HOMOERECTUS

Discovery Date: 1891

Where Lived: Northern, Eastern, and Southern Africa; Western Asia (Dmanisi, Republic of Georgia); East Asia
(China and Indonesia)
When Lived: Between about 1.89 million and 110,000 years ago
Homo erectus lived between about 1.89 million and 110,000 years ago.
Height: Ranges from 4 ft 9 in - 6 ft 1 in (145 - 185 cm)
Weight: Ranges from 88 - 150 lbs (40 - 68 kg)
Overview
Early African Homo erectus fossils (sometimes called Homo ergaster) are the oldest known early humans to
have possessed modern human-like body proportions with relatively elongated legs and shorter arms compared
to the size of the torso. These features are considered adaptations to a life lived on the ground, indicating the
loss of earlier tree-climbing adaptations, with the ability to walk and possibly run long distances. Compared
with earlier fossil humans, note the expanded braincase relative to the size of the face. The most complete fossil
individual of this species is known as the ‘Turkana Boy’ – a well-preserved skeleton (though minus almost all
the hand and foot bones), dated around 1.6 million years old. Microscopic study of the teeth indicates that he
grew up at a growth rate similar to that of a great ape. There is fossil evidence that this species cared for old and
weak individuals. The appearance of Homo erectus in the fossil record is often associated with the earliest
handaxes, the first major innovation in stone tool technology.
Early fossil discoveries from Java (beginning in the 1890s) and China (‘Peking Man’, beginning in the 1920s)
comprise the classic examples of this species. Generally considered to have been the first species to have
expanded beyond Africa, Homo erectus is considered a highly variable species, spread over two continents (it's
not certain whether it reached Europe), and possibly the longest lived early human species - about nine times as
long as our own species, Homo sapiens, has been around!
History of Discovery:
Eugène Dubois, a Dutch surgeon, found the first Homo erectus individual (Trinil 2) in Indonesia in 1891. In
1894, Dubois named the species Pithecanthropus erectus, or ‘erect ape-man.’ At that time, Pithecanthropus
(later changed to Homo) erectus was the most primitive and smallest-brained of all known early human species;
no early human fossils had even been discovered in Africa yet
How They Survived:
The tall bodies and large brains of Homo erectus individuals required a lot of energy on a regular basis to
function. Eating meat and other types of protein that could be quickly digested made it possible to absorb
nutrients with a shorter digestive tract, making more energy available faster. There is also speculation that
honey and underground tubers may have been significant food sources for Homo erectus.
Soon after we see evidence in the fossil record of the earliest Homo erectus fossils (by about 1.9 million years
ago), we see evidence in the archeological record for the first major innovation in stone tool technology (by
about 1.76 million years ago). Known as the Acheulean stone tool industry, it consisted of the creation of large
cutting tools like handaxes and cleavers. Increased reliance on a broader set of tools may have helped Homo
erectus survive during changing climates.
The earliest evidence of hearths (campfires) occur during the time range of Homo erectus. While we have
evidence that hearths were used for cooking (and probably sharing) food, they are likely to have been places for
social interaction, and also used for warmth and to keep away large predators.

Another Overview:
Homo erectus, (Latin: “upright man”) extinct species of the human genus (Homo), perhaps an ancestor of
modern humans (Homo sapiens). H. erectus most likely originated in Africa, though Eurasia cannot be ruled
out. Regardless of where it first evolved, the species seems to have dispersed quickly, starting about 1.9 million
years ago (mya) near the middle of the Pleistocene Epoch, moving through the African tropics, Europe, South
Asia, and Southeast Asia. This history has been recorded directly if imprecisely by many sites that have yielded
fossil remains of H. erectus. At other localities, broken animal bones and stone tools have indicated the presence
of the species, though there are no traces of the people themselves. H. erectus was a human of medium stature
that walked upright. The braincase was low, the forehead was receded, and the nose, jaws, and palate were
wide. The brain was smaller and the teeth larger than in modern humans. H. erectus appears to have been the
first human species to control fire, some 1,000,000 years ago. The species seems to have flourished until some
200,000 years ago (200 kya) or perhaps later before giving way to other humans including Homo sapiens.
FOSSIL EVIDENCE
The earliest finds
The first fossils attributed to Homo erectus were discovered by a Dutch army surgeon, Eugène Dubois, who
began his search for ancient human bones on the island of Java (now part of Indonesia) in 1890. Dubois found
his first specimen in the same year, and in 1891 a well-preserved skullcap was unearthed at Trinil on the Solo
River. Considering its prominent browridges, retreating forehead, and angled rear skull, Dubois concluded that
the Trinil cranium showed anatomic features intermediate between those of humans (as they were then
understood) and those of apes. Several years later, near where the skull was discovered, he found a remarkably
complete and modern-looking femur (thighbone). Since this bone was so similar to a modern human femur,
Dubois decided that the individual to which it belonged must have walked erect. He adopted the name
Pithecanthropus (coined earlier by the German zoologist Ernst Haeckel) and called his discoveries
Pithecanthropus erectus (“upright ape-man”), but the colloquial term became “Java man.” Only a few other
limb fragments turned up in the Trinil excavations, and it would be some three decades before more substantial
evidence appeared. Most paleontologists now regard all of this material as H. erectus, and the name
Pithecanthropus has been dropped.
Fossil Discoveries: https://australian.museum/learn/science/human-evolution/homo-erectus/
After years of searching Indonesia for ‘the missing link’, Dutchman Eugene Dubois finally uncovered part of a
skull in 1891 (known as ‘Java Man’). He believed this fossil belonged to an ancient and ‘upright’ human and so
coined the species name erectus. Other scientists dismissed this interpretation, preferring to emphasise its ape-
like qualities. Dubois’ opinion was validated when a series of similar fossils were uncovered in China during
the 1920s and 1930s.
Important Homo erectus specimens
Sangiran 17 – a 1.2 million-year-old skull discovered in 1969 in Sangiran, Indonesia. This adult male skull is
the best preserved Homo erectus skull from Java.
Zhoukoudian 3 – a skullcap discovered in 1929 in Zhoukoudian, China. This adolescent’s skullcap was
originally found in fragments. When the pieces were fitted together, they showed that this young individual had
a brain size of 915 cubic centimetres.
‘Java Man’ or Trinil 2 – a skullcap discovered in 1891 by Eugene Dubois in Trinil, Indonesia. This fossil was
nicknamed ‘Java Man’ because it was found on the island of Java. It is the ‘type specimen’ or official
representative for the species.
Sangiran 2 – a 1 million-year-old skullcap discovered in 1937 in Sangiran, Indonesia
‘Solo Man’ or Ngandong – a skull cap discovered in 1932 in Ngandong, Indonesia. Because its exact original
location is unknown, published dates have ranged from 35,000 to 500,000 years old. ‘Solo Man’ shares
similarities with earlier Homo erectus specimens from Sangiran and is considered to be a late Homo erectus.
‘Mojokerto’ or Perning 1 – a juvenile skull discovered in 1936 in Mojokerto, Indonesia. Radiometric dates have
suggested this child’s skull may be as old as 1.8 million years, which significantly increases the previous dates
for Homo erectus in Asia. However, this date is debated as the sediment sample taken for dating was taken
about 60 years after the skull was collected and the two may have come from different levels.
Zhoukoudian 5 – a partial skull discovered in Zhoukoudian, China. This skull was reconstructed from several
pieces found in 1934, 1936 and 1966.
‘Peking Man’ discovered in Zhoukoudian, China. The original ‘Peking Man’ skull was reconstructed using a
mixture of male and female fossils whereas the modern reconstruction by I. Tattersal and G. Sawyer combines
fossil pieces from males only.
Sangiran 4 – a 1.5 million-year-old upper jaw discovered in 1939 in Sangiran, Indonesia. The canine teeth were
larger than those found in modern humans. This is one of the oldest specimens from Sangiran.
Sangiran 1 – a 1.5 million-year-old partial lower jaw discovered in 1936 in Sangiran, Indonesia. This is the first
human fossil discovered at Sangiran.
What the name Homo erectus means
Homo, is a Latin word meaning ‘human’ or ‘man’ and is the genus or group name of this species.
The second word in this species’ scientific name is erectus. This name was selected to indicate that this species’
ability to stand and walk with an upright or erect stance.
Homo erectus Key physical features
This species had a robust skeleton that was generally similar to those of modern humans. However, the skulls of
Homo erectus were quite different to those of modern humans.
Body size and shape
 the body (known only from the Chinese specimens) tended to be shorter and stockier than those of
modern humans.
Brain
 showed an increase in size over earlier species and averaged about 1050 cubic centimetres
structure of the brain was similar to that of modern humans
Skull
  face was large with a low, sloping forehead, a massive brow ridge and a broad, flat nose
 skull was broad and long with sharp angles at the rear, unlike the curve found in modern humans
 bones of the skull were very thick and formed a small central ridge, known as a midline keel, along the
top of the skull
Jaws and teeth
 jaw was large and thick without a pointed chin
 molar teeth had large roots but were decreasing toward a more modern size
Limbs
 limbs were like those of modern humans although the bones were thicker, suggesting a physically
demanding lifestyle.
Homo erectus Lifestyle
Culture
The oldest known stone tools used by Homo erectus were made in China about one million years ago. These
tools were simple choppers and flakes (Mode 1 technology). Tools gradually became smaller over time and
came to include a greater variety of designs. More complex bifacial tools (Mode 2 technology) like those made
by Homo heidelbergensis people from Eurasia made a limited appearance in parts of northern China. This may
indicate a brief period of contact between these different peoples.
Relatively few stone tools have been found in east Asia compared with western Asia, Africa and Europe but
tools made from non-durable materials such as bamboo may have been manufactured instead.
Burnt stones and animal bones, charcoal and ash deposits indicate these people may have used fire about
500,000 years ago but it is difficult to prove whether this use was controlled.
Environment and diet
China underwent significant climatic changes during the period that Zhoukoudian was occupied. These changes
included three cold glacial periods with harsh, winter temperatures. The cooling and drying that occurred in
these glacial periods brought an expansion of open habitats, with grasslands and mixed steppes. These
environments favoured large grazing animals, which would have been hunted by Homo erectus.
Java, on the other hand, had a warmer climate. Low sea levels about 1.6 million years ago (when this area was
first occupied by H.erectus) may have seen Indonesia joined to mainland southeast Asia.
The remains of meals have been found at some Homo erectus sites in China. These show that they ate large
amounts of meat supplemented with plant foods and, in general, had a diet similar to that of early modern
humans.
Another Overview: https://www.nature.com/scitable/knowledge/library/homo-erectus-a-bigger-smarter-
97879043/
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.
History and Geography
Eugene Dubois first identified and described a new human-like set of Indonesian fossils at the end of the 19th
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 cm3, 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 cm3, 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.