Sima del elefante and Gran Dolina. There is clear evidence at Sima del elefante
that hominins had migrated to western Europe by 1.2 million years ago. At this site, a
hominin jaw, stone tools, and butchered animal remains were excavated. This hominin
fossil was identified as Homo antecessor. Scientists debate the taxonomic classification of
these hominin fossils. The excavators suggest that Homo antecessor could represent an
early population expansion out of Africa and the formation of a distinct species that
occurred in Eurasia leading to the Homo antecessor populations discovered at Sima del
elefante and Gran Dolina (Carbonell et al. 2008, Bermúdez de Castro 2017). It should be
noted, however, that this is not universally accepted, and some (including textbook
author Kenneth Feder) consider Homo antecessor a western European variant of Homo
erectus. Homo antecessor exhibits some ancestral characteristics that are similar to Homo
erectus while others link it more closely to modern forms of the genus Homo.
Gran Dolina, Spain is an 800,000-year-old site that is remarkable both for its age and for
the unusual assemblage that was recovered. Like Sima del elefante, the hominin fossils at
Gran Dolina are classified as Homo antecessor by the excavators. Interestingly, skeletal
analysis revealed evidence of cannibalism. Hominin carcasses at Gran Dolina were
butchered to obtain meat and marrow (Carbonell et al. 2010). This is the earliest
evidence of cannibalism in the archaeological record. The treatment and disposal pattern
of the butchered hominins indicate that cannibalism was part of the subsistence strategy
at this site. The hominin and animal bones have similar cut marks and breakage patterns,
and both human and animal remains seem to represent food waste that was disposed of
in the same manner. Regardless of whether the earliest hominin fossils are identified
as Homo erectus or Homo antecessor, there is clear evidence of an early hominin
occupation in western Europe by 1.2 million years ago.
At this site, a hominin jaw, stone tools, and butchered animal remains were excavated.
This hominin fossil was identified as Homo antecessor. Scientists debate the taxonomic
classification of these hominin fossils. The excavators suggest that Homo
antecessor could represent an early population expansion out of Africa and the formation
of a distinct species that occurred in Eurasia leading to the Homo antecessor populations
discovered at Sima del elefante and Gran Dolina (Carbonell et al. 2008, Bermúdez de
Castro 2017). It should be noted, however, that this is not universally accepted, and
some (including textbook author Kenneth Feder) consider Homo antecessor a western
European variant of Homo erectus. Homo antecessor exhibits some ancestral
characteristics that are similar to Homo erectus while others link it more closely to
modern forms of the genus Homo.
Gran Dolina, Spain is an 800,000-year-old site that is remarkable both for its age and for
the unusual assemblage that was recovered. Like Sima del elefante, the hominin fossils at
Gran Dolina are classified as Homo antecessor by the excavators. Interestingly, skeletal
analysis revealed evidence of cannibalism. Hominin carcasses at Gran Dolina were
butchered to obtain meat and marrow (Carbonell et al. 2010). This is the earliest
evidence of cannibalism in the archaeological record. The treatment and disposal pattern
of the butchered hominins indicate that cannibalism was part of the subsistence strategy
at this site. The hominin and animal bones have similar cut marks and breakage patterns,
and both human and animal remains seem to represent food waste that was disposed of
in the same manner. Regardless of whether the earliest hominin fossils are identified
as Homo erectus or Homo antecessor, there is clear evidence of an early hominin
occupation in western Europe by 1.2 million years ago.
Video 2

the earliest human in Europe

archaeological site of uh turf worker
research at this archaeological site
Ataturk burka in central Spain has
revealed extraordinary information about
European prehistoric settlement
excavation work has been conducted here
for the past thirty years this is the
leader of the team
archaeologists you dulled carbonelle
even before large-scale excavations
began he had a strong intuition that
there would be important finds to be
made here at Atapuerca a team of
archaeology students helped him with the
excavation work today
they are mostly volunteers the
excavation work only takes place between
June and September anyone digging here
is almost short to make an interesting
find many famous items have been already
discovered here
in 1992 the skull of a human ancestor
half a million years old was unearthed
here it was widely believed that
Neanderthals were the only human species
linked to modern man to evolve on the
European continent
however the skull turned out to be from
an earlier period and probably belonged
to a different type of human then a
jawbone dating from 800,000 years ago
was found these discoveries confirmed
that there were indeed other human
species apart from Neanderthals in
Europe the unknown species of human
which existed before Neanderthal was
named Homo antecessor a day in March
2008 became a day to remember
food alt carbonyl a human jaw bone from
1.2 million years ago was found here in
at apakah the find was reported around
the world with a headline that read the
first European now archaeologists are
looking forward to developing the
knowledge of how early humans came lived
and settled in Western Europe
Homo erectus Technology
The Acheulean Tool Tradition
Some early Homo erectus sites in Africa, and sites outside Africa such as Dmanisi, contain
Oldowan tools, but the emergence of Homo erectus coincides with the development of a
new stone tool technology called Acheulean. The Acheulean tool tradition is named after
the site of Saint-Acheul, France where the tools were first identified and described. The
earliest Acheulean tools in Africa date to 1.76 million years ago from the Kokiselei 4 Site,
West Turkana, Kenya. This tool technology spread to Europe and parts of Asia by about
one million years ago (Lepre et al. 2011). This Acheulean tool tradition persisted for
about 1.5 million years and in some areas, until about 160,000 years ago (Schick and
Toth 2013).
The characteristic Acheulean tool is a bifacial, leaf or teardrop shaped handaxe that has flakes
removed from both sides (see Figure 5-9). Experiments conducted by Schick and Toth (1993)
determine that Acheulean handaxes and cleavers were used primarily for butchering animal
carcasses. Microwear analysis shows a pattern consistent with animal butchering, and this is
interpreted as representing a shift towards more habitual and systematic butchering (Schick and
Toth 1993).

Acheulean handaxes are recovered from many sites in Africa and Europe. Sites in east
Asia, however, rarely contain handaxes. There are rare exceptions such as 800,000-year-
old handaxes from the Bose site in China (Yamei et al. 2000), but most of the tools that
are found at Homo erectus sites in east Asia are like the Oldowan tools used by Homo
habilis. At Zhoukoudian, there are no handaxes, and in Java, there are very few stone
tools at all (Rightmire 1990). The absence of stone tools in Indonesia suggests that Homo
erectus tools were manufactured from perishable materials such as wood or bamboo
(Schick and Toth 1993). Archaeologists consider this a likely explanation for the rarity of
handaxes in parts of south and east Asia.
Controlled Use of Fire
Precisely when hominins first harnessed fire is difficult to pinpoint, but the earliest
convincing evidence for the controlled use of fire dates to the time of Homo
erectus. There have been a number of claims for sites with evidence of the controlled use
of fire by early humans. Not all claims are widely accepted because scientists must be
able to rule out natural causes of fire. Some of the most convincing evidence for the
controlled use of fire comes from the Wonderwerk cave site in South Africa. Burned
bone and plant materials were found within discrete concentrations in Acheulean levels
dating to one million years ago (Berna et al. 2012). Analysis of the burned plant material
reveals the fuel source was leaves and grasses. Microscopic sediment analysis reveals
that this site contains in situ burning of material within the cave in association with
hominin occupation. There have been earlier claims for the controlled use of fire by
hominins, but these can be difficult to confirm because often natural causes of
accumulation of ash and burned material cannot be ruled out. Wonderwerk cave
provides unambiguous evidence of intentional use of fire (Berna et al. 2012). Another
site with evidence of intentional use of fire is Gesher Benot Ya’aqov, Israel (Goren-Inbar
et al. 2004). At this Acheulean site, dating to 790,000 years ago, archaeologists
uncovered burned seeds, burned wood, and burned flint. These materials were found in
distinct concentrations within the site. They are interpreted as hearths and a clear
representation of hominin activity. The archaeologists were able to rule out natural
causes as the source of burned material based on distinct concentrations of burned
materials (Goren-Inbar et al. 2004). In the case of natural fires, one would expect to find
high concentrations of burned material widely dispersed throughout the site. Regardless
of when hominins first used fire, it would have been a technology that would have
significant consequences for early hominins.

Why Fire Makes Us Human

Cooking may be more than just a part of your daily routine,
it may be what made your brain as powerful as it is
Wherever humans have gone in the world, they have carried with them two
things, language and fire. As they traveled through tropical forests they
hoarded the precious embers of old fires and sheltered them from
downpours. When they settled the barren Arctic, they took with them the
memory of fire, and recreated it in stoneware vessels filled with animal fat.
Darwin himself considered these the two most significant achievements of
humanity. It is, of course, impossible to imagine a human society that does
not have language, but—given the right climate and an adequacy of raw wild
food—could there be a primitive tribe that survives without cooking? In fact,
no such people have ever been found. Nor will they be, according to a
provocative theory by Harvard biologist Richard Wrangham, who believes
that fire is needed to fuel the organ that makes possible all the other
products of culture, language included: the human brain.

Every animal on earth is constrained by its energy budget; the calories

obtained from food will stretch only so far. And for most human beings, most
of the time, these calories are burned not at the gym, but invisibly, in
powering the heart, the digestive system and especially the brain, in the
silent work of moving molecules around within and among its 100 billion
cells. A human body at rest devotes roughly one-fifth of its energy to the
brain, regardless of whether it is thinking anything useful, or even thinking at
all. Thus, the unprecedented increase in brain size that hominids embarked
on around 1.8 million years ago had to be paid for with added calories either
taken in or diverted from some other function in the body. Many
anthropologists think the key breakthrough was adding meat to the diet. But
Wrangham and his Harvard colleague Rachel Carmody think that’s only a
part of what was going on in evolution at the time. What matters, they say, is
not just how many calories you can put into your mouth, but what happens
to the food once it gets there. How much useful energy does it provide, after
subtracting the calories spent in chewing, swallowing and digesting? The real
breakthrough, they argue, was cooking.

Wrangham, who is in his mid-60s, with an unlined face and a modest

demeanor, has a fine pedigree as a primatologist, having studied
chimpanzees with Jane Goodall at Gombe Stream National Park. In pursuing
his research on primate nutrition he has sampled what wild monkeys and
chimpanzees eat, and he finds it, by and large, repellent. The fruit of the
Warburgia tree has a “hot taste” that “renders even a single fruit impossibly
unpleasant for humans to ingest,” he writes from bitter experience. “But
chimpanzees can eat a pile of these fruits and look eagerly for more.”
Although he avoids red meat ordinarily, he ate raw goat to prove a theory
that chimps combine meat with tree leaves in their mouths to facilitate
chewing and swallowing. The leaves, he found, provide traction for the teeth
on the slippery, rubbery surface of raw muscle.

Food is a subject on which most people have strong opinions, and

Wrangham mostly excuses himself from the moral, political and aesthetic
debates it provokes. Impeccably lean himself, he acknowledges blandly that
some people will gain weight on the same diet that leaves others thin. “Life
can be unfair,” he writes in his 2010 book Catching Fire, and his shrug is
almost palpable on the page. He takes no position on the philosophical
arguments for and against a raw-food diet, except to point out that it can be
quite dangerous for young children. For healthy adults, it’s “a terrific way to
lose weight.”

Which is, in a way, his point: Human beings evolved to eat cooked food. It is
literally possible to starve to death even while filling one’s stomach with raw
food. In the wild, people typically survive only a few months without cooking,
even if they can obtain meat. Wrangham cites evidence that urban raw-
foodists, despite year-round access to bananas, nuts and other high-quality
agricultural products, as well as juicers, blenders and dehydrators, are often
underweight. Of course, they may consider this desirable, but Wrangham
considers it alarming that in one study half the women were malnourished to
the point they stopped menstruating. They presumably are eating all they
want, and may even be consuming what appears to be an adequate number
of calories, based on standard USDA tables. There is growing evidence that
these overstate, sometimes to a considerable degree, the energy that the
body extracts from whole raw foods. Carmody explains that only a fraction of
the calories in raw starch and protein are absorbed by the body directly via
the small intestine. The remainder passes into the large bowel, where it is
broken down by that organ’s ravenous population of microbes, which
consume the lion’s share for themselves. Cooked food, by contrast, is mostly
digested by the time it enters the colon; for the same amount of calories
ingested, the body gets roughly 30 percent more energy from cooked oat,
wheat or potato starch as compared to raw, and as much as 78 percent from
the protein in an egg. In Carmody’s experiments, animals given cooked food
gain more weight than animals fed the same amount of raw food. And once
they’ve been fed on cooked food, mice, at least, seemed to prefer it.

In essence, cooking—including not only heat but also mechanical processes

such as chopping and grinding—outsources some of the body’s work of
digestion so that more energy is extracted from food and less expended in
processing it. Cooking breaks down collagen, the connective tissue in meat,
and softens the cell walls of plants to release their stores of starch and fat.
The calories to fuel the bigger brains of successive species of hominids came
at the expense of the energy-intensive tissue in the gut, which was shrinking
at the same time—you can actually see how the barrel-shaped trunk of the
apes morphed into the comparatively narrow-waisted Homo sapiens. Cooking
freed up time, as well; the great apes spend four to seven hours a day just
chewing, not an activity that prioritizes the intellect

The trade-off between the gut and the brain is the key insight of the
“expensive tissue hypothesis,” proposed by Leslie Aiello and Peter Wheeler in
1995. Wrangham credits this with inspiring his own thinking—except that
Aiello and Wheeler identified meat-eating as the driver of human evolution,
while Wrangham emphasizes cooking. “What could be more human,” he
asks, “than the use of fire?”

Unsurprisingly, Wrangham’s theory appeals to people in the food world. “I’m

persuaded by it,” says Michael Pollan, author of Cooked, whose opening
chapter is set in the sweltering, greasy cookhouse of a whole-hog barbecue
joint in North Carolina, which he sets in counterpoint to lunch with
Wrangham at the Harvard Faculty Club, where they each ate a salad. “Claude
Lévi-Strauss, Brillat-Savarin treated cooking as a metaphor for culture,”
Pollan muses, “but if Wrangham is right, it’s not a metaphor, it’s a
precondition.” (Read about what it's like to have dinner with Pollan)

Wrangham, with his hard-won experience in eating like a chimpanzee, tends

to assume that—with some exceptions such as fruit—cooked food tastes
better than raw. But is this an innate mammalian preference, or just a
human adaptation? Harold McGee, author of the definitive On Food and
Cooking, thinks there’s an inherent appeal in the taste of cooked food,
especially so-called Maillard compounds. These are the aromatic products of
the reaction of amino acids and carbohydrates in the presence of heat,
responsible for the tastes of coffee and bread and the tasty brown crust on a
roast. “When you cook food you make its chemical composition more
complex,” McGee says. “What’s the most complex natural, uncooked food?
Fruit, which is produced by plants specifically to appeal to animals. I used to
think it would be interesting to know if humans are the only animals that
prefer cooked food, and now we’re finding out it’s a very basic preference.”

Among Wrangham’s professional peers, his theory elicits skepticism, mainly

because it implies that fire was mastered around the time Homo
erectus appeared, roughly 1.8 million years ago. Until recently, the earliest
human hearths were dated to about 250,000 B.C.; last year, however, the
discovery of charred bone and primitive stone tools in a cave in South Africa
tentatively pushed the time back to roughly one million years ago, closer to
what Wrangham’s hypothesis demands but still short. He acknowledges that
this is a problem for his theory. But the number of sites dating from that
early period is small, and the evidence of fire might not have been preserved.
Future excavations, he hopes, will settle the issue.
In Wrangham’s view, fire did much more than put a nice brown crust on a
haunch of antelope. Fire detoxifies some foods that are poisonous when
eaten raw, and it kills parasites and bacteria. Again, this comes down to the
energy budget. Animals eat raw food without getting sick because their
digestive and immune systems have evolved the appropriate defenses.
Presumably the ancestors of Homo erectus—say, Australopithecus—did as
well. But anything the body does, even on a molecular level, takes energy; by
getting the same results from burning wood, human beings can put those
calories to better use in their brains. Fire, by keeping people warm at night,
made fur unnecessary, and without fur hominids could run farther and faster
after prey without overheating. Fire brought hominids out of the trees; by
frightening away nocturnal predators, it enabled Homo erectus to sleep safely
on the ground, which was part of the process by which bipedalism (and
perhaps mind-expanding dreaming) evolved. By bringing people together at
one place and time to eat, fire laid the groundwork for pair bonding and,
indeed, for human society.

We will now, in the spirit of impartiality, acknowledge all the ways in which
cooking is a terrible idea. The demand for firewood has denuded forests. As
Bee Wilson notes in her new book, Consider the Fork, the average open
cooking fire generates as much carbon dioxide as a car. Indoor smoke from
cooking causes breathing problems, and heterocyclic amines from grilling or
roasting meat are carcinogenic. Who knows how many people are burned or
scalded, or cut by cooking utensils, or die in cooking-related house fires? How
many valuable nutrients are washed down the sink along with the water in
which vegetables were boiled? Cooking has given the world junk food, 17-
course tasting menus at restaurants where you have to be a movie star to
get a reservation, and obnoxious, overbearing chefs berating their sous-chefs
on reality TV shows. Wouldn’t the world be a better place without all that?

Raw-food advocates are perfectly justified in eating what makes them feel
healthy or morally superior, but they make a category error when they
presume that what nourished Australopithecus should be good enough
for Homo sapiens. We are, of course, animals, but that doesn’t mean we have
to eat like one. In taming fire, we set off on our own evolutionary path, and
there is no turning back. We are the cooking animal.
Homo floresiensis
Homo floresiensis is a unique hominin species that was discovered in 2003 when
excavations at Liang Bua (see Figure 5-10) on the Indonesian island of Flores uncovered
the remains of a tiny, small brained hominin (Brown et al. 2004). A remarkably complete
skeleton and cranium (LB1) was the first to be discovered (See Figure 5-11). This
hominin is unlike any other. A unique combination of primitive and derived
characteristics led to its identification as a new species belonging to the genus Homo.
Recent dating of sediments at Liang Bua, indicate Homo floresiensis lived very recently
between 100,000 and 60,000 years ago (Sutikna et al. 2016).
The shape of the pelvis indicates that LB1 is a female. Her height is estimated at 106 cm and her
weight between 16 to 28.7 kg (Brown et al. 2004). Despite the small size, she is clearly an adult.
This is based on the presence of all the permanent teeth, the degree of tooth wear, and the
degree of epiphyseal fusion. LB1 has a cranial capacity of 417 cm3 which is similar to the size of
an australopithecine (Falk et al. 2005; see Figure 5-1), but although the size of the brain is similar
to an australopithecine, the shape of the endocast is more like Homo erectus. The overall shape of
the cranium and face is also similar to Homo erectus. The face is less prognathic than the
australopithecines, with smaller teeth and jaws (Brown et al. 2004). In 2004, additional
discoveries of Homo floresiensis were made at Liang Bua, all exhibiting similar characteristics to
LB1. A total of nine individuals with the same characteristics confirm that LB1 is not a
pathological individual. She is one of many hominins identified as Homo floresiensis that share the
same physical characteristics (Morwood et al. 2005). Another remarkable discovery at Liang Bua
is the discovery of Oldowan-like stone tools in the same sediments as the Homo
floresiensis remains. This indicates that a large brain size is not a requirement for the production
of core and flake tools (Morwood and Junger 2009). Other significant finds include evidence of
the use of fire, butchered animal bone, and fire cracked rock (Morwood et al. 2005). It appears
that Homo floresiensis hunted and butchered animals, including dwarf elephants. These are the
types of activities that are usually associated with large brained hominins.

An explanation for the small size of the body and brain of Homo floresiensis is insular
dwarfing (Brown et al. 2004). It is possible that the small size of Homo floresiensis was selected
for in an island habitat with limited resources. In such an environment, a small body size with
lower energy requirements is advantageous. Scientists suggest that the ancestors of Homo
floresiensis may have been larger than Homo erectus, and that it became smaller in a habitat with
limited resources (Brown et al. 2004). Not everyone agrees with this hypothesis. When Homo
floresiensis was discovered, some suggested that it did not represent a new form of hominin but
was instead a modern human with microcephaly, which is a pathological condition affecting
growth and brain development (Jacob et al. 2006, Martin et al. 2006). However, comparison
of Homo floresiensis to modern human skeletal materials with a variety of conditions that affect
stature and brain growth indicate no similarity of Homo floresiensis to modern humans with
pathological conditions (Brown et al. 2004, Falk et al. 2005, Argue et al. 2017).

In 2015, additional Homo floresiensis fossils were identified at a second site in Flores called Mata
Menge. Like the younger Homo floresiensis discovery at Liang Bua, the Mata Menge site had
hominin fossils alongside stone tools similar to those used by later Homo floresiensis (van den
Bergh et al. 2016). The Mata Menge site, however, is much older dating to 700,000 years ago
(Brumm et al. 2016). The finds at Mata Menge are derived compared to both australopithecines
and Homo habilis and support the view that Homo floresiensis is a dwarfed descendent of
Asian Homo erectus (van den Burgh et al. 2016). In their discussion of the Mata Menge hominins
and the evolutionary relationship of Homo floresiensis, van den Bergh and colleagues suggest that
the physical characteristics of the Mata Menge fossils are consistent with the hypothesis
that Homo floresiensis originated from a population with closest affinities to Homo erectus from
Java (van den Bergh et al. 2016). The Homo floresiensis fossils found in 700,000-year-old
sediments seem to suggest that small body size was present in Flores hominins at a very early
date (van den Bergh et al. 2016).

Homo floresiensis: Making Sense of the Small-Bodied Hominin

Fossils from Flores
Are the bones of several tiny individuals from the island of Flores the newest
addition to our family tree, or are they the remains of diseased humans only
masquerading as an extinct species?

It was 2004 when a team of joint Australian-Indonesian researchers announced the

discovery of fossils and simple (Oldowan-like) stone tools from Liang Bua cave (which
translates to "cool cave") on the remote Indonesian island of Flores. The fossils
represent a small-bodied and small-brained hominin, named Homo floresiensis, but
better known as the "Hobbit." The position of these fossils on the human evolutionary
tree remains unclear. In fact, since the 2004 discovery, there has been an unending
series of controversies surrounding these specimens.
What is perhaps most surprising is the young geological age of these fossils. The fossils
range in age from 74 to 17 ka (Brown et al., 2004), which falls well within the range of
modern humans elsewhere in the Old and quite possibly New World. However, only the
younger layers above the Homo floresiensis fossils show evidence of modern
human occupation at Liang Bua.
Fossil and Archaeological Evidence
The LB1 skeleton (type specimen of the new species) preserves a nearly complete
skull, a partial pelvis, several limb bones, and bones of the hands and feet. This
individual is estimated to have stood about 1.06 m (3'6") tall, which is roughly the size of
a 3–4 year old modern human child. To date, LB1 has the only complete skull to be
recovered, but a second mandible (lower jaw) and numerous skeletal elements from a
second individual (LB6) have been described (Brown et al., 2004). Fossils of at least
four other individuals have also been recovered, confirming that this was a population of
small-bodied individuals and that LB1 was not an anomaly.
Initial descriptions emphasized the mix of ancestral traits that remained unchanged
from more ancient species and derived traits that linked it with more recent ones. The
skull resembles those belonging to extinct species of our own genus Homo (Brown et
al., 2004; Baab and McNulty, 2009). The skeleton, however, is considerably more
primitive and in some respects aligns the LB1 specimen and the other Flores fossils
with older and even more primitive species like those belonging to Australopithecus
afarensis (which includes "Lucy"; Tocheri et al., 2007; Jungers et al., 2009) (link to
Ward and Hammond article; link to Schrein article). Taken together, this is a puzzling
pattern: a population that existed between 74 and 18 ka, with a skull that most closely
resembles the much older Homo habilis or Homo erectus, and a skeleton that retains
some features normally associated with australopith species at least 3 Ma.
Presumably Homo floresiensis made the simple Oldowan-like tools (the oldest and most
primitive type known in the archaeological record) found in the same layers as the
skeletal remains at Liang Bua cave (Moore and Brumm, 2009). These stone tools
resemble those found elsewhere on the island at sites that are closer to a million years
in age (Brumm et al., 2006; 2010). In addition to tools, there is also evidence in the form
of cut marks on some Stegodon bones indicating that the hominins were butchering
these animals (Morwood et al., 2005). There are also some burnt bones and pebbles,
but whether this was the result of intentional or accidental fire is still unknown
(Westaway et al., 2009)
Evolutionary Scenarios
As a result of the primitive morphology but surprisingly young geological age of the
fossils, many scenarios have been proposed to explain the presence of these fossils on
Flores. Homo erectus occupied Southeast Asia from about 1.5 million years to perhaps
as recently as half a million years ago or even 50 thousand years ago (Larick et al.,
2001; Huffman et al., 2010; Indriati et al., 2011). The low neurocranium (the part
surrounding the brain) with a flat and sloping forehead, thick cranial bones, short and
flat face, and other details of LB1's skull anatomy (e.g., an occipital torus and
a mastoid fissure), as well as the shape of the brain provide a link to Homo erectus,
but both the small body size and brain size are outside of the expected range for that
species (Brown et al., 2004; Falk et al., 2005; Baab and McNulty, 2009). Therefore,
based primarily on the cranial evidence, the original description proposed that these
fossils represented a new species, Homo floresiensis, that was a dwarfed descendent
of Homo erectus. Additional aspects of anatomy also resemble that known for Homo
erectus, including the shape of the brain and the shoulder (Falk et al., 2005; Larson et
al., 2009). According to this theory, the ancestors of Homo floresiensis somehow made
the treacherous water crossing to reach Flores and over time there was a reduction in
body size (Meijer et al., 2010).
The phenomenon of island dwarfing has been documented for a number of other
large-bodied mammals, including primates, mammoths, and deer. There are also
numerous instances of human populations becoming dwarfed, including on the islands
of Southeast Asia. The impetus for island dwarfing is often linked to reduced availability
of resources in the environment. In either case, being small-bodied may be more
advantageous on an island than on the mainland. Flores exists in a region known
as Wallacea where there is a low level of migration from the Asian and Australasian
faunas (collections of animals) to the east and west, respectively. These islands appear
to have been isolated by strong currents, even when sea levels were low. Along with the
hominins, a few other large-bodied mammals have been documented in this region,
including Stegodon, a type of extinct type of elephant, and komodo dragons. Both of
these species were found in deposits coeval with Homo floresiensis, however, their
rarity is further confirmation that it was difficult for large terrestrial animals to reach
islands in Wallacea. Interestingly, the particular subspecies of Stegodon found in the
same layers of Liang Bua cave as Homo floresiensis, Stegodon florensis insularis, is
itself a dwarfed form whose closest relative is the distinctly larger and geologically
older Stegodon florensis florensis (van den Bergh et al., 2008).
An argument against island dwarfing as an explanation for Homo floresiensis stems
from the relationship between brain size and body size. LB1's endocranial volume,
calculated from CT scans of the skull, is only 417 cm3, close to a third of the average
modern human value (Falk et al., 2005). This small brain size is difficult to explain
based on typical patterns of island dwarfing assuming that the ancestor resembled a
modern human or even Homo erectus (Martin et al., 2006). This is because brain size
typically "dwarfs" less than overall body size. For example, despite having bodies that
are much smaller than their neighbors, modern human pygmies have brains which are
only slightly smaller. However, more drastic decreases in brain size compared to body
size during island dwarfing are not entirely without precedent and may have occurred in
at least two other groups: Myotragus ("cave goat") on Mallorca and Hippopotamus
lemerlei on Madagascar (Köhler and Moyà-Solà, 2004; Weston and Lister, 2009).
A second model advanced to explain the presence of the hominin fossils on the island
of Flores in the Pleistocene states that this population was the offshoot of a more
primitive, pre-erectus hominin species with a small body size and small brain. Evidence
from the mandible and the rest of the skeleton supports this hypothesis (Argue et al.,
2009). The size and morphology of the teeth and mandible share more resemblances
to Australopithecus and the earliest Homo species than to Homo erectus (Brown and
Maeda, 2009). In particular, the very short legs (relative both to the arms and to the
feet) are a pattern seen in apes and australopiths rather than Homo erectus (a
good Homo habilis skeletal comparison has not yet been discovered). LB1 was also
disproportionately heavy for her height — a pattern closely approximated by the famous
3.2 million year old Australopithecus afarensis skeleton of "Lucy" (Jungers and Baab,
2009). Despite being only 106 cm in height, LB1 is estimated to have weighed close to
32.5 kg (71.7 lbs.). The carpal bones (bones of the wrist) in Homo floresiensis look
more like those of chimpanzees than of modern humans (Tocheri et al., 2007). While
the evolution of the wrist is not well documented in early members of the
genus Homo (e.g., Homo habilis and Homo erectus), it is clear that the wrist
morphology is more primitive than that of modern humans and Neanderthals. Like the
wrist, the foot morphology, although exhibiting some human-like traits (including a non-
grasping big toe), also retains several quite primitive features: a long forefoot with
curved toes and the lack of a medial longitudinal arch (Jungers et al., 2009). The
latter trait is indicated by the particular shape of the navicular bone, one of the bones
close to the heel. This medial longitudinal arch is located on the inside aspect of the
foot, forming the instep, and acts both as a shock absorber and a spring during walking.
It seems likely that the relatively long foot and particular combination of traits present in
the foot would have resulted in a slightly different walking gait than seen in modern
humans. This combination of traits in the skeleton appears to place the ancestry of
the Homo floresiensis skeleton earlier than Homo erectus.
One major challenge to the idea that Homo floresiensis has an ancestry deeper
than Homo erectus is the absence of fossils of any such species in either island or
mainland Southeast Asia. The hominin fossil record prior to Homo erectus is found only
in Africa. A second complication is that the fossil record of postcranial anatomy for pre-
erectus species of Homo is poor and their morphology is not as well documented as
other species, so comparison with Homo floresiensis is limited. As a result, it is not clear
whether a species such as Homo habilis, which precedes Homo
erectus geochronologically, is in fact a good model for the ancestor of Homo
floresiensis.
Pathological Hypotheses
In contrast to the two evolutionary hypotheses, a third theory has been advanced: the
Flores remains represent modern humans suffering from some type of pathological
condition (a disease). The initial suggestion was that LB1 was a modern human that
suffered from microcephaly, a condition in which the neurocranium is considerably
smaller than that of normal, healthy people (Henneberg and Thorne, 2004; Martin et al.,
2006; Jacob et al., 2006). This can be the result of a genetic disorder whose main
symptoms are under-development of the brain and the overlying cranial vault — a
condition known as primary microcephaly. However, microcephaly is a symptom of a
range of other disorders, some of which include short stature. The proponents of the
pathology hypothesis have thus far failed to identify exactly what disorder can account
for the large number of apparently primitive traits in the LB1 skeleton. There is a clear
similarity in the shape of the skull between archaic species of Homo and humans with
microcephaly — primarily in the high ratio of facial to neurocranial size. This
convergence in shape occurs for two very different reasons: microcephalic humans
have pathologically underdeveloped brains whereas early hominins belonged to species
normally characterized by smaller brains than those of modern humans. These
similarities are only superficial, however, and more detailed examination of the cranial
vault shape, as well as the shape of the underlying brain, shows that LB1 shares
important characteristics with fossil Homo species (Falk et al., 2005, 2007; Baab, 2010).
In addition to microcephaly, two other very different pathologies have been implicated:
myxoedematous endemic hypothyroidism ("cretinism"; Obendorf et al., 2007; Oxnard et
al., 2010) and Laron Syndrome (Hershkovitz et al., 2007). The former is caused by a
lack of iodine in the diet during both pre- and post-natal development (Halpern et al.,
1991); iodine is crucial for the proper production of hormones in the thyroid gland and
for the development of the central nervous system. The most notable symptoms of this
disease include mental retardation and other neurological defects as well as short
stature due to delayed skeletal maturity (Boyages et al, 1988). The skull is often
described as brachycephalic, and the sutures between individual bones of the skull
frequently remain open even once brain growth has ended (Koplik, 1918, p. 44).
However, the LB1 skull is not brachycephalic, and many of its cranial sutures are fully
fused. In fact, the skulls of cretins are not particularly small, and their shape more
closely resembles healthy humans than LB1 (Baab, 2010). Some skulls of patients
with congenital hypothyroidism exhibit an enlarged pituitary fossa. This feature was
initially attributed to the skull of LB1 (Obendorf et al., 2007), but this has been refuted by
measurements from CT scans of the skull, as have a number of other alleged
similarities between LB1 and humans with myxoedematous hypothyroidism (Jungers et
al., 2009). While the skeletons of both human cretins and Homo floresiensis differ from
that of healthy modern humans, they do not resemble one another. For example, the
limb proportions of LB1 are not comparable to those of humans with cretinism
(Jungers et al., 2009; Groves and Fitzgerald, 2010).
Laron Syndrome is characterized by insensitivity to growth hormones; although the
body manufactures the correct growth hormones, the receptors in the body do not
respond properly to them. As in myexedemotous hypothyroidism, patients suffering from
Laron Syndrome are of shorter than average stature. Although the original paper
proposing that LB1 had Laron Syndrome found a number of similarities between the
LB1 skeleton and individuals with Laron Syndrome (Hershkovitz et al., 2007), it is now
clear that the case was overstated (Falk et al., 2009), and that the evidence for this
particular syndrome in LB1 is nonexistent. For example, with regards to the skull,
patients with Laron Syndrome typically have a protruding forehead, underdeveloped
facial bones such that the face looks small compared to the rest of the head, and a skull
that is disproportionately wide across the parietal bones compared to the base of the
skull. The LB1 skull shows the opposite pattern: the forehead slants backwards rather
than protruding, the face is large relative to the rest of the skull, and it is wide at the
base rather than the parietals (Falk et al., 2009; Baab, 2010).
Conclusions
The rather unusual combination of short stature, a small brain, primitive skeletal
anatomy, simple stone tools, all occurring at a time when fully modern humans
inhabited other parts of the globe has led to a wide range of explanations (summarized
above and in more detail in Aiello [2010]). Consensus among researchers regarding the
interpretation of the Liang Bua fossils remains elusive as the study of the fossils is
ongoing and apparently contradictory interpretations have been published. How can we
make sense of this? Both cretinism and Laron Syndrome result in humans of small body
size, which is consistent with LB1. However, the other symptoms of these diseases are
not observed in the LB1 skeleton. Furthermore, LB1 was not an isolated find, and the
other individuals found in Liang Bua cave are of a similarly diminutive size. As it is
highly improbable that all members of this population were diseased, short stature
should be viewed as a characteristic of the population rather than a pathological
symptom of a single individual. Microcephaly is a more difficult argument to dismiss
because it is not a disease per se but rather a symptom that accompanies many
disorders. However, the shape of the LB1 neurocranium does not resemble skulls of
human microcephalics thus far examined (Baab, 2010) and a single disorder that
explains the apparently primitive features found elsewhere in the skeleton has yet to be
described. Taken together, the weight of evidence does not support a pathological
explanation for the particular characteristics found in LB1 and her kin in Liang Bua cave.
Rather, the various lines of evidence, from the simple stone tools, the Homo-like skull
morphology, and the even more primitive skeletal anatomy, paint a more complicated
evolutionary picture. The two most popular evolutionary hypotheses position Homo
floresiensis as the dwarfed descendant of Homo erectus or descendant of an even
more primitive species. If the former is correct, then Homo floresiensis certain skeletal
traits reappeared in this lineage that were seen in earlier australopith species but lost
prior to or at the origin of Homo erectus. If the latter is true, then Homo floresiensis was
descended from a species such as Homo habilis for which there is no evidence
elsewhere in Asia. Only additional fossils or analyses will determine the evolutionary
history of the "Hobbit" of Flores island.
Figure 5: The LB1 cranium (center) compared to Homo habilis from Kenya (1.9
Ma) (left) and the slightly younger Homo erectus cranium from the Republic of
Georgia (1.8 Ma) (right).
Previous investigations of the shape of the LB1 neurocranium indicate close similarities
to this Georgian Homo erectus individual.
Glossary
ancestral trait: Characteristic shared between two or more groups inherited from an
ancestor prior to their most recent common ancestor
Australopithecus: An extinct hominin genus found only in Africa that preceded the
genus Homo chronologically and contains several species spanning the time interval of
4.4 to 2.0 Ma
Australopithecus afarensis: An extinct hominin species in the
genus Australopithecus from East Africa in the time period 3.9 to 3 Ma; the most famous
representative of this species is the well-known fossil skeleton named "Lucy" from
Ethiopia
brachycephalic: Term referring to skulls that are relatively wide from left to right
compared to their length from front to back
cranial: Refers to the cranium (the bony skull minus the lower jaw)
CT: Acronym for computed tomography which is an X-ray-based imaging technique
used to see both the external and internal anatomy of structures such as skulls and
teeth (also known as CAT scanning in a clinical context)
derived trait: A characteristic shared between two or more groups that was inherited
from their most recent common ancestor; this trait indicates a close evolutionary
relationship among those groups that share the trait
hominin: Any member of the tribe Hominini, the group that includes modern humans
and those fossil species that are more closely related to humans than any other animal
Homo: An extinct hominin genus that includes our own species (Homo sapiens) among
others. Members of this genus first appear in the fossil record about 2.3 Ma in Africa.
The Liang Bua fossils have been assigned to this genus.
Homo erectus: An extinct species of hominin that is in the same genus as modern
humans (Homo) that was found in Africa, Eurasia and east Asia from 1.8 Ma to 0.5 or
0.05 Ma
Homo habilis: An extinct species of hominin that is in the same genus as modern
humans (Homo) that was found in Africa from 2.3–1.4 Ma
island dwarfing: Phenomenon where large-bodied mammals may become dwarfed on
after immigrating to an island
ka: Acronym referring to kiloannum (thousand years); used to express ages in
thousands of years (e.g., a skull that is 20 ka is 20,000 years old)
lineage: A restricted segment of the phylogenetic tree that includes an ancestor and
those species that it gave rise to, either directly or indirectly (can also refer to only a
single species)
Ma: Acronym referring to megaannum (million years); used to express ages in millions
of years (e.g., a skull that is 1.2 Ma is 1,200,000 years old)
mastoid fissure: A groove located on the underside of the skull between the mastoid
process and the petrosal crest of the tympanic bone (common in skulls of Homo
erectus)
morphology: Pertaining to the form or appearance of an organism or a particular
anatomical structure
occipital torus: Elevated mound of bone that crosses the occipital bone located at the
back of the skull (common in skulls of Homo erectus)
Oldowan: Oldest known stone tool technology (most common in Africa 1.6–1.6 Ma);
flakes are produced by knocking them off of a core using a hammerstone
parietal bones: The large and fairly flat bones that surround the brain on the right and
left sides; they are joined by the sagittal suture which traverses the midline of the skull
on its top surface
pituitary fossa: A depression inside of the skull (on the sphenoid bone) where the
pituitary gland (hypophysis) sits during life
Pleistocene: An epoch (a unit of geological time) that spans 1.8 Ma through 11.7 ka
sutures: A joint between bones of the skull; these are generally open during growth of
the skull and begin to fuse later in life, after growth has ended
type specimen: The specimen for which a particular species is named for; intended to
typify the characteristics of that species
Wallacea: Geographic region in Southeast Asia delimited by two imaginary lines:
Wallace's Line to the west and Lydekker's line to the east. The region between these
lines contains animals that are distinct from those in Asia (to the west) or Australia (to
the east). See Figure 1.

Homo naledi
Homo naledi is a recent addition to the genus Homo that was discovered in South Africa.
It is unique from Homo erectus and demonstrates the diversity of hominins assigned to
the genus Homo. Homo naledi was discovered in 2013 in the Dinaledi Chamber within
the Rising Star Cave System in South Africa. Excavations in 2013 and 2014 yielded the
remains of over 1,500 fossils representing at least 15 individuals including children and
adults (Figure 5-12). This hominin has a mix of australopithecine-like and Homo-like
characteristics, as well as some features that are not seen in any other hominin species
(Berger et al. 2015). The fossil assemblage includes most skeletal elements and four
skulls with a cranial capacity between 465 cm and 560 cm (see Figure 5-1). The crania
3 3

are small in size but similar in structure to the genus Homo. The teeth and mandibles are
smaller than those of australopithecines (Berger et al. 2015).
When the fossils were first described, a date had not yet been assigned to the hominins
within the Dinaledi Chamber. It was suggested, based on physical characteristics,
that Homo naledi could potentially be older than two million years old (Berger et al.
2015). However, dates recently obtained using a variety of dating methods reveal the
fossils are between 236,000 to 335,000 years old (Dirks et al. 2017). This is much
younger than was expected based on their physical characteristics. These new findings
demonstrate the difficulty of trying to predict the age of a fossil based only on its
appearance and emphasize the importance of dating specimens using independent tests
(Dirks et al. 2017).
The deposition of the fossils deep within a cave is unusual (see Figure 5-13), and it has
been suggested that the individuals may have been deliberately placed in the cave after
death (Dirks et al. 2015). Perhaps this is evidence of intentional disposal of the bodies.
The remains do not show evidence of carnivore chewing, and the skeletal remains in the
cave are almost exclusively hominin. This seems to rule out carnivore predation as the
reason for the accumulation of these materials. There are no signs of water flow washing
them into the cave system, and there is not even any evidence of a direct vertical
passage from the surface into the Dinaledi chamber (Dirks et al. 2015).
Further exploration of the Rising Star cave system has turned up remains of additional Homo
naledi fossils from another chamber called the Lesedi Chamber (Hawks et al. 2017). In this
chamber, 30 metres beneath the ground surface, three additional individuals (including two
adults and one child) were found in a separate, undated chamber. The physical characteristics of
the hominins are like those from the Dinaledi Chamber and are also assigned to Homo naledi. Like
those from the Dinaledi chamber, the Lesedi chamber hominins appeared to have been intact
when deposited in the chamber (Hawks et al. 2017). Paleoanthropologists will undoubtedly
continue to learn more about Homo naledi as new fossils are discovered and analyse
Video 4

we do have our genus

what these discoveries telling us is
that there's a lot out there to be found
that we actually don't have the whole
story of human evolution i mean it looks
like it might be a fragment of like the
super orbital taurus or something oh my
god
the
homo naledi fossils are extraordinary in
a wide variety of ways we know they have
the characters of an early member of the
genus homo
by comparing their morphology to
other early hominids we can see that we
would place them right at the base of
the lineage that leads to
us
we've never seen
a non-human
that shares so many primitive and yet
sometimes advanced characters tiny brain
curved fingers but a generally
human-like hand long legs and a human
foot
you're looking at well over a dozen in
fact probably around 18 different
individuals representing all different
age spans from near fetal age to senile
individuals who were at the last stages
of their lives
when
i first
saw these images by my exploration team
i knew that we had to act the rising
star cave system is one of the best
known in south africa it's been
caved continuously for more than 50
years
i had this fossil lying on the surface
and i could see that it had been damaged
so i called national geographic to put
together an expedition fast
the entrance is very very difficult
to get to it you have to move through a
seven and a half inch slot
wiggling your way across sharp rocks
before you drop into
this remarkable
little chamber
and the floor is quite
literally comprised of parts of the
bodies of these human ancestors
look at this
tell them they're a go to
all collect
you got the fossil fuel
it appears based upon the context
that we have discovered this incredible
assemblage
of hominin fossils in
is that they were deliberately placed
there by their next of kin after death
homo naledi was doing something that
until this moment we thought
was unique to modern humans that is
deliberate disposal of the dead
if this hypothesis holds true
that's an extraordinary thing
what Naledi has taught us
is that there is clearly more out there
that we didn't know
you
Homo erectus is a highly variable species spanning a broad temporal and geographic
range. It is the first hominin to migrate beyond the African continent and did so shortly
after its initial appearance in Africa almost 1.9 million years ago. With its use of
technology and larger brain and body, Homo erectus was able to adapt to a wide variety
of habitats. Homo erectus developed a more sophisticated stone tool technology than
earlier hominins. The Acheulean tool industry likely allowed Homo erectus to utilize a
wider variety of resources than earlier hominins, particularly animal protein. There is
ample evidence of butchering at many Homo erectus sites, and several later sites reveal
evidence for the controlled use of fire. Perhaps it is these behavioural changes that
allowed Homo erectus to successfully adapt to the diverse environments of Africa,
Eurasia, and eventually Western Europe. Paleoanthropology is an ever-changing field of
study, and recent excavations of unusually small-brained hominins surviving late into
the Pleistocene reveal the diversity of hominin species that existed. Discoveries
like Homo naledi and Homo floresiensis demonstrate the presence of seemingly isolated
populations that exhibit unique suites of ancestral and derived physical characteristics,
even at a time when anatomically modern humans were emerging elsewhere.

Learning Activity 5-9

Add to the summary table of key archaeological sites that you worked on in a previous
module by adding more information about the following sites:

▪ Nariokotome, West Turkana, Kenya

▪ Dmanisi, Georgia
▪ Trinil, Java, Indonesia
▪ Zhoukoudian, China
▪ Atapuerca, Spain (Sima del elefante, Gran Dolina)
▪ Kokiselei 4, West Turkana, Kenya
▪ Wonderwerk Cave, South Africa
▪ Gesher Benot Ya’aqov, Israel
▪ Flores, Indonesia (Liang Bua, Mata Menge)
▪ Rising Star Cave System, South Africa (Dinaledi Chamber, Lesedi Chamber)
Using the information presented in this module, create a summary table describing the
following for each site:
 1. The age of the site;
2. what fossils and/or artifacts were found at the site; and
3. the significance of these discoveries.
Keep your completed summary table and use it as a study tool for this module.
Book
IN 1918 A FOSSIL LOCALITY SOUTHWEST OF Beijing, China, was explored by Swedish geologist
Johan Gunnar Andersson. Andersson took advantage of a local Chinese belief that fossil bones
were actually the remnants of dragons and that powder made from ground-up “dragon bones” was
a cure-all. Many local druggists collected such bones for use in their medicines. Even today,
paleontologists rely on local druggists for leads in their search for fossil (“dragon”) bones (Jian and
Rice 1990).

In 1918 Andersson was directed by a druggist to a hill called Jigushan (Chicken Bone Hill) near the
village of Zhoukoudian (Jia and Huang 1990). Convinced there was a rich array of fossil bones in the
surrounding region, he began excavating on another hill, called Longgoshan—Dragon Bone Hill. In
1926 two humanlike teeth were recovered; in 1929, with the dig now led by British scientist
Davidson Black, a nearly intact skull was encountered in a caveat the top of the hill (Locality 1). The
fossil was recognizably different from almost everything that had been found previously, with the
possible exception ofJava Man. A new species was named and defined: Sinanthropus pekinensis.
The specimens from Zhoukoudian are now included in the species Homo erectus, but in the
popular mind both then and now, they may forever be known as “Peking Man” (see Figure 4.8,
bottom). The cave at Dragon Bone Hill was spectacularly productive by any standards. By the time
excavations were finished at Zhoukoudian, the expedition had recovered, along with thousands of
specimens of ancient animals, 15 fragmentary skulls, 6 more complete crania, 13 fragmentary
mandibles, 3 upperjaws, some postcranial bones (including pieces of femur, upper arms, toe
bones, numerous teeth), and a single vertebra of Peking Man ( Jia and Huang 1990:161-62). All
told,the remains of more than 40 hominin individuals were recovered from deposits in the cave.
Though dating of the hominin occupation of the cave has been fraught with uncertainty, a recent
redating of the site shows that the cave was first occupied approximately 770,000 years ago
(Guanjun et al. 2009). Being so numerous and discovered so early in our thinking about human
evolution, the Peking Man fossils played an important historical role in the scientific
conceptualization of human evolution and in interpretations of the culture of ancient human
beings. Tragically, the Zhoukoudian hominin assemblage was lost during World War II when the
fossils were being removed from China by U.S. Marines in an attempt to keep them away from
Japanese invaders, coincidentally on the same day that Pear] Harbor was attacked. The Marines
were captured and imprisoned, and to this day no one knows what became of their precious fossil
cargo. (It may have been destroyed by Japanese troops or simply lost, or it may have been found
later by Chinese druggists who ground the bones up for medicine. There is even a very slight
possibility that some are still hidden away in China, Japan, or the United States.

SUMMARY

Sometime after 1.8 million years ago, Homo habilis was replaced by a new hominin species, Homo
erectus. Erectus possessed a larger brain than habilis; its mean brain size of just under 1,000 cc is
two-thirds the modern human mean. With its larger brain and attendant greater intelligence, Homo
erectus was able to adapt to the changing environmental conditions posed by the Pleistocene
epoch. Homo erectus was the first ancestral human being to expand beyond the borders of our
hominin family’s African birthplace and nursery. Following the most reasonable trail beyond the
borders ofAfrica, Homo erectus fossils and their tools are found in southwest Asia and Eurasia
more than 1.7 million years ago, very soon after they first appeared in Africa. For reasons that are
still debated, these SUMMARY 137 African migrants did not enter into western Europe until later,
perhaps about 1.2 million years ago. It was intelligence and not any physical adaptation that
enabled Homo erectus to adapt to the diversity of habitats offered throughout Asia. New and more
sophisticated tools, new methods of hunting, and the use offire were all part of the Homo erectus
behavioral repertoire. Homo erectus was a stable and long-lived species. Fossils from Africa to east
Asia show a consistent morphology from close to 1.8 million to 400,000 years ago. After 400,000
years ago, brain size,relatively stable during the existence of erectus, exhibits a rapid increase,
signifying the evolution of the first Homo sapiens from an erectus base

HOMO ERECTUS: THE TOOLMAKER

In a class I teach called “Experimental Archaeology,” we spend a lot of time trying to replicate, as
authentically as possible, various stone tools made by prehistoric people. We follow a
chronological, evolutionary sequence,first replicating the Oldowan tools ofHomo habilis and then
making copies ofthe Acheulean handaxe (named for the French site of Saint-Acheul, where they
were first identified) that typifies Homo erectus, at least in Africa and Europe. Students generally
have little trouble making impressive versions of Oldowan choppers and flake tools—with a little
effort and after mastering the proper striking angle of hammerstone on core. This is not the case,
however, for the Homo erectus handaxe, which is not that easy to make, at least not without lots of
practice, knowledge, and time. Only a few of my students develop proficiency in handaxe
production. I joke in the class that when it comes to “evolving” our abilities at making ever more
complex stone tools “we leave no hominin behind.” But the truth is, sometimes we do. The oldest
handaxes have been found at the Kokiselei 4 site, located on the western margin of Lake Turkana in
Kenya and dating to 1.76 million years ago (Figure 4.12; Lepre et al. 2011). This date corresponds
very closely with the age of the oldest Homo erectus specimens discussed earlier in this chapter.
Even to an untrained eye, handaxes look much more tightly patterned and even more aesthetically
pleasing than the Oldowan tools discussed in Chapter 3. In fact, when shown an Oldowan tool
many think it just looks like a beat-up rock. Everyone, when shown an Acheulean handaxe,
immediately recognizes it as a human-made tool.

In a truly amazing research project, well-known stone tool replicator Bruce Bradley was hooked up
to a device (a positron emission tomography scanner) that tracked his brain activity while making
an Oldowan chopper and a late-style Acheulean handaxe (Normille 2012). Researchers determined
that while Bradley was making the Oldowan tool, the scanner showed brain activity normally
associated with hand gripping and coordination. That’s not surprising. However, when Bradley was
making the handaxe, the areas in his brain associated with abstract thought, hierarchical
organization, and language lit up. In other words, all kinds of stuff is going on in the brain of an
individual making a complex stone tool like a handaxe; you can’t accomplish that without a brain
capable of such activity.

Through a process ofbifacial flaking, a symmetrical, finely made handaxe was produced. Compare
this to the process of making Oldowantools (see Figure 3.18). (Noel G. Coonce) They're okay, and
my students are marginally impressed, but I wouldn't display them in a museum. Later Acheulean
tradition handaxes especially are symmetrical, finely flaked, and often aesthetically exquisite (see
Figure 4.13). Dozens offlakes are removed from the core, notjust a few (Figure 4.14). Even quite
simple handaxes can take 25 individual hammerstrikes. The best-made examples, which date to
after 1 million B.P, in Africa and more recently in Europe, took nearly three times that number
(Constable 1973:128). Each flake blow must be located precisely in order to allow for the proper
positioning of the next strike. The stone must be turned over again and again between
hammerstrikes to maintain symmetry and to keep the edge of the tool straight. All—or, at least,
most—of the exterior rind, or cortex, of the object piece was removed in order to keep the tool
relatively thin and light, so flakes needed to shoot across the face of the axe at the same time that
the edge was being maintained. This takes great skill, precision, and strength.

Experimental archaeologist Mark Newcomer (1971) has replicated handaxes, determining that at
least some were made in three separate steps. First, a blank, or preform, was roughed out with a
stone hammer into the general shape of the desired end product. Then the preform was refined via
a second stage of percussion with a softer stone or even a piece of antler used in thinning the tool.
Finally, the edges were straightened and sharpened in one last application of percussion. All the
work was worth it: For the same mass of stone, a handaxe produces about four times more cutting
edge than an Oldowan chopper and, at the same time, yields far more usable, sharp flakes. During
the production of a single handaxe, Newcomer produced more than 50 flakes usable for cutting or
scraping. The handaxe appears to have been an all-purpose tool (a colleague of minecalls them the
“Swiss Army rocks”ofthe Pleistocene). Its sharp tip was used for piercing, the thin edges for cutting,
and the steeper-angled edges toward the butt of the tool for scraping or chopping.

SUBSISTENCE

Remember one major issue concerning the subsistence base of Homo habilis: Did the members
ofthis species hunt big game or merely scavenge remnants of kills left by carnivores? As a whole,
the evidence supports the view that Homo habilis had a broad and opportunistic subsistence
strategy: Big-game hunting was probably not preeminent, but hunting, scavenging, and gathering
wild plants together provided subsistence for Homo habilis. Evidence for the importance of hunting
in Homo erectus is more substantial. For example, the remains of four butchered rhinoceroses
were found at the Box grove Quarry site in England. The 150 handaxes found at the site would have
served well in butchering the thick-skinned rhinos (Pitts and Roberts 2000). The researchers at
Boxgrove have demonstrated that butchering marks were made on the animals’ vertebrae, just
where you would expect for the most efficient dismembering of the animal. Beyond this, the
excavators recovered a horse scapula at Boxgrove exhibiting a wound that appears to have been
made with a spear (Pitts and Roberts 2000:260). There is clear evidence that Homo erectus
butchered animals at Aridos 1 and 2, 18 km (11 mi) southeast of Madrid, Spain (Villa 1990). Both
sites produced a butchered elephant, dating to probably about 350,000 years ago. Mixed in with the
bones were the tools used to cut off the meat, as well as the waste flakes produced in sharpening
the cutting and scraping tools. While there is no evidence of hunting at either Aridos site—no
spearpoints were found, for example—there also is no evidence that Homo erectus gained access
to the carcasses after carnivores had their fill—there are no tooth or gnaw marks on the bones.
Thefacts that the flint serving as the raw material for the tools at Aridos came from deposits at a 3-
km distance and that much of the toolmaking occurred away from the butchery site, with mostly
sharpening conducted on-site, offer evidence for what researcher Paola Villa calls “planning
depth.” In other words, the hominins who butchered the elephant carcasses at Aridos knew that
elephants roamed the area and that there was always a chance one or more of them might die at
any given time—the Aridos 1 elephant was a juvenile and Aridos 2 was an old male, both part of
subpopulations with high natural mortality rates. The Homo erectus population in the area seemsto
have planned for the lucky occasions when a carcass became available by collecting the raw
material and making the tools in advance. Planning ahead for a future eventuality is a human
behavior, and the evidence at Aridos suggests that Homo erectus was capable of such behavior.
Additional evidence of the importance of meat in the diet of Homo erectus comes in the form of
weapons that appear to bear witness to hunting. At the Kathu Pan site in South Africa, researchers
have recovered what appear to be stone points, the business end ofspears dating to about 500,000
years ago (Wilkins et al. 2012). Each of the artifacts is roughly triangular with thin lateral edges, a
sharp, pointed tip, and a dull, broad base. So they look like spearpoints. Beyond this, there is
evidence of intentional flaking and thinning toward the base of the artifacts. This likely was the
result of the makers trying to make it easier to attach the points to a long wooden shaft. Finally,
there is clear evidence of damage to the working tips of these artifacts, breakage that likely resulted
from their having banged against something hard, like the bone of a prey animal, or the rocky
ground after a missed throw or thrust. Applying the methods of experimental archaeology (see
Chapter 2), researchers made replicas of the artifacts, hafted them onto wooden shafts, and then
used them as spears. In those experiments, various