Chapter Summary and Key Concepts

Learning Objectives

After reading this chapter, students should be able to:

  • recount the history of the discovery of Lower Paleolithic hominins and how this relates to our understanding of evolution
  • describe the evolution of hominins from 1.7–1.6 million years ago to 250,000 years ago, the Lower Paleolithic/Early Stone Age
  • explain the distinguishing features of H. ergaster
  • understand the relationships between H. ergaster and later hominins
  • describe the relevant biological and morphological differences that relate to each species, and why current phylogenies are constructed as they are
  • be conversant with the important fossils and sites from which evidence is drawn
  • extrapolate from biological and artifactual evidence what we know about the social and cultural behavior of various hominins of this period
  • explain hominin migrations, and the differing implications of early and later dates for hominin migration out of Africa
  • describe the differences between Acheulean African, European, and Southwest Asian people and non-Acheulean/Acheulean age people found in East Asia and Indonesia
  • describe Acheulean culture, including tools, behavior, and biology
  • relate what is known about early, failed attempts at European colonization
  • understand the implications of new finds at Dmanisi, Denisova, dates from Java, and other current controversies
  • be conversant with the debates over hunting vs. scavenging, site modification, fire use, symbolic and other current issues

Chapter Summary

Homo ergaster

Homo ergaster, an African species, existed between 1.8–1.7 million and 600,000 years ago. Important sites include Koobi Fora (East Turkana) and Nariokotome III (West Turkana). H. ergaster, discovered in the 1970s, was originally classified as H. erectus due to shared morphological features with the East Asian hominin discovered in 1891, but despite some dissent, due to their relative ages, most now believe that that H. erectus originated in Africa, later migrating east. In addition, African H. ergaster skulls are more primitive than H. erectus.

It was once believed that H. erectus was directly ancestral to H. sapiens. If so, H. ergaster would be an early stage of H. erectus. But fossils dating after 600,000 years ago now indicate that H. sapiens evolved in Africa while H. erectus continued on largely unchanged in East Asia. As H. sapiens emerged in Africa, H. neanderthalensis, was evolving in Europe. H. sapiens and H. neanderthalensis last shared a common ancestor about 600,000–500,000 years ago; this ancestor has been designated H. heidelbergensis. Thus H. ergaster is ancestral to H. erectus and H. heidelbergensis and its descendants, H. sapiens and H. neanderthalensis.


Homo ergaster displayed cognitive and behavioral advances over H. habilis, linked to its ability to create new tools, colonize arid, seasonal environments, and expand out of Africa. H. ergaster meticulously shaped cores into the characteristic hand axe: completely bifacially flaked, teardrop-shaped with a broad base and rounded point. Ovals, triangles, and sometimes cleavers were common. These define the Acheulean cultural tradition, spanning over a million years and three continents.

While “hand axe” implies a hand-held chopper, many are too large, and their precise use remains a puzzle, a problem increased at sites with hundreds of closely packed hand axes with no obvious signs of use. Most sites have smaller numbers of hand axes that show signs of use. They came in many sizes and shapes, probably serving multiple utilitarian functions. Some may have been hurled, others used as portable sources of flakes, others to chop or scrape wood. Experiments show that they are effective butchering tools.

Homo erectus

To understand the controversies surrounding H. erectus and H. ergaster, the discovery and dating of its East Asian fossils must be examined.

THE DISPERSION OF Homo ergaster AND THE FATE OF Homo erectus

Archaeology shows that about 1.5 million years ago, shortly after appearing in Africa, H. ergaster colonized the northern and southern margins of the continent, passing through a more hospitable Saharan region. A group on the periphery might periodically outgrow its resources, and a splinter party could find empty territory nearby. From northeastern Africa, such groups probably colonized eastward toward China and Indonesia and northwestward toward Europe without even knowing that they had left Africa.

Dispersal into eastern Asia occurred by 1 million years ago, perhaps earlier. Until recently, the oldest accepted European evidence was about 800,000 years old, from Gran Dolina cave, Spain. However, recent discoveries at Dmanisi, Republic of Georgia, may be earlier. Dmanisi lies 1500 km (940 miles) north of ‘Ubeidiya, and yielded more than 1000 artifacts and many human fossils.

The Persistence of Homo erectus in Java

By 600,000–500,000 years ago, along with more sophisticated hand axes, hominins with larger, more modern braincases appeared in Africa, probably evolving abruptly from Homo ergaster. They closely resembled Europeans of 500,000–400,000 years ago, so are grouped together as Homo heidelbergensis. H. heidelbergensis’ expansion from Africa probably explains the introduction of Acheulean artifacts to Europe about 500,000 years ago. Homo heidelbergensis may be the last shared ancestor of the Neanderthals and modern humans.


No European site is indisputably older than 800,000 years, and only one or two are older than 500,000 years. This implies obstacles to early human settlement, particularly glacial conditions. Fossils from Petralona, Greece, and Arago, France, suggest descent from Homo heidelbergensis: expanding Africans who brought the late Acheulean tradition to Europe c. 500,000 years ago.

Homo heidelbergensis had large, prognathic faces, massive, chinless jaws with big teeth, large browridges, low, flat foreheads, and thick skulls. Yet they had a much enlarged brain, averaging over 1200 cc. Their anatomy and geographic distribution make them a plausible common ancestor for Neanderthals and modern humans.

Brain Expansion and Change within the Hand Axe Tradition

The Acheulean tradition, as noted, had an early phase, before 600,000 years ago, with thick, asymmetric hand axes and a later phase, after 600,000 years ago, with thinner, trimmed, more symmetrical examples. This greater technological sophistication may have been crucial for colonization of Europe.

The Evolution of the Neanderthals in Europe

After 500,000 years ago, Neanderthals evolved in Europe while modern humans evolved in Africa. Older finds from Swanscombe, England, and Steinheim, Germany, augmented with recent finds at Sima de los Huesos, Atapuerca, Spain, shed light on Neanderthal development.


Knowledge of Acheulean people comes from fewer than 50 archaeological sites scattered in time and space, not always well-dated. The behavioral implications are often ambiguous. Despite this, we can say a few things about early human behavior apart from stone tools.

Raw Materials besides Stone

Animal bones at many sites suggest their use as hammers, retouchers, anvils, or cutting boards; a few were percussion-flaked to produce bifaces, scrapers, and choppers. Bamboo was likely used in eastern Asia, where hand axes are lacking. Actual bamboo artifacts remain unknown.

Site Modification

Shelters must have been needed in Eurasia, but apparently were ephemeral, leaving ambiguous traces. Patterned arrangements of large rocks found at several sites may be foundations for huts or windbreaks, but natural processes cannot be ruled out. At other sites, oval or round clusters of artifacts, bones, and debris could mark hut interiors. Evidence for housing becomes abundant and unambiguous after the advent of fully modern humans between 50,000 and 40,000 years ago.


Homo ergaster (or H. erectus) may have mastered fire for warmth and cooking before colonizing Eurasia, but direct evidence is tenuous. The oldest widely accepted evidence comes from Zhoukoudian Locality 1 in China dated to between 500,000 and 240,000 years ago. In Europe, several sites have evidence for fire dated between 400,000 and 300,000 years ago, and others date between 186,000 and 127,000 years ago.


Three examples of possible art are most convincing. First, fragments of humanly introduced ocher have been found with Acheulean artifacts and animal bones at Kapthurin, Kenya, Duinefontein 2, South Africa, and Twin Rivers, Zambia. Second, an elephant bone fragment from Bilzingsleben, Germany has incised fanlike lines (not butchery marks) created simultaneously with one stone tool. Third, a small, definitely humanly modified lava pebble from Berekhat Ram (Syrian/Israeli border) may represent a crude human figurine. Clear artistic objects become commonplace only after 50,000 years ago, with modern humans.

Diet and Food Procurement

Remains of edible plants are found at some Acheulean-age sites, but may not be linked to human activity. Archaeologists once simply assumed that Achuelean people were big-game hunters, killing and butchering animals with their tools. In the 1970s, specialized zooarchaeological analyses revealed that animal bones could represent human kills or carnivore kills and natural deaths, subsequently scavenged by people. While this is still debated, current consensus is that Acheuleans hunted large animals.

Box Features

Key Controversy: The Dating of Javan Homo erectus

Key Controversy: The “Hobbit”, Homo floresiensis: a Unique Species?

Key Controversy: When did Humans First Colonize Europe?

Key Controversy: Is Homo erectus Represented by DNA from Denisova Cave?

Key Discovery: The Discovery of the Turkana Boy

Key Discovery: The Acheulean Hand Axe Tradition

Key Method: Electron Spin Resonance (ESR) Dating

Key Method: Uranium-series Dating

Key Method: Luminescence Dating

Key Site: The Gran Dolina TD6 and the History of Cannibalism

Key Concepts

Hominin Species

  • Hominin species
  • Homo ergaster
  • Homo erectus
  • Homo heidelbergensis
  • Homo neanderthalensis
  • Homo antecessor
  • Pithecanthropus erectus
  • Sinanthropus pekinensis


  • Lake Turkana
  • East Turkana, West Turkana, Kenya
  • Olduvai Gorge, Tanzania
  • Koobi Fora, Kenya
  • Nariokotome III, Kenya
  • Konso, Ethiopia
  • Karari Escarpment, Kenya
  • Peninj (Lake Natron), Tanzania
  • Melka Kunturé, Ethiopia
  • Olorgesailie, Kenya
  • Isimila, Tanzania
  • Kalambo Falls, Zambia
  • Trinil, Java, Indonesia
  • Sangiran, Java, Indonesia
  • Ngandong, Java, Indonesia
  • Sambungmacan, Java, Indonesia
  • Zhoukoudian, Locality 1, China
  • Gongwangling, China
  • Lontandong Cave, China
  • Nihewan Basin, China
  • ‘Ubeidiya, Israel
  • Gran Dolina, TD6, Atapuerca, Spain
  • Dmanisi, Republic of Georgia
  • Petralona, Greece
  • Denisova, Russia
  • Arago, France
  • Sima de los Huesos, Atapuerca, Spain
  • Ceprano, Italy
  • Swanscombe, England
  • Steinheim, Germany
  • Fontana Ranuccio, Italy
  • Malagrotta, Italy
  • Castel di guido, Italy
  • La polledrara, Italy
  • Bilzingsleben, Germany
  • Gesher Benot Ya’aqov, Israel
  • Clacton-on-Sea, England
  • Happisburgh, England
  • Schöningen, Germany
  • Soleihac, France
  • Latamne, Syria
  • Terra Amata, France
  • Ariendorf 1, Germany
  • Le Lazaret Cave, France
  • La Baume Bonne, France
  • Orgnac, France
  • Chesowanja, Kenya
  • Montagu Cave, South Africa
  • Swartkrans cave, South Africa
  • Cave of Hearths, South Africa
  • Prezletice, Czech Republic
  • Vértesszöllös, Hungary
  • Menez-Dregan, France
  • Pech de l’Azé, France
  • La Cotte de St. Brelade, Jersey
  • Kapthurin, Kenya
  • Duinefontein 2, South Africa
  • Twin Rivers, Zambia
  • Berekhat Ram, Syrian/Israeli border

Important fossils

  • Turkana Boy
  • KNM WT 15000
  • D2700, 2280, and D2282
  • KNM-ER-1813
  • Olduvai hominid 9
  • Buia and Daka skulls
  • KNM-ER 1808


  • Eugène Dubois
  • G.H.R. von Koenigswald
  • J.G. Andersson
  • Davidson Black
  • Franz Weidenreich
  • Hallam L. Movius

Dating methods

  • potassium-argon
  • fission-track
  • paleomagnetism
  • biostratigraphy
  • electron spin resonance
  • climate dating
  • sediment accumulation rate
  • uranium series

Hominin morphology terms

  • braincase
  • forehead (frontal bone)
  • browridge
  • forward projecting nose
  • barrel-shaped chest
  • narrowed hips
  • prognathic face
  • chinless jaw
  • mastoid process
  • constricted birth canal
  • infant dependency
  • digestive tract
  • tall, slim, heat dissipation
  • moisture condensation
  • perspiration
  • sexual dimorphism


  • Acheulean cultural tradition
  • hand axes, cleavers, large bifacial ovals, triangles
  • Early vs. Late Acheulean
  • Levallois technique
  • colonization of hot, arid, seasonal environments
  • colonization of cold, glacial environments
  • Movius line
  • bamboo
  • early dispersion vs. technological bottleneck
  • population budding
  • bone tools
  • wooden tools
  • shelters
  • fire
  • ocher
  • lava pebble figurine
  • incised elephant tibia
  • plant foods
  • yaws
  • vitamin A
  • hunting and scavenging
  • big-game hunters
  • skeletal part representation
  • cutmarks, butchering