Ice Age

megafauna and time notes

Contents
1

Ice age

1.1

Origin of ice age theory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.2

Evidence for ice ages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.3

Major ice ages

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.4

Glacials and interglacials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.5

Positive and negative feedback in glacial periods . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.5.1

Positive feedback processes

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.5.2

Negative feedback processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Causes of ice ages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.6.1

Changes in Earths atmosphere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.6.2

Position of the continents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.6.3

Fluctuations in ocean currents

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.6.4

Uplift of the Tibetan plateau and surrounding mountain areas above the snowline . . . . . .

1.6.5

Variations in Earths orbit (Milankovitch cycles) . . . . . . . . . . . . . . . . . . . . . . .

1.6.6

Variations in the Suns energy output . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.6.7

Volcanism

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Recent glacial and interglacial phases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.7.1

Glacial stages in North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.7.2

Last Glacial Period in the semiarid Andes around Aconcagua and Tupungato . . . . . . . .

1.8

Eects of glaciation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1.9

See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

10

1.6

1.7

1.10 References

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

10

1.11 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

13

Megafauna

14

2.1

Ecological strategy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

14

2.2

Evolution of large body size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

14

2.2.1

In terrestrial mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

15

2.2.2

In marine mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

15

2.2.3

In ightless birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

15

Megafaunal mass extinctions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

16

2.3.1

Timing and possible causes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

16

2.3.2

Consequences of depletion of megafauna . . . . . . . . . . . . . . . . . . . . . . . . . . .

17

2.3

ii

CONTENTS
2.4

Examples . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

17

2.5

Gallery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

21

2.5.1

Extinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

21

2.5.2

Living . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

22

2.6

See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

23

2.7

Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

23

2.8

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

23

2.9

External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

26

Pleistocene

27

3.1

Dating . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

27

3.2

Paleogeography and climate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

28

3.2.1

Glacial features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

28

3.2.2

Major events . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

28

3.2.3

Palaeocycles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

29

3.3

Fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

30

3.4

Humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

30

3.5

Deposits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

31

3.6

See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

31

3.7

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

31

3.8

External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

32

Prehistoric mammal

33

4.1

List of prehistoric mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

33

4.2

See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

33

Stone Age

34

5.1

Historical signicance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

34

5.2

The Stone Age in archaeology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

34

5.2.1

Beginning of the Stone Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

34

5.2.2

End of the Stone Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

35

5.2.3

The concept of Stone Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

35

5.2.4

The three-stage system . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

36

5.2.5

The problem of the transitions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

37

Chronology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

37

5.3.1

Three-age chronology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

37

5.3.2

Three-stage chronology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

43

Material culture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

43

5.4.1

Tools . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

43

5.4.2

Food and drink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

43

5.4.3

Shelter and habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

44

5.4.4

Art . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

44

5.3

5.4

CONTENTS
5.4.5

Stone Age rituals and beliefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

45

5.5

Modern popular culture and the Stone Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

45

5.6

See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

45

5.7

Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

46

5.8

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

47

5.9

Further reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

47

5.10 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

47

Woolly mammoth

49

6.1

Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

49

6.1.1

Etymology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

50

6.1.2

Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

50

Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

51

6.2.1

Coat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

52

6.2.2

Dentition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

53

Palaeobiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

54

6.3.1

Diet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

54

6.3.2

Growth and reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

55

6.4

Distribution and habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

56

6.5

Relationship with humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

57

6.5.1

Exploitation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

57

6.6

Extinction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

58

6.7

Frozen specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

59

6.7.1

Recreating the species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

61

Cultural signicance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

62

6.8.1

Cryptozoology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

62

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

63

6.10 Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

67

Woolly rhinoceros

68

7.1

Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

68

7.2

Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

68

7.3

Behavior and habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

68

7.3.1

Diet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

69

7.4

Extinction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

69

7.5

See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

70

7.6

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

70

7.7

External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

70

7.8

Text and image sources, contributors, and licenses . . . . . . . . . . . . . . . . . . . . . . . . . .

71

7.8.1

Text . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

71

7.8.2

Images . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

74

7.8.3

Content license . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

78

6.2

6.3

6.8
6.9

iii

Chapter 1

Ice age
4

This article is about a generic geological period of temperature reduction. For the most recent glacial period
commonly referred to as the Ice Age, see Last glacial period. For other uses, see Ice age (disambiguation).
An ice age is a period of long-term reduction in the

Temperature variation (T)

0
2
4
6
8
10

400

350

300

250

200

150

100

300

50

Carbon Dioxide

280

ppmv

260
240
220
200

ppm

180
1.8
1.6
1.4
1.2
1
0.8
0.6
0.4
0.2
0

400

350

300

250

200

150

100

50

Dust concentration

400

350

300

250

200

150

100

50

Thousands of years ago

Variations in temperature, CO
2, and dust from the Vostok ice core over the last 400,000 years

temperature of Earth's surface and atmosphere, resulting

in the presence or expansion of continental and polar ice
sheets and alpine glaciers. Within a long-term ice age,
individual pulses of cold climate are termed "glacial periods" (or alternatively glacials or glaciations or colloquially as ice age), and intermittent warm periods are
called "interglacials". Glaciologically, ice age implies the
presence of extensive ice sheets in the northern and southAn artists impression of ice age Earth at glacial maximum. ern hemispheres.[1] By this denition, we are in an interBased on: Crowley, T.J. (1995). Ice age terrestrial carbon glacial periodthe holoceneof the ice age that began
changes revisited. Global Biogeochemical Cycles 9 (3): 377
2.6 million years ago at the start of the Pleistocene epoch,
389. Bibcode:1995GBioC...9..377C. doi:10.1029/95GB01107.
because the Greenland, Arctic, and Antarctic ice sheets
still exist.[2]

1.1 Origin of ice age theory

In 1742 Pierre Martel (17061767), an engineer and geographer living in Geneva, visited the valley of Chamonix
in the Alps of Savoy.[3][4] Two years later he published
an account of his journey. He reported that the inhabitants of that valley attributed the dispersal of erratic
boulders to the fact that the glaciers had once extended
much farther.[5][6] Later similar explanations were reported from other regions of the Alps. In 1815 the carpenter and chamois hunter Jean-Pierre Perraudin (1767
1858) explained erratic boulders in the Val de Bagnes in
the Swiss canton of Valais as being due to glaciers previ-

The Antarctic ice sheet. Ice sheets expand during an ice age.

2
ously extending further.[7] An unknown woodcutter from
Meiringen in the Bernese Oberland advocated a similar idea in a discussion with the Swiss-German geologist
Jean de Charpentier (17861855) in 1834.[8] Comparable explanations are also known from the Val de Ferret
in the Valais and the Seeland in western Switzerland[9]
and in Goethe's Scientic Work.[10] Such explanations
could also be found in other parts of the world. When the
Bavarian naturalist Ernst von Bibra (18061878) visited
the Chilean Andes in 18491850 the natives attributed
fossil moraines to the former action of glaciers.[11]
Meanwhile, European scholars had begun to wonder what
had caused the dispersal of erratic material. From the
middle of the 18th century some discussed ice as a means
of transport. The Swedish mining expert Daniel Tilas
(17121772) was, in 1742, the rst person to suggest
drifting sea ice in order to explain the presence of erratic
boulders in the Scandinavian and Baltic regions.[12] In
1795, the Scottish philosopher and gentleman naturalist,
James Hutton (17261797), explained erratic boulders in
the Alps with the action of glaciers.[13] Two decades later,
in 1818, the Swedish botanist Gran Wahlenberg (1780
1851) published his theory of a glaciation of the Scandinavian peninsula. He regarded glaciation as a regional
phenomenon.[14] Only a few years later, the DanishNorwegian Geologist Jens Esmark (17621839) argued
a sequence of worldwide ice ages. In a paper published in
1824, Esmark proposed changes in climate as the cause
of those glaciations. He attempted to show that they originated from changes in Earths orbit.[15] During the following years, Esmarks ideas were discussed and taken
over in parts by Swedish, Scottish and German scientists. At the University of Edinburgh Robert Jameson
(17741854) seemed to be relatively open to Esmarks
ideas, as reviewed by Norwegian professor of glaciology Bjrn G. Andersen (1992).[16] Jamesons remarks
about ancient glaciers in Scotland were most probably
prompted by Esmark.[17] In Germany, Albrecht Reinhard Bernhardi (17971849), a geologist and professor
of forestry at an academy in Dreissigacker, since incorporated in the southern Thuringian city of Meiningen,
adopted Esmarks theory. In a paper published in 1832,
Bernhardi speculated about former polar ice caps reaching as far as the temperate zones of the globe.[18]

CHAPTER 1. ICE AGE

pentier presented his paper before the Schweizerische
Naturforschende Gesellschaft.[20] In the meantime, the
German botanist Karl Friedrich Schimper (18031867)
was studying mosses which were growing on erratic boulders in the alpine upland of Bavaria. He began to wonder
where such masses of stone had come from. During the
summer of 1835 he made some excursions to the Bavarian Alps. Schimper came to the conclusion that ice must
have been the means of transport for the boulders in the
alpine upland. In the winter of 1835 to 1836 he held
some lectures in Munich. Schimper then assumed that
there must have been global times of obliteration (Verdungszeiten) with a cold climate and frozen water.[21]
Schimper spent the summer months of 1836 at Devens,
near Bex, in the Swiss Alps with his former university
friend Louis Agassiz (18011873) and Jean de Charpentier. Schimper, de Charpentier and possibly Venetz
convinced Agassiz that there had been a time of glaciation. During Winter 1836/7 Agassiz and Schimper developed the theory of a sequence of glaciations. They mainly
drew upon the preceding works of Venetz, de Charpentier and on their own eldwork. There are indications that
Agassiz was already familiar with Bernhardis paper at
that time.[22] At the beginning of 1837 Schimper coined
the term ice age (Eiszeit).[23] In July 1837 Agassiz presented their synthesis before the annual meeting of the
Schweizerische Naturforschende Gesellschaft at Neuchtel. The audience was very critical or even opposed the
new theory because it contradicted the established opinions on climatic history. Most contemporary scientists
thought that the earth had been gradually cooling down
since its birth as a molten globe.[24]
In order to overcome this rejection, Agassiz embarked
on geological eldwork. He published his book Study on
glaciers ("tudes sur les glaciers) in 1840.[25] De Charpentier was put out by this as he had also been preparing
a book about the glaciation of the Alps. De Charpentier
felt that Agassiz should have given him precedence as it
was he who had introduced Agassiz to in-depth glacial
research.[26] Besides that, Agassiz had, as a result of personal quarrels, omitted any mention of Schimper in his
book.[27]
All together, it took several decades until the ice age theory was fully accepted. This happened on an international
scale in the second half of the 1870s following the work
of James Croll including the publication of Climate and
Time, in Their Geological Relations in 1875 which provided a credible explanation for the causes of ice ages.[28]

Independently of these debates, the Swiss civil engineer

Ignaz Venetz (17881859) in 1829, explained the dispersal of erratic boulders in the Alps, the nearby Jura
Mountains and the North German Plain as being due
to huge glaciers. When he read his paper before the
Schweizerische Naturforschende Gesellschaft, most scientists remained sceptical.[19] Finally, Venetz managed
to convince his friend Jean de Charpentier. De Char- 1.2 Evidence for ice ages
pentier transformed Venetzs idea into a theory with a
glaciation limited to the Alps. His thoughts resembled There are three main types of evidence for ice ages: geWahlenbergs theory. In fact, both men shared the same ological, chemical, and paleontological.
volcanistic, or in de Charpentiers case rather plutonistic
assumptions, about the earths history. In 1834, de Char- Geological evidence for ice ages comes in various forms,
including rock scouring and scratching, glacial moraines,

1.3. MAJOR ICE AGES

drumlins, valley cutting, and the deposition of till or

tillites and glacial erratics. Successive glaciations tend to
distort and erase the geological evidence, making it dicult to interpret. Furthermore, this evidence was dicult
to date exactly; early theories assumed that the glacials
were short compared to the long interglacials. The advent of sediment and ice cores revealed the true situation:
glacials are long, interglacials short. It took some time for
the current theory to be worked out.
The chemical evidence mainly consists of variations in
the ratios of isotopes in fossils present in sediments and
sedimentary rocks and ocean sediment cores. For the
most recent glacial periods ice cores provide climate
proxies from their ice, and atmospheric samples from included bubbles of air. Because water containing heavier
isotopes has a higher heat of evaporation, its proportion
decreases with colder conditions.[29] This allows a temperature record to be constructed. However, this evidence can be confounded by other factors recorded by
isotope ratios.
The paleontological evidence consists of changes in the
geographical distribution of fossils. During a glacial period cold-adapted organisms spread into lower latitudes,
and organisms that prefer warmer conditions become extinct or are squeezed into lower latitudes. This evidence is
also dicult to interpret because it requires (1) sequences
of sediments covering a long period of time, over a wide
range of latitudes and which are easily correlated; (2) ancient organisms which survive for several million years
without change and whose temperature preferences are
easily diagnosed; and (3) the nding of the relevant fossils.
Despite the diculties, analysis of ice core and ocean
sediment cores[30] has shown periods of glacials and interglacials over the past few million years. These also
conrm the linkage between ice ages and continental
crust phenomena such as glacial moraines, drumlins, and
glacial erratics. Hence the continental crust phenomena
are accepted as good evidence of earlier ice ages when
they are found in layers created much earlier than the time
range for which ice cores and ocean sediment cores are
available.

1.3 Major ice ages

There have been at least ve major ice ages in the earths
past (the Huronian, Cryogenian, Andean-Saharan, Karoo
Ice Age and the Quaternary glaciation). Outside these
ages, the Earth seems to have been ice-free even in high
latitudes.[31][32]
Rocks from the earliest well established ice age, called
the Huronian, formed around 2.4 to 2.1 Ga (billion years)
ago during the early Proterozoic Eon. Several hundreds
of km of the Huronian Supergroup are exposed 10
100 km north of the north shore of Lake Huron ex-

Ice age map of northern Germany and its northern neighbours.

Red: maximum limit of Weichselian glacial; yellow: Saale
glacial at maximum (Drenthe stage); blue: Elster glacial maximum glaciation.

Timeline of glaciations, shown in blue.

tending from near Sault Ste. Marie to Sudbury, northeast of Lake Huron, with giant layers of now-lithied till
beds, dropstones, varves, outwash, and scoured basement
rocks. Correlative Huronian deposits have been found
near Marquette, Michigan, and correlation has been made
with Paleoproterozoic glacial deposits from Western Australia.
The next well-documented ice age, and probably the most
severe of the last billion years, occurred from 850 to 630
million years ago (the Cryogenian period) and may have
produced a Snowball Earth in which glacial ice sheets
reached the equator,[33] possibly being ended by the accumulation of greenhouse gases such as CO
2 produced by volcanoes. The presence of ice on the
continents and pack ice on the oceans would inhibit both
silicate weathering and photosynthesis, which are the two
major sinks for CO
2 at present.[34] It has been suggested that the end of this
ice age was responsible for the subsequent Ediacaran and
Cambrian Explosion, though this model is recent and controversial.
The Andean-Saharan occurred from 460 to 420 million
years ago, during the Late Ordovician and the Silurian
period.
The evolution of land plants at the onset of the Devonian
period caused a long term increase in planetary oxygen
levels and reduction of CO
2 levels, which resulted in the Karoo Ice Age. It is named
after the glacial tills found in the Karoo region of South
Africa, where evidence for this ice age was rst clearly
identied. There were extensive polar ice caps at intervals

CHAPTER 1. ICE AGE

from 360 to 260 million years ago in South Africa during

the Carboniferous and early Permian Periods. Correlatives are known from Argentina, also in the center of the
ancient supercontinent Gondwanaland.
100 kyr cycle

2.5

-2
-4
-6
-8

Five Million Years of

Climate Change
From Sediment Cores

3
3.5
4

18

41 kyr cycle

O Benthic
Carbonate (per mil)

Equivalent
Vostok T (C)

4.5
Millions of Years Ago

Sediment records showing the uctuating sequences of glacials

and interglacials during the last several million years.

Shows the pattern of temperature and ice volume changes associated with recent glacials and interglacials

The current ice age, the Pliocene-Quaternary glaciation, started about 2.58 million years ago during the late
Pliocene, when the spread of ice sheets in the Northern
Hemisphere began. Since then, the world has seen cycles of glaciation with ice sheets advancing and retreating
on 40,000- and 100,000-year time scales called glacial
periods, glacials or glacial advances, and interglacial periods, interglacials or glacial retreats. The earth is currently in an interglacial, and the last glacial period ended
about 10,000 years ago. All that remains of the continental ice sheets are the Greenland and Antarctic ice sheets
and smaller glaciers such as on Ban Island.
Ice ages can be further divided by location and time;
for example, the names Riss (180,000130,000 years bp)
and Wrm (70,00010,000 years bp) refer specically to
glaciation in the Alpine region. The maximum extent of
the ice is not maintained for the full interval. The scouring action of each glaciation tends to remove most of the
evidence of prior ice sheets almost completely, except in
regions where the later sheet does not achieve full coverage.
Minimum (interglacial, black) and maximum (glacial, grey)
glaciation of the northern hemisphere

1.4 Glacials and interglacials

See also: Glacial period and Interglacial
Within the ice ages (or at least within the current one),
more temperate and more severe periods occur. The changes in Earths orbit called Milankovitch cycles.
colder periods are called glacial periods, the warmer pe- The earth has been in an interglacial period known as
riods interglacials, such as the Eemian Stage.
the Holocene for more than 11,000 years. It was conGlacials are characterized by cooler and drier climates ventional wisdom that the typical interglacial period lasts
over most of the earth and large land and sea ice masses about 12,000 years, but this has been called into quesextending outward from the poles. Mountain glaciers in tion recently. For example, an article in Nature[35] arotherwise unglaciated areas extend to lower elevations gues that the current interglacial might be most analodue to a lower snow line. Sea levels drop due to the re- gous to a previous interglacial that lasted 28,000 years.
moval of large volumes of water above sea level in the Predicted changes in orbital forcing suggest that the next
icecaps. There is evidence that ocean circulation patterns glacial period would begin at least 50,000 years from now,
are disrupted by glaciations. Since the earth has signi- even in absence of human-made global warming[36] (see
cant continental glaciation in the Arctic and Antarctic, we Milankovitch cycles). Moreover, anthropogenic forcare currently in a glacial minimum of a glaciation. Such ing from increased greenhouse gases might outweigh ora period between glacial maxima is known as an inter- bital forcing for as long as intensive use of fossil fuels
glacial. The glacials and interglacials also coincided with continues.[37]

1.6. CAUSES OF ICE AGES

5
rent projected consequences of global warming include
a largely ice-free Arctic Ocean within 520 years, see
Arctic shrinkage.) Additional fresh water owing into
the North Atlantic during a warming cycle may also reduce the global ocean water circulation (see Shutdown of
thermohaline circulation). Such a reduction (by reducing the eects of the Gulf Stream) would have a cooling
eect on northern Europe, which in turn would lead to
increased low-latitude snow retention during the summer.
It has also been suggested that during an extensive glacial,
glaciers may move through the Gulf of Saint Lawrence,
extending into the North Atlantic ocean far enough to
block the Gulf Stream.

1.5.2 Negative feedback processes

Minimum (interglacial, black) and maximum (glacial, grey)

glaciation of the southern hemisphere

1.5 Positive and negative feedback

in glacial periods

Ice sheets that form during glaciations cause erosion of

the land beneath them. After some time, this will reduce land above sea level and thus diminish the amount
of space on which ice sheets can form. This mitigates the
albedo feedback, as does the lowering in sea level that
accompanies the formation of ice sheets.

Another factor is the increased aridity occurring with

glacial maxima, which reduces the precipitation available
to maintain glaciation. The glacial retreat induced by this
Each glacial period is subject to positive feedback which or any other process can be amplied by similar inverse
makes it more severe and negative feedback which miti- positive feedbacks as for glacial advances.
gates and (in all cases so far) eventually ends it.
According to research published in Nature Geoscience,
human emissions of carbon dioxide will defer the next ice
age. Researchers used data on Earths orbit to nd the his1.5.1 Positive feedback processes
torical warm interglacial period that looks most like the
current one and from this have predicted that the next ice
Ice and snow increase Earths albedo, i.e. they make it
age would usually begin within 1,500 years. They go on
reect more of the suns energy and absorb less. Hence,
to say that emissions have been so high that it will not.[39]
when the air temperature decreases, ice and snow elds
grow, and this continues until competition with a negative
feedback mechanism forces the system to an equilibrium.
Also, the reduction in forests caused by the ices expan- 1.6 Causes of ice ages
sion increases albedo.
Another theory proposed by Ewing and Donn in 1956[38] The causes of ice ages are not fully understood for either
hypothesized that an ice-free Arctic Ocean leads to in- the large-scale ice age periods or the smaller ebb and ow
creased snowfall at high latitudes. When low-temperature of glacialinterglacial periods within an ice age. The conice covers the Arctic Ocean there is little evaporation or sensus is that several factors are important: atmospheric
sublimation and the polar regions are quite dry in terms composition, such as the concentrations of carbon dioxof precipitation, comparable to the amount found in ide and methane (the specic levels of the previously
mid-latitude deserts. This low precipitation allows high- mentioned gases are now able to be seen with the new
Dome C in Antarctica over
latitude snowfalls to melt during the summer. An ice- ice core samples from EPICA
[40]
);
changes
in the earths orbit
the
past
800,000
years
free Arctic Ocean absorbs solar radiation during the long
around
the
Sun
known
as
Milankovitch
cycles; the mosummer days, and evaporates more water into the Arctic
tectonic
plates
resulting
in
changes
in the relation
of
atmosphere. With higher precipitation, portions of this
tive
location
and
amount
of
continental
and
oceanic
crust
snow may not melt during the summer and so glacial ice
ocean
curon
the
earths
surface,
which
aect
wind
and
can form at lower altitudes and more southerly latitudes,
rents;
variations
in
solar
output;
the
orbital
dynamics
reducing the temperatures over land by increased albedo
as noted above. Furthermore, under this hypothesis the of the Earth-Moon system; and the impact of relatively
lack of oceanic pack ice allows increased exchange of wa- large meteorites, and volcanism including eruptions of
ters between the Arctic and the North Atlantic Oceans, supervolcanoes.
warming the Arctic and cooling the North Atlantic. (Cur- Some of these factors inuence each other. For exam-

CHAPTER 1. ICE AGE

ple, changes in Earths atmospheric composition (especially the concentrations of greenhouse gases) may alter
the climate, while climate change itself can change the
atmospheric composition (for example by changing the
rate at which weathering removes CO
2).

to have a signicant global impact on the earths climate

and ecosystems, did not begin in the 18th century with the
advent of the Industrial Era, but dates back to 8,000 years
ago, due to intense farming activities of our early agrarian
ancestors. It was at that time that atmospheric greenhouse
gas concentrations stopped following the periodic pattern
Maureen Raymo, William Ruddiman and others propose of the Milankovitch cycles. In his overdue-glaciation hythat the Tibetan and Colorado Plateaus are immense CO pothesis Ruddiman states that an incipient glacial would
probably have begun several thousand years ago, but the
2 scrubbers with a capacity to remove enough CO
glacial was forestalled by the ac2 from the global atmosphere to be a signicant causal arrival of that scheduled[45]
tivities of early farmers.
factor of the 40 million year Cenozoic Cooling trend.
They further claim that approximately half of their up- At a meeting of the American Geophysical Union (Delift (and CO
cember 17, 2008), scientists detailed evidence in sup2 scrubbing capacity) occurred in the past 10 million port of the controversial idea that the introduction of
years.[41][42]
large-scale rice agriculture in Asia, coupled with extensive deforestation in Europe began to alter world climate
by pumping signicant amounts of greenhouse gases
into the atmosphere over the last 1,000 years. In turn,
1.6.1 Changes in Earths atmosphere
a warmer atmosphere heated the oceans making them
much less ecient storehouses of carbon dioxide and reThere is considerable evidence that over the very recent
inforcing global warming, possibly forestalling the onset
period of the last 1001000 years, the sharp increases
of a new glacial age.[46]
in human activity, especially the burning of fossil fuels,
has caused the parallel sharp and accelerating increase in
atmospheric greenhouse gases which trap the suns heat.
1.6.2 Position of the continents
The consensus theory of the scientic community is that
the resulting greenhouse eect is a principal cause of
The geological record appears to show that ice ages start
the increase in global warming which has occurred over
when the continents are in positions which block or rethe same period, and a chief contributor to the accelerduce the ow of warm water from the equator to the poles
ated melting of the remaining glaciers and polar ice. A
and thus allow ice sheets to form. The ice sheets increase
2012 investigation nds that dinosaurs released methane
Earths reectivity and thus reduce the absorption of solar
through digestion in a similar amount to humanitys curradiation. With less radiation absorbed the atmosphere
rent methane release, which could have been a key faccools; the cooling allows the ice sheets to grow, which
[43]
tor to the very warm climate 150 million years ago.
further increases reectivity in a positive feedback loop.
There is evidence that greenhouse gas levels fell at the The ice age continues until the reduction in weathering
start of ice ages and rose during the retreat of the ice causes an increase in the greenhouse eect.
sheets, but it is dicult to establish cause and eect (see
There are three known congurations of the continents
the notes above on the role of weathering). Greenhouse
which block or reduce the ow of warm water from the
gas levels may also have been aected by other factors
equator to the poles:
which have been proposed as causes of ice ages, such as
the movement of continents and volcanism.
A continent sits on top of a pole, as Antarctica does
The Snowball Earth hypothesis maintains that the severe
today.
freezing in the late Proterozoic was ended by an increase
in CO
A polar sea is almost land-locked, as the Arctic
2 levels in the atmosphere, and some supporters of SnowOcean is today.
ball Earth argue that it was caused by a reduction in atmospheric CO
A supercontinent covers most of the equator, as
2. The hypothesis also warns of future Snowball Earths.
Rodinia did during the Cryogenian period.
In 2009, further evidence was provided that changes in
solar insolation provide the initial trigger for the earth to
warm after an Ice Age, with secondary factors like increases in greenhouse gases accounting for the magnitude
of the change.[44]

Since todays Earth has a continent over the South Pole

and an almost land-locked ocean over the North Pole,
geologists believe that Earth will continue to experience
glacial periods in the geologically near future.

William Ruddiman has proposed the early anthropocene

hypothesis, according to which the anthropocene era, as
some people call the most recent period in the earths history when the activities of the human species rst began

Some scientists believe that the Himalayas are a major

factor in the current ice age, because these mountains
have increased Earths total rainfall and therefore the rate
at which carbon dioxide is washed out of the atmosphere,

1.6. CAUSES OF ICE AGES

decreasing the greenhouse eect.[42] The Himalayas formation started about 70 million years ago when the IndoAustralian Plate collided with the Eurasian Plate, and the
Himalayas are still rising by about 5 mm per year because
the Indo-Australian plate is still moving at 67 mm/year.
The history of the Himalayas broadly ts the long-term
decrease in Earths average temperature since the midEocene, 40 million years ago.

1.6.3

Fluctuations in ocean currents

Another important contribution to ancient climate

regimes is the variation of ocean currents, which are
modied by continent position, sea levels and salinity,
as well as other factors. They have the ability to cool
(e.g. aiding the creation of Antarctic ice) and the ability to warm (e.g. giving the British Isles a temperate as
opposed to a boreal climate). The closing of the Isthmus
of Panama about 3 million years ago may have ushered in
the present period of strong glaciation over North America by ending the exchange of water between the tropical
Atlantic and Pacic Oceans.[47]
Analyses suggest that ocean current uctuations can adequately account for recent glacial oscillations. During the
last glacial period the sea-level has uctuated 2030 m as
water was sequestered, primarily in the northern hemisphere ice sheets. When ice collected and the sea level
dropped suciently, ow through the Bering Strait (the
narrow strait between Siberia and Alaska is ~50 m deep
today) was reduced, resulting in increased ow from the
North Atlantic. This realigned the thermohaline circulation in the Atlantic, increasing heat transport into the
Arctic, which melted the polar ice accumulation and reduced other continental ice sheets. The release of water
raised sea levels again, restoring the ingress of colder water from the Pacic with an accompanying shift to northern hemisphere ice accumulation.[48]

1.6.4

Uplift of the Tibetan plateau and surrounding mountain areas above the
snowline

Matthias Kuhles geological theory of Ice Age development was suggested by the existence of an ice sheet covering the Tibetan plateau during the Ice Ages (Last Glacial
Maximum?). According to Kuhle, the plate-tectonic uplift of Tibet past the snow-line has led to a surface of c.
2,400,000 square kilometres (930,000 sq mi) changing
from bare land to ice with a 70% greater albedo. The reection of energy into space resulted in a global cooling,
triggering the Pleistocene Ice Age. Because this highland
is at a subtropical latitude, with 4 to 5 times the insolation
of high-latitude areas, what would be Earths strongest
heating surface has turned into a cooling surface.
Kuhle explains the interglacial periods by the 100,000-

7
year cycle of radiation changes due to variations in Earths
orbit. This comparatively insignicant warming, when
combined with the lowering of the Nordic inland ice areas
and Tibet due to the weight of the superimposed ice-load,
has led to the repeated complete thawing of the inland ice
areas.[49][50][51][52]

1.6.5 Variations in Earths orbit (Milankovitch cycles)

The Milankovitch cycles are a set of cyclic variations in
characteristics of the Earths orbit around the Sun. Each
cycle has a dierent length, so at some times their eects
reinforce each other and at other times they (partially)
cancel each other.

Past and future of daily average insolation at top of the atmosphere on the day of the summer solstice, at 65 N latitude.

There is strong evidence that the Milankovitch cycles

aect the occurrence of glacial and interglacial periods
within an ice age. The present ice age is the most studied and best understood, particularly the last 400,000
years, since this is the period covered by ice cores that
record atmospheric composition and proxies for temperature and ice volume. Within this period, the match of
glacial/interglacial frequencies to the Milankovi orbital
forcing periods is so close that orbital forcing is generally accepted. The combined eects of the changing distance to the Sun, the precession of the Earths axis, and
the changing tilt of the Earths axis redistribute the sunlight received by the Earth. Of particular importance are
changes in the tilt of the Earths axis, which aect the intensity of seasons. For example, the amount of solar inux in July at 65 degrees north latitude varies by as much
as 22% (from 450 W/m to 550 W/m). It is widely believed that ice sheets advance when summers become too
cool to melt all of the accumulated snowfall from the previous winter. Some workers believe that the strength of
the orbital forcing is too small to trigger glaciations, but
feedback mechanisms like CO
2 may explain this mismatch.
While Milankovitch forcing predicts that cyclic changes
in the Earths orbital elements can be expressed in the
glaciation record, additional explanations are necessary to
explain which cycles are observed to be most important
in the timing of glacialinterglacial periods. In particular, during the last 800,000 years, the dominant period
of glacialinterglacial oscillation has been 100,000 years,
which corresponds to changes in Earths orbital eccentricity and orbital inclination. Yet this is by far the weakest
of the three frequencies predicted by Milankovitch. Dur-

CHAPTER 1. ICE AGE

ing the period 3.00.8 million years ago, the dominant

which occurred during the coldest part of the Little
pattern of glaciation corresponded to the 41,000-year peIce Age.
riod of changes in Earths obliquity (tilt of the axis). The
reasons for dominance of one frequency versus another The long-term increase in the Suns output cannot be a
are poorly understood and an active area of current re- cause of ice ages.
search, but the answer probably relates to some form of
resonance in the Earths climate system.
The traditional Milankovitch explanation struggles to
explain the dominance of the 100,000-year cycle over the
last 8 cycles. Richard A. Muller, Gordon J. F. MacDonald,[53][54][55] and others have pointed out that those calculations are for a two-dimensional orbit of Earth but the
three-dimensional orbit also has a 100,000-year cycle of
orbital inclination. They proposed that these variations
in orbital inclination lead to variations in insolation, as
the Earth moves in and out of known dust bands in the
solar system. Although this is a dierent mechanism to
the traditional view, the predicted periods over the last
400,000 years are nearly the same. The Muller and MacDonald theory, in turn, has been challenged by Jose Antonio Rial.[56]

1.6.7 Volcanism

Volcanic eruptions may have contributed to the inception and/or the end of ice age periods. At times during the paleoclimate, carbon dioxide levels were two or
three times greater than today. Volcanoes and movements in continental plates contributed to high amounts
of CO2 in the atmosphere. Carbon dioxide from volcanoes probably contributed to periods with highest overall temperatures.[60] One suggested explanation of the
Paleocene-Eocene Thermal Maximum is that undersea
volcanoes released methane from clathrates and thus
caused a large and rapid increase in the greenhouse effect. There appears to be no geological evidence for such
eruptions at the right time, but this does not prove they
Another worker, William Ruddiman, has suggested did not happen.
a model that explains the 100,000-year cycle by the
modulating eect of eccentricity (weak 100,000-year cycle) on precession (26,000-year cycle) combined with 1.7 Recent glacial and interglacial
greenhouse gas feedbacks in the 41,000- and 26,000phases
year cycles. Yet another theory has been advanced by
Peter Huybers who argued that the 41,000-year cycle has
always been dominant, but that the Earth has entered
a mode of climate behavior where only the second or
third cycle triggers an ice age. This would imply that
the 100,000-year periodicity is really an illusion created
by averaging together cycles lasting 80,000 and 120,000
years.[57] This theory is consistent with a simple empirical
multi-state model proposed by Didier Paillard.[58] Paillard suggests that the late Pleistocene glacial cycles can be
seen as jumps between three quasi-stable climate states.
The jumps are induced by the orbital forcing, while in
the early Pleistocene the 41,000-year glacial cycles resulted from jumps between only two climate states. A
dynamical model explaining this behavior was proposed
by Peter Ditlevsen.[59] This is in support of the suggestion
that the late Pleistocene glacial cycles are not due to the Northern hemisphere glaciation during the last ice ages. The set
weak 100,000-year eccentricity cycle, but a non-linear re- up of 3 to 4 km thick ice sheets caused a sea level lowering of
sponse to mainly the 41,000-year obliquity cycle.
about 120 m.

1.6.6

Variations in the Suns energy output

Main article: Timeline of glaciation

There are at least two types of variation in the Suns en1.7.1

ergy output

Glacial stages in North America

In the very long term, astrophysicists believe that the The major glacial stages of the current ice age in North
Suns output increases by about 7% every one billion America are the Illinoian, Sangamonian and Wisconsin
stages. The use of the Nebraskan, Afton, Kansan,
(109 ) years.
and Yarmouthian (Yarmouth) stages to subdivide the ice
Shorter-term variations such as sunspot cycles, and age in North America have been discontinued by Qualonger episodes such as the Maunder minimum, ternary geologists and geomorphologists. These stages

1.8. EFFECTS OF GLACIATION

have all been merged into the Pre-Illinoian Stage in the 8.4C here was an increase in precipitation. Accord1980s.[61][62][63]
ingly, at glacial times the humid climatic belt that today
further to the S, was
During the most recent North American glaciation, dur- is situated several latitude degrees
[67][68]
shifted
much
further
to
the
N.
ing the latter part of the Wisconsin Stage (26,000 to
13,300 years ago), ice sheets extended to about 45 degrees north latitude. These sheets were 3 to 4 km
thick.[62]

1.8 Eects of glaciation

This Wisconsin glaciation left widespread impacts on

the North American landscape. The Great Lakes and
the Finger Lakes were carved by ice deepening old valleys. Most of the lakes in Minnesota and Wisconsin were
gouged out by glaciers and later lled with glacial meltwaters. The old Teays River drainage system was radically
altered and largely reshaped into the Ohio River drainage
system. Other rivers were dammed and diverted to new
channels, such as the Niagara, which formed a dramatic
waterfall and gorge, when the waterow encountered a
limestone escarpment. Another similar waterfall, at the
present Clark Reservation State Park near Syracuse, New
York, is now dry.
The area from Long Island to Nantucket was formed from
glacial till, and the plethora of lakes on the Canadian
Shield in northern Canada can be almost entirely attributed to the action of the ice. As the ice retreated and
the rock dust dried, winds carried the material hundreds
of miles, forming beds of loess many dozens of feet thick
in the Missouri Valley. Isostatic rebound continues to reshape the Great Lakes and other areas formerly under the
weight of the ice sheets.
The Driftless Zone, a portion of western and southwestern Wisconsin along with parts of adjacent Minnesota,
Scandinavia exhibits some of the typical eects of ice age glaciaIowa, and Illinois, was not covered by glaciers.
See also: Glacial history of Minnesota

tion such as fjords and lakes.

See also: Glacial landforms

Although the last glacial period ended more than 8,000
years ago, its eects can still be felt today. For example, the moving ice carved out the landscape in Canada
(See Canadian Arctic Archipelago), Greenland, northern Eurasia and Antarctica. The erratic boulders, till,
A specially interesting climatic change during glacial drumlins, eskers, fjords, kettle lakes, moraines, cirques,
times has taken place in the semi-arid Andes. Be- horns, etc., are typical features left behind by the glaciers.
side the expected cooling down in comparison with the
current climate, a signicant precipitation is concerned The weight of the ice sheets was so great that they dehere. So, researches in the presently semiarid subtropic formed the Earths crust and mantle. After the ice sheets
Aconcagua-massif (6,962 m) have shown an unexpect- melted, the ice-covered land rebounded. Due to the high
edly extensive glacial glaciation of the type ice stream viscosity of the Earths mantle, the ow of mantle rocks
network.[64][65][66][67][68] The connected valley glaciers which controls the rebound process is very slowat a rate
exceeding 100 km in length, owed down on the East- of about 1 cm/year near the center of rebound area today.
side of this section of the Andes at 3234S and 69 During glaciation, water was taken from the oceans to
71W as far as a height of 2,060 m and on the western form the ice at high latitudes, thus global sea level
lu-side still clearly deeper.[68][69] Where current glaciers dropped by about 110 meters, exposing the continental
scarcely reach 10 km in length, the snowline (ELA) runs shelves and forming land-bridges between land-masses
at a height of 4,600 m and at that time was lowered to for animals to migrate. During deglaciation, the melted
3,200 m asl, i.e. about 1,400 m. From this follows that ice-water returned to the oceans, causing sea level to rise.
beside of an annual depression of temperature about c. This process can cause sudden shifts in coastlines and

1.7.2

Last Glacial Period in the semiarid Andes around Aconcagua and

Tupungato

10
hydration systems resulting in newly submerged lands,
emerging lands, collapsed ice dams resulting in salination
of lakes, new ice dams creating vast areas of freshwater,
and a general alteration in regional weather patterns on
a large but temporary scale. It can even cause temporary reglaciation. This type of chaotic pattern of rapidly
changing land, ice, saltwater and freshwater has been proposed as the likely model for the Baltic and Scandinavian
regions, as well as much of central North America at the
end of the last glacial maximum, with the present-day
coastlines only being achieved in the last few millennia
of prehistory. Also, the eect of elevation on Scandinavia submerged a vast continental plain that had existed
under much of what is now the North Sea, connecting the
British Isles to Continental Europe.[70]
The redistribution of ice-water on the surface of the
Earth and the ow of mantle rocks causes changes in the
gravitational eld as well as changes to the distribution
of the moment of inertia of the Earth. These changes to
the moment of inertia result in a change in the angular
velocity, axis, and wobble of the Earths rotation.

CHAPTER 1. ICE AGE

[2] Gribbin, J.R. (1982). Future Weather: Carbon Dioxide, Climate and the Greenhouse Eect. Penguin. ISBN
0140224599.
[3] Rmis, F.; Testus, L.; Testut (2006). Mais comment scoule donc un glacier ?
Aperu historique (PDF). C. R. Geoscience (in French) 338
(5):
368385.
Bibcode:2006CRGeo.338..368R.
doi:10.1016/j.crte.2006.02.004. Note: p. 374
[4] Montgomery 2010
[5] Martel, Pierre (1898). Appendix: Martel, P. (1744) An
account of the glacieres or ice alps in Savoy, in two letters,
one from an English gentleman to his friend at Geneva ;
the other from Pierre Martel , engineer, to the said English
gentleman. In Mathews, C.E. The annals of Mont Blanc.
London: Unwin. p. 327. See (Montgomery 2010) for a
full bibliography
[6] Krger, Tobias (2013). Discovering the Ice Ages. International Reception and Consequences for a Historical Understanding of Climate (German editon: Basel 2008). Leiden.
p. 47. ISBN 978-90-04-24169-5.

The weight of the redistributed surface mass loaded the [7] Krger 2013, pp. 78-83
lithosphere, caused it to ex and also induced stress within
the Earth. The presence of the glaciers generally sup- [8] Krger 2013, p. 150
pressed the movement of faults below.[71][72][73] How- [9] Krger 2013, pp. 83, 151
ever, during deglaciation, the faults experience accelerated slip triggering earthquakes. Earthquakes triggered [10] Goethe, Johann Wolfgang von: Geologische Probleme
und Versuch ihrer Ausung, Mineralogie und Geologie
near the ice margin may in turn accelerate ice calving and
in Goethes Werke, Weimar 1892, ISBN 3-423-05946-X,
[74]
may account for the Heinrich events. As more ice is rebook 73 (WA II,9), p. 253, 254.
moved near the ice margin, more intraplate earthquakes
are induced and this positive feedback may explain the [11] Krger 2013, p. 83
fast collapse of ice sheets.
In Europe, glacial erosion and isostatic sinking from
weight of ice made the Baltic Sea, which before the Ice
Age was all land drained by the Eridanos River.

1.9 See also

Global cooling
International Union for Quaternary Research
Irish Sea Glacier
Late Glacial Maximum
Little Ice Age
Post-glacial rebound
Timeline of glaciation

1.10 References
[1] Imbrie, J.; Imbrie, K.P (1979). Ice ages: solving the mystery. Short Hills NJ: Enslow Publishers. ISBN 978-089490-015-0.

[12] Krger 2013, p. 38

[13] Krger 2013, pp. 61-2
[14] Krger 2013, pp. 8890
[15] Krger 2013, pp. 91-6
[16] Andersen, Bjrn G. (1992). "Jens Esmarka pioneer in
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[17] Davies, Gordon L. (1969). The Earth in Decay. A History of British Geomorphology 15781878. London. pp.
267f.
Cunningham, Frank F. (1990). James David Forbes. Pioneer Scottish Glaciologist. Edinburgh: Scottish Academic
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[18] Krger 2013, pp. 142-47
[19] Krger 2013, pp. 10405
[20] Krger 2013, pp. 15053
[21] Krger 2013, pp. 15559
[22] Krger 2013, pp. 167-70
[23] Krger 2013, p. 173
[24] Krger 2008, pp. 17778

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1.11. EXTERNAL LINKS

1.11 External links

Cracking the Ice Age from PBS
Montgomery, Keith (2010). Development of the
glacial theory, 18001870. Historical Simulation
Raymo, M. (July 2011). Overview of the UpliftWeathering Hypothesis.
Eduard Y. Osipov ., Oleg M. Khlystov. Glaciers
and meltwater ux to Lake Baikal during the Last
Glacial Maximum.
Black, R. (9 January 2012). Carbon emissions 'will
defer Ice Age'". BBC News: Science and Environment.

13

Chapter 2

Megafauna
This article is about large land animals. For naked-eye
visible bottom-dwelling animals, see Macrobenthos. For
giant animals in mythology, see Megafauna (mythology).
In terrestrial zoology, megafauna (Ancient Greek megas

Eurasia, the Americas and Australia became extinct as

recently as 10,00040,000 years ago.[4] It is also commonly used for the largest extant wild land animals, especially elephants, giraes, hippopotamuses, rhinoceroses,
and large bovines. Megafauna may be subcategorized
by their trophic position into megaherbivores (e.g., elk),
megacarnivores (e.g., lions), and, more rarely, megaomnivores (e.g., bears).
Other common uses are for giant aquatic species, especially whales, any larger wild or domesticated land animals such as larger antelope and cattle, as well as numerous dinosaurs and other extinct giant reptilians.
The term is also sometimes applied to animals (usually
extinct) of great size relative to a more common or surviving type of the animal, for example the 1 m (3 ft)
dragonies of the Carboniferous period.

2.1 Ecological strategy

Megafauna in the sense of the largest mammals and
birds are generally K-strategists, with high longevity,
slow population growth rates, low mortality rates, and (at
least for the largest) few or no natural predators capable of
killing adults. These characteristics, although not exclusive to such megafauna, make them vulnerable to human
overexploitation, in part because of their slow population
recovery rates.
The African bush elephant, Earths largest living land animal

large + New Latin fauna animal) are large or giant

animals. The most common thresholds used are 45 kilograms (100 lb)[1][2] or 100 kilograms (220 lb).[2][3] This
thus includes many species not popularly thought of as
overly large, such as white-tailed deer, red kangaroo, and
humans.
In practice, the most common usage encountered in academic and popular writing describes land animals roughly
larger than a human that are not (solely) domesticated.
The term is especially associated with the Pleistocene
megafauna the land animals often larger than modern
counterparts considered archetypical of the last ice age,
such as mammoths, the majority of which in northern

2.2 Evolution of large body size

One observation that has been made about the evolution
of larger body size is that rapid rates of increase that are
often seen over relatively short time intervals are not sustainable over much longer time periods. In an examination of mammal body mass changes over time, the
maximum increase possible in a given time interval was
found to scale with the interval length raised to the 0.25
power.[5] This is thought to reect the emergence, during a trend of increasing maximum body size, of a series of anatomical, physiological, environmental, genetic
and other constraints that must be overcome by evolu-

14

2.2. EVOLUTION OF LARGE BODY SIZE

tionary innovations before further size increases are possible. A strikingly faster rate of change was found for
large decreases in body mass, such as may be associated
with the phenomenon of insular dwarsm. When normalized to generation length, the maximum rate of body
mass decrease was found to be over 30 times greater than
the maximum rate of body mass increase for a ten-fold
change.[5]

2.2.1

In terrestrial mammals

Subsequent to the CretaceousPaleogene extinction event

that eliminated the non-avian dinosaurs about 66 Ma ago,
terrestrial mammals underwent a nearly exponential increase in body size as they diversied to occupy the ecological niches left vacant.[6] Starting from just a few kg
before the event, maximum size had reached ~50 kg a
few million years later, and ~750 kg by the end of the
Paleocene. This trend of increasing body mass appears
to level o about 40 Ma ago (in the late Eocene), suggesting that physiological or ecological constraints had been
reached, after an increase in body mass of over three orders of magnitude.[6] However, when considered from the
standpoint of rate of size increase per generation, the exponential increase is found to have continued until the
appearance of Indricotherium 30 Ma ago. (Since generation time scales with body mass0.259 , increasing generation times with increasing size cause the log mass vs. time
plot to curve downward from a linear t.)[5]
Megaherbivores eventually attained a body mass of over
10 000 kg. The largest of these, indricotheres and
proboscids, have been hindgut fermenters, which are believed to have an advantage over foregut fermenters in
terms of being able to accelerate gastrointestinal transit in order to accommodate very large food intakes.[7]
A similar trend emerges when rates of increase of maximum body mass per generation for dierent mammalian clades are compared (using rates averaged over
macroevolutionary time scales). Among terrestrial mammals, the fastest rates of increase of body mass0.259 vs.
time (in Ma) occurred in perissodactyls (a slope of 2.1),
followed by rodents (1.2) and proboscids (1.1),[5] all of
which are hindgut fermenters. The rate of increase for
artiodactyls (0.74) was about a third that of perissodactyls. The rate for carnivorans (0.65) was slightly lower
yet, while primates, perhaps constrained by their arboreal
habits, had the lowest rate (0.39) among the mammalian
groups studied.[5]
Terrestrial mammalian carnivores from several eutherian
groups (the mesonychid Andrewsarchus, the creodonts
Megistotherium and Sarkastodon, and the carnivorans
Amphicyon and Arctodus) all reached a maximum size
of about 1000 kg[6] (the carnivoran Arctotherium apparently actually got somewhat larger). The largest known
metatherian carnivore, Proborhyaena gigantea, apparently reached 600 kg, also close to this limit.[8] A similar
theoretical maximum size for mammalian carnivores has

15
been predicted based on the metabolic rate of mammals,
the energetic cost of obtaining prey, and the maximum estimated rate coecient of prey intake.[9] It has also been
suggested that maximum size for mammalian carnivores
is constrained by the stress the humerus can withstand at
top running speed.[8]
Analysis of the variation of maximum body size over the
last 40 Ma suggests that decreasing temperature and increasing continental land area are associated with increasing maximum body size. The former correlation would
be consistent with Bergmanns rule,[10] and might be related to the thermoregulatory advantage of large body
mass in cool climates,[6] better ability of larger organisms to cope with seasonality in food supply,[10] or other
factors;[10] the latter correlation could be explainable in
terms of range and resource limitations.[6] However, the
two parameters are interrelated (due to sea level drops accompanying increased glaciation), making the driver of
the trends in maximum size more dicult to identify.[6]

2.2.2 In marine mammals

The ancestors of cetaceans are believed to have been the
semiaquatic pakicetids, no larger than wolves, of about
53 million years (Ma) ago.[11] By 40 Ma ago, cetaceans
had attained a length of 20 m or more in Basilosaurus,
an elongated, serpentine whale that diered from modern whales in many respects and was not ancestral to
them. Following this, the evolution of large body size
in cetaceans appears to have come to a temporary halt,
and then to have backtracked, although the available fossil
records are limited. However, in the period from 31 Ma
ago (in the Oligocene) to the present, cetaceans underwent a signicantly more rapid sustained increase in body
mass (a rate of increase in body mass0.259 of a factor of 3.2
per million years) than achieved by any group of terrestrial mammals.[5] This trend led to the largest animal of
all time, the modern blue whale. Several reasons for the
more rapid evolution of large body size in cetaceans are
possible. Fewer biomechanical constraints on increases
in body size may be associated with suspension in water
as opposed to standing against the force of gravity, and
with swimming movements as opposed to terrestrial locomotion. Also, the greater heat capacity and thermal
conductivity of water compared to air may increase the
thermoregulatory advantage of large body size in marine
endotherms, although diminishing returns apply.[5]
Cetaceans are not the only marine mammals to reach
unprecedented size in the modern era. The largest
carnivoran of all time is the mostly aquatic modern
southern elephant seal.

2.2.3 In ightless birds

Because of the small initial size of all mammals following the extinction of the dinosaurs, nonmammalian ver-

16

CHAPTER 2. MEGAFAUNA

tebrates had a roughly ten million year long window of

opportunity (during the Paleocene) for evolution of gigantism without much competition.[12] During this interval, apex predator niches were often occupied by reptiles,
such as terrestrial crocodilians (e.g. Pristichampsus),
large snakes (e.g. Titanoboa) or varanid lizards, or by
ightless birds[6] (e.g. Gastornis in Europe and North
America, Paleopsilopterus in South America). This is
also the period when ightless herbivorous paleognath
birds evolved to large size on a number of Gondwanan
land masses. These birds, termed ratites, have traditionally been viewed as representing a lineage separate
from that of their small ighted relatives, the Neotropic
tinamous. However, recent genetic studies have found
that tinamous nest well within the ratite tree, and are the
sister group of the extinct moa of New Zealand.[13][12][14]
Similarly, the small kiwi of New Zealand have been found
to be the sister group of the extinct elephant birds of
Madagascar.[12] These ndings indicate that ightlessness
and gigantism arose independently multiple times among
ratites via parallel evolution.
In the northern continents, large predatory birds were displaced when large eutherian carnivores evolved. In isolated South America, the phorusrhacids could not be outcompeted by the local metatherian sparassodonts and remained dominant until advanced eutherian predators arrived from North America (as part of the Great American Interchange) during the Pliocene. However, none
of the largest predatory (Brontornis), possibly omnivorous (Dromornis[15] ) or herbivorous (Aepyornis) ightless
birds of the Cenozoic ever grew to masses much above
500 kg, and thus never attained the size of the largest
mammalian carnivores, let alone that of the largest mammalian herbivores. It has been suggested that the increasing thickness of avian eggshells in proportion to egg mass
with increasing egg size places an upper limit on the size
of birds.[16][note 1] The largest species of Dromornis, D.
stirtoni, may have gone extinct after it attained the maximum avian body mass and was then outcompeted by
marsupial diprotodonts that evolved to sizes several times
larger.[19]

2.3 Megafaunal mass extinctions

2.3.1

Timing and possible causes

A well-known mass extinction of megafauna, the

Holocene extinction (see also Quaternary extinction
event), occurred at the end of the last ice age glacial period (a.k.a. the Wrm glaciation) and wiped out many
giant ice age animals, such as woolly mammoths, in
the Americas and northern Eurasia. Various theories
have attributed the wave of extinctions to human hunting, climate change, disease, a putative extraterrestrial
impact, or other causes. However, this extinction pulse
near the end of the Pleistocene was just one of a series

of megafaunal extinction pulses that have occurred during the last 50,000 years over much of the Earths surface,
with Africa and southern Asia being largely spared. The
latter areas did suer a gradual attrition of megafauna,
particularly of the slower-moving species (a class of vulnerable megafauna epitomized by giant tortoises), over
the last several million years.[20][21]
Outside the mainland of Afro-Eurasia, these megafaunal extinctions followed a highly distinctive landmass-bylandmass pattern that closely parallels the spread of humans into previously uninhabited regions of the world,
and which shows no correlation with climatic history
(which can be visualized with plots over recent geological time periods of climate markers such as marine oxygen isotopes or atmospheric carbon dioxide levels).[22][23]
Australia was struck rst around 45,000 years ago,[24]
followed by Tasmania about 41,000 years ago (after
formation of a land bridge to Australia about 43,000
years ago),[25][26][27] Japan apparently about 30,000 years
ago,[28] North America 13,000 years ago, South America about 500 years later,[29][30] Cyprus 10,000 years
ago,[31][32] the Antilles 6000 years ago,[33] New Caledonia[34] and nearby islands[35] 3000 years ago, Madagascar
2000 years ago,[36] New Zealand 700 years ago,[37] the
Mascarenes 400 years ago,[38] and the Commander Islands 250 years ago.[39] Nearly all of the worlds isolated islands could furnish similar examples of extinctions occurring shortly after the arrival of Homo sapiens, though most of these islands, such as the Hawaiian
Islands, never had terrestrial megafauna, so their extinct
fauna were smaller.[22][23]
An analysis of Sporormiella fungal spores (which derive mainly from the dung of megaherbivores) in swamp
sediment cores spanning the last 130,000 years from
Lynchs Crater in Queensland, Australia showed that
the megafauna of that region virtually disappeared about
41,000 years ago, at a time when climate changes were
minimal; the change was accompanied by an increase
in charcoal, and was followed by a transition from rainforest to re-tolerant sclerophyll vegetation. The highresolution chronology of the changes supports the hypothesis that human hunting alone eliminated the megafauna,
and that the subsequent change in ora was most likely
a consequence of the elimination of browsers and an increase in re.[40][41][42] The increase in re lagged the disappearance of megafauna by about a century, and most
likely resulted from accumulation of fuel once browsing stopped. Over the next several centuries grass increased; sclerophyll vegetation increased with a lag of
another century, and a sclerophyll forest developed after about another thousand years.[42] During two periods
of climate change about 120 and 75 thousand years ago,
sclerophyll vegetation had also increased at the site in response to a shift to cooler, drier conditions; neither of
these episodes had a signicant impact on megafaunal
abundance.[42] Similar conclusions regarding the culpability of human hunters in the disappearance of Pleis-

2.4. EXAMPLES

17

tocene megafauna were obtained via an analysis of a large

collection of eggshell fragments of the ightless Australian bird Genyornis newtoni[43] and from analysis of
Sporormiella fungal spores from a lake in eastern North
America.[44][45]

of the time (up to 10 C warmer than at present).[50][51]

This large emission follows from the enormous estimated
biomass of sauropods, and because methane production
of individual herbivores is believed to be almost proportional to their mass.[50]

Continuing human hunting and environmental disturbance has led to additional megafaunal extinctions in the
recent past, and has created a serious danger of further
extinctions in the near future (see examples below).

Recent studies have indicated that the extinction of

megafaunal herbivores may have caused a reduction
in atmospheric methane. This hypothesis is relatively
new.[52] One study examined the methane emissions from
the bison that occupied the Great Plains of North America before contact with European settlers. The study estimated that the removal of the bison caused a decrease
of as much as 2.2 million tons per year.[53] Another study
examined the change in the methane concentration in the
atmosphere at the end of the Pleistocene epoch after the
extinction of megafauna in the Americas. After early humans migrated to the Americas about 13,000 BP, their
hunting and other associated ecological impacts led to
the extinction of many megafaunal species there. Calculations suggest that this extinction decreased methane
production by about 9.6 million tons per year. This suggests that the absence of megafaunal methane emissions
may have contributed to the abrupt climatic cooling at the
onset of the Younger Dryas.[52] The decrease in atmospheric methane that occurred at that time, as recorded
in ice cores, was 2-4 times more rapid than any other decrease in the last half million years, suggesting that an
unusual mechanism was at work.[52]

A number of other mass extinctions occurred earlier in

Earths geologic history, in which some or all of the
megafauna of the time also died out. Famously, in the
CretaceousPaleogene extinction event the dinosaurs and
most other giant reptilians were eliminated. However, the
earlier mass extinctions were more global and not so selective for megafauna; i.e., many species of other types,
including plants, marine invertebrates[46] and plankton,
went extinct as well. Thus, the earlier events must have
been caused by more generalized types of disturbances to
the biosphere.

2.3.2

Consequences
megafauna

of

depletion

of

Eect on nutrient transport

Megafauna play a signicant role in the lateral transport of mineral nutrients in an ecosystem, tending to
translocate them from areas of high to those of lower
abundance. They do so by their movement between the
time they consume the nutrient and the time they release it through elimination (or, to a much lesser extent,
through decomposition after death).[47] In South Americas Amazon Basin, it is estimated that such lateral diffusion was reduced over 98% following the megafaunal
extinctions that occurred roughly 12,500 years ago.[48][49]
Given that phosphorus availability is thought to limit productivity in much of the region, the decrease in its transport from the western part of the basin and from oodplains (both of which derive their supply from the uplift
of the Andes) to other areas is thought to have signicantly impacted the regions ecology, and the eects may
not yet have reached their limits.[49]
Eect on methane emissions
Large populations of megaherbivores have the potential
to contribute greatly to the atmospheric concentration of
methane, which is an important greenhouse gas. Modern ruminant herbivores produce methane as a byproduct of foregut fermentation in digestion, and release it
through belching. Today, around 20% of annual methane
emissions come from livestock methane release. In the
Mesozoic, it has been estimated that sauropods could
have emitted 520 million tons of methane to the atmosphere annually,[50] contributing to the warmer climate

2.4 Examples
The following are some notable examples of animals often considered as megafauna (in the sense of the large
animal denition). This list is not intended to be exhaustive:
Clade Synapsida
Class Mammalia (phylogenetically, a clade
within Therapsida; see below)
Infraclass Metatheria
Order Diprotodontia
The red kangaroo (Macropus rufus) is the largest living Australian
mammal and marsupial at a weight
of up to 85 kg (187 lb). However, its extinct relative, the giant
short-faced kangaroo Procoptodon
goliah reached 230 kg (510 lb),
while extinct diprotodonts attained
the largest size of any marsupial in
history, up to an estimated 2,750 kg
(6,060 lb). The extinct marsupial
lion (Thylacleo carnifex), at up to
160 kg (350 lb) was much larger
than any extant carnivorous marsupial.

18

CHAPTER 2. MEGAFAUNA
Infraclass Eutheria
Superorder Afrotheria
Order Proboscidea
Elephants are the largest living
land animals. They and their relatives arose in Africa, but until
recently had a nearly worldwide
distribution. The African bush
elephant (Loxodonta africana) has
a shoulder height of up to 4.3
m (14 ft) and weighs up to 13
tons.
Among recently extinct
proboscideans, mammoths (Mammuthus) were close relatives of elephants, while mastodons (Mammut) were much more distantly
related. The steppe mammoth
(M. trogontherii) is estimated to
have commonly weighed around
10 tonnes, making it possibly the
largest proboscid, which would
make it the second largest land
mammal after indricotherines.
Order Sirenia
The largest sirenian at up to 1500
kg is the West Indian manatee
(Trichechus manatus). Stellers
sea cow (Hydrodamalis gigas) was
probably around ve times as massive, but was exterminated by humans within 27 years of its discovery o the remote Commander
Islands in 1741. In prehistoric
times this sea cow also lived along
the coasts of northeastern Asia
and northwestern North America;
it was apparently eliminated from
these more accessible locations by
aboriginal hunters.
Superorder Xenarthra
Order Cingulata
The glyptodonts were a group of
large, heavily armored ankylosaurlike xenarthrans related to living armadillos. They originated
in South America, invaded North
America during the Great American Interchange, and went extinct at the end of the Pleistocene
epoch.[54]
Order Pilosa
Ground sloths were another group
of slow, terrestrial xenarthrans,
related to modern tree sloths.
They had a similar history, although they reached North America earlier, and spread farther

north (e.g., Megalonyx).

The
largest genera, Megatherium and
Eremotherium, reached sizes comparable to elephants.[54]
Superorder Euarchontoglires
Order Primates
The largest living primate, at up
to 266 kg (586 lb), is the gorilla
(Gorilla beringei and Gorilla gorilla, with three of four subspecies being critically endangered). The extinct Malagasy
sloth lemur Archaeoindris reached
a similar size, while the extinct Gigantopithecus blacki of
Southeast Asia is believed to have
been several times larger. Some
populations of archaic Homo were
signicantly larger than recent
Homo sapiens;[55][56] for example,
Homo heidelbergensis in southern Africa may have commonly
reached 7 feet (2.1 m) in height,[57]
while Neanderthals were about
30% more massive.[58]
Order Rodentia
The extant capybara (Hydrochoerus hydrochaeris) of South
America, the largest living rodent,
weighs up to 65 kg (143 lb).
Several recently extinct North
American forms were larger: the
capybara Neochoerus pinckneyi
(another neotropic migrant) was
about 40% heavier; the giant
beaver (Castoroides ohioensis) was
similar. The extinct blunt-toothed
giant hutia (Amblyrhiza inundata)
of several Caribbean islands may
have been larger still. However,
several million years ago South
America harbored much more
massive rodents. Phoberomys pattersoni, known from a nearly full
skeleton, probably reached 700 kg
(1,500 lb). Fragmentary remains
suggest that Josephoartigasia
monesi grew to upwards of 1,000
kg (2,200 lb).
Superorder Laurasiatheria
Order Carnivora
Big cats include the tiger (Panthera tigris) and lion (Panthera
leo). The largest subspecies, at up
to 306 kg (675 lb), is the Siberian
tiger (P. tigris altaica), in accord
with Bergmanns rule. Members

2.4. EXAMPLES

19
of Panthera are distinguished by
morphological features which enable them to roar. Larger extinct felids include the American
lion (Panthera leo atrox) and the
South American saber-toothed cat
Smilodon populator.
Bears are large carnivorans of the
caniform suborder. The largest
living forms are the polar bear (Ursus maritimus), with a body weight
of up to 680 kg (1,500 lb), and
the similarly sized Kodiak bear
(Ursus arctos middendor), again
consistent with Bergmanns rule.
Arctotherium augustans, an extinct short-faced bear from South
America, was the largest predatory
land mammal ever with an estimated average weight of 1,600 kg
(3,500 lb).[59]
Seals, sea lions, and walruses
are amphibious marine carnivorans that evolved from bearlike
ancestors.
The southern elephant seal (Mirounga leonina) of
Antarctic and subantarctic waters
is the largest carnivoran of all time,
with bull males reaching a maximum length of 67 m (2023 ft)
and maximum weight of 5,000 kg
(11,000 lb).
Order Perissodactyla
Tapirs are browsing animals, with
a short prehensile snout and piglike form that appears to have
changed little in 20 million years.
They inhabit tropical forests of
Southeast Asia and South and
Central America, and include the
largest surviving land animals of
the latter two regions. There are
four species.

Rhinoceroses are odd-toed ungulates with horns made of keratin,

the same type of protein composing hair. They are among the
largest living land mammals after elephants (hippos attain a similar size). Three of ve extant
species are critically endangered.
Their extinct central Asian relatives the indricotherines were the
largest terrestrial mammals of all
time.
Order Artiodactyla (or cladistically,
Cetartiodactyla)
Giraes (Giraa camelopardalis)
are the tallest living land animals,
reaching heights of up to nearly 6
m (20 ft).
Bovine ungulates include the
largest surviving land animals of
Europe and North America. The
water bualo (Bubalis arnee),
bison (Bison bison and B. bonasus), and gaur (Bos gaurus) can all
grow to weights of over 900 kg
(2,000 lb).
The semiaquatic hippopotamus
(Hippopotamus amphibius) is the
heaviest living even-toed ungulate;
it and the critically endangered
pygmy hippo (Choeropsis liberiensis) are believed to be the closest
extant relatives of cetaceans.
Hippos are among the megafaunal species most dangerous to
humans.[60]
Order Cetacea (or cladistically,
Cetartiodactyla)
Whales, dolphins, and porpoises
are marine mammals. The blue
whale (Balaenoptera musculus) is
the largest baleen whale and the
largest animal that has ever lived,
at 30 metres (98 ft)[61] in length
and 170 tonnes (190 short tons)[62]
or more in weight. The sperm
whale (Physeter macrocephalus) is
the largest toothed whale, as well
as the planets loudest and brainiest
animal (with a brain about ve
times as massive as a humans).
The killer whale (Orcinus orca) is
the largest dolphin.
Order Pelycosauria (traditional; paraphyletic)

Rhinoceros, from Drers woodcut

Cotylorhynchus was a large, big-clawed,

herbivorous caseid of Early Permian

20

CHAPTER 2. MEGAFAUNA
North America, reaching 6 m (20 ft) and
2 tonnes.
Order Therapsida
Anteosaurus was a headbutting, semiaquatic, carnivorous dinocephalian of
Middle Permian South Africa. It reached
56 m (1620 ft) long, and weighed about
500600 kg (1,1001,300 lb).[63]

Clade Sauropsida
Class Aves (phylogenetically, a clade within
Coelurosauria, a taxon within the order
Saurischia; see below)
Order Struthioniformes
The ratites are an ancient and
diverse group of ightless birds
that are found on fragments of the
former supercontinent Gondwana.
The largest living bird, the ostrich
(Struthio camelus) was surpassed by
the extinct Aepyornis of Madagascar,
the heaviest of the group (400 kg
(880 lb)), and the extinct giant moa
(Dinornis) of New Zealand, the
tallest, growing to heights of 3.4 m
(11 ft). The latter two are examples
of island gigantism.
Order Anseriformes
Extinct dromornithids of Australia
such as Dromornis may have exceeded the largest ratites in size.
(Due to its small size for a continent
and its isolation, Australia is sometimes viewed as the worlds largest island; thus, these species could also be
considered insular giants.)
Class Reptilia (traditional; paraphyletic)
Order Crocodilia
Alligators and crocodiles are large
semiaquatic reptiles, the largest
of which, the saltwater crocodile
(Crocodylus porosus), can grow to
a weight of 1,360 kg (3,000 lb).
Crocodilians distant ancestors and
their kin, the crurotarsans, dominated the world in the late Triassic,
until the TriassicJurassic extinction
event allowed dinosaurs to overtake them. They remained diverse
during the later Mesozoic, when
crocodyliforms such as Deinosuchus
and Sarcosuchus reached lengths
of 12 m. Similarly large crocodilians, such as Mourasuchus and
Purussaurus, were present as recently as the Miocene in South
America.

Order Saurischia
Saurischian dinosaurs of the Jurassic
and Cretaceous include sauropods,
the longest (at up to 40 m or 130 ft)
and most massive terrestrial animals
known (Argentinosaurus reached 80
100 metric tonnes, or 90110 tons),
as well as theropods, the largest terrestrial carnivores (Spinosaurus grew
to 79 tonnes; the more famous
Tyrannosaurus, to 6.8 tonnes).
Order Squamata
While the largest extant lizard, the
Komodo dragon (Varanus komodoensis), another island giant, can
reach 3 m (10 ft) in length, its extinct
Australian relative Megalania may
have reached more than twice that
size. These monitor lizards' marine
relatives, the mosasaurs, were apex
predators in late Cretaceous seas.
The heaviest extant snake is considered to be the green anaconda (Eunectes murinus), while the reticulated
python (Python reticulatus), at up
to 8.7 m or more, is considered
the longest. An extinct Australian
Pliocene species of Liasis, the Blu
Downs giant python, reached 10 m,
while the Paleocene Titanoboa of
South America reached lengths of
1215 m and an estimated weight of
about 1135 kilograms (2500 lb).
Order Testudines
The largest turtle is the critically endangered marine leatherback turtle
(Dermochelys coriacea), weighing
up to 900 kg (2,000 lb). It is distinguished from other sea turtles by
its lack of a bony shell. The most
massive terrestrial chelonians are
the giant tortoises of the Galpagos
Islands (Chelonoidis nigra) and
Aldabra Atoll (Aldabrachelys gigantea), at up to 300 kg (660 lb). These
tortoises are the biggest survivors
of an assortment of giant tortoise
species that were widely present
on continental landmasses[64][65]
and additional islands[64] during the
Pleistocene.
Class Amphibia (in the wide, probably paraphyletic,
sense)
Order Temnospondyli (relationship to extant
amphibians is unclear)

2.5. GALLERY
The
Permian
temnospondyl
Prionosuchus, the largest amphibian
known, reached 9 m in length and
was an aquatic predator resembling a
crocodilian. After the appearance of
real crocodilians, temnospondyls such as
Koolasuchus (5 m long) had retreated to
the Antarctic region by the Cretaceous,
before going extinct.
Class Actinopterygii
Order Tetraodontiformes
The largest extant bony sh is the ocean
sunsh (Mola mola), whose average adult
weight is 1,000 kg (2,200 lb). While phylogenetically a bony sh, its skeleton is
primarily cartilage (which is lighter than
bone). It has a disk-shaped body, and propels itself with its long, thin dorsal and
anal ns; it feeds primarily on jellysh.
In these three respects (as well as in its
size and diving habits), it resembles a
leatherback turtle.
Order Acipenseriformes
The critically endangered beluga (European sturgeon, Huso huso) at up to
1,476 kg (3,254 lb) is the largest sturgeon
(which are also mostly cartilaginous) and
is considered the largest anadromous sh.
Order Siluriformes
The critically endangered Mekong giant
catsh (Pangasianodon gigas), at up to
293 kg (646 lb), is often viewed as the
largest freshwater sh.
Class Chondrichthyes
Order Lamniformes
The largest living predatory sh, the great
white shark (Carcharodon carcharias),
reaches weights up to 2,240 kg (4,940
lb). Its extinct relative C. megalodon (the
disputed genus being either Carcharodon
or Carcharocles) was more than an order
of magnitude larger, and is the largest
predatory shark or sh of all time (and
possibly the largest predator in vertebrate
history); it preyed on whales and other
marine mammals.
Order Orectolobiformes

21
Order Rajiformes
The manta ray (Manta birostris) is another
lter feeder and the largest ray, growing
to up to 2300 kg.
Class Placodermi
Order Arthrodira
The largest armored sh, Dunkleosteus,
arose during the late Devonian. At up
to 10 metres (33 ft) in length[66] and 3.6
tonnes (4.0 short tons) in mass,[67] it was a
hypercarnivorous apex predator that employed suction feeding.[68][69] Its contemporary, Titanichthys, apparently an early
lter feeder, rivaled it in size. The anthrodires were eliminated by the environmental upheavals of the Late Devonian
extinction, after existing for only about 50
million years.
Class Cephalopoda
Order Teuthida
A number of deep ocean creatures exhibit abyssal gigantism. These include
the giant squid (Architeuthis) and colossal
squid (Mesonychoteuthis hamiltoni); both
(although rarely seen) are believed to attain lengths of 12 m (39 ft) or more. The
latter is the worlds largest invertebrate,
and has the largest eyes of any animal.
Both are preyed upon by sperm whales.
Subphylum Chelicerata
Order Eurypterida
Eurypterids (sea scorpions) were a diverse group of aquatic and possibly amphibious predators that included the most
massive arthropods to have existed. They
survived over 200 million years, but nally died out in the PermianTriassic extinction event along with trilobites and
most other forms of life present at the
time, including most of the dominant terrestrial therapsids. The Early Devonian
Jaekelopterus reached an estimated length
of 2.5 m (8.2 ft), not including its
raptorial chelicerae, and is thought to
have been a freshwater species.

The largest extant shark, cartilaginous 2.5 Gallery

sh, and sh overall is the whale shark
(Rhincodon typus), which reaches weights 2.5.1 Extinct
in excess of 21.5 tonnes (47,000 pounds).
Some Paleozoic sea scorpions (Eurypterus shown)
Like baleen whales, it is a lter feeder and
primarily consumes plankton.
were larger than a man.

22

CHAPTER 2. MEGAFAUNA

Dunkleosteus was a 10 m (33 ft) long toothless 2.5.2 Living

armored predatory Devonian placoderm sh.
The gorilla is the largest and one of the most
endangered primates on the planet.
Sail-backed
pelycosaur
Dimetrodon
and
temnospondyl Eryops from North Americas
Siberian tigers are the biggest living cats, exempliPermian.
fying Bergmanns rule.
Pliosaur Pliosaurus (right) harassing the lter feeder
sh Leedsichthys during the Jurassic.

Polar bears, the largest bears and semiaquatic carnivores, are vulnerable to global warming.

Macronarian sauropods; from left, Camarasaurus,

Brachiosaurus, Giraatitan, Euhelopus.

The critically endangered black rhinoceros, up to 14

feet (4.3 m) long, is threatened by poaching.

Tyrannosaurus was a 12.3 m (40 ft) long theropod

dinosaur, an apex predator of west North America.
Indricotheres, the land mammals closest to
sauropods in size and lifestyle, were Asian rhinos.
The Late Miocene teratorn Argentavis of South
America had a 7 m (23 ft) wingspan.
C. megalodon (two possible sizes) with a whale
shark, great white shark and human for scale.
Deinotherium had downward-curving tusks and
ranged widely over Afro-Eurasia.
Titanis walleri, the only terror bird known to have
invaded North America, was 2.5 m (8 ft 2 in) tall.
Hippo-sized Diprotodon of Australia, the largest
marsupial of all time, went extinct 40,000 years ago.
Megalania, a giant carnivorous goanna of Australia,
might have grown to 7 metres long.

Wild Bactrian camels are critically endangered.

Their ancestors originated in North America.
Unlike woolly rhinos and mammoths, muskoxen
narrowly survived the Quaternary extinctions.[1]
Hippos, the heaviest and most aquatic even-toed ungulates, are whales' closest living relatives.
A lter feeder up to 33 m (108 ft) long, the blue
whale is the largest animal of all time.
The orca, the largest dolphin and pack predator, is
highly intelligent and lives in complex societies.
The ostrich is the largest ratite, the heaviest living bird and, at 70 km/h,[2] the fastest running
biped.[3][note 1]
The saltwater crocodile is the largest living reptile
and a dangerous predator of humans.
The Komodo dragon, an insular giant, is the largest
lizard and has infectious and venomous saliva.
The green anaconda, an aquatic constrictor, is the
heaviest snake, weighing up to 97.5 kg (215 lb).

Elephant-sized Megatherium, from South Americas

Pleistocene, was the largest sloth.[1]

The deep-diving ocean sunsh is the largest bony

sh, but its skeleton is mostly cartilaginous.

Toxodon, one of South Americas largest and last

notoungulates.[1] It had a relative in Mexico.

The Nile perch, one of the largest freshwater sh, is

also a damaging invasive species.[note 2]

American lions exceeded extant lions in size and

ranged over two continents until 10,000 BP.
Woolly mammoths vanished after humans invaded
their habitat in Eurasia and N. America.[2]
Haasts eagle, the largest eagle known, attacking
moa (which included the tallest bird known).
1. ^ Cite error:
The named reference
Fari.C3.B1a2013 was invoked but never dened
(see the help page).
2. ^ Cite error: The named reference Stuart was invoked but never dened (see the help page).

The whale shark is the largest extant shark or sh

species, growing up to 12.6 m (41 ft) in length.
The manta, a lter feeder, is the largest ray at up to
7.6 m across, yet can breach clear of the water.
Examination of a 9 m giant squid, an abyssal giant
and the second largest cephalopod.
1. ^ Cite error: The named reference Stuart was invoked but never dened (see the help page).
2. ^ Davies, S.J.J.F. (2003). Birds I Tinamous and
Ratites to Hoatzins. In Hutchins, Michael. Grzimeks Animal Life Encyclopedia 8 (2 ed.). Farmington Hills, MI: Gale Group. pp. 99101. ISBN
0-7876-5784-0.

2.8. REFERENCES
3. ^ Cite error: The named reference Stewart2006 was
invoked but never dened (see the help page).
4. ^ Penny, M. (2002). The Secret World of Kangaroos. Austin TX: Raintree Steck-Vaughn. ISBN 07398-4986-7.
Cite error: There are <ref group=note> tags on this
page, but the references will not show without a {{reist|group=note}} template (see the help page).

2.6 See also

Australian megafauna
Bergmanns rule
Charismatic megafauna
Copes rule
Deep-sea gigantism
Fauna
Giant animals in ction and mythology
Island dwarsm
Island gigantism
Largest organisms
Largest prehistoric organisms
List of megafauna discovered in modern times
Megafauna (categories)
Africa
Australia
Eurasia
North America
South America
New World Pleistocene extinctions
Pleistocene megafauna
Quaternary extinction event

2.7 Notes
[1] Nonavian dinosaur size was not similarly constrained because they had a dierent relationship between body mass
and egg size than birds. The 400 kg Aepyornis had larger
eggs than nearly all dinosaurs.[17][18]

23

2.8 References
[1] Stuart, A. J. (November 1991). Mammalian extinctions
in the Late Pleistocene of northern Eurasia and North
America. Biological Reviews (Wiley) 66 (4): 453562.
doi:10.1111/j.1469-185X.1991.tb01149.x.
[2] Johnson, C. N. (2002-09-23). Determinants of Loss of
Mammal Species during the Late Quaternary 'Megafauna'
Extinctions: Life History and Ecology, but Not Body
Size. Proceedings of the Royal Society of London B (The
Royal Society) 269 (1506): 22212227 (see p. 2225).
doi:10.1098/rspb.2002.2130. JSTOR 3558643.
[3] Martin, P. S.; Steadman, D. W. (1999-06-30).
Prehistoric extinctions on islands and continents.
In MacPhee, R. D. E. Extinctions in near time: causes,
contexts and consequences. Advances in Vertebrate
Paleontology 2. New York: Kluwer/Plenum. pp. 1756.
ISBN 978-0-306-46092-0. OCLC 41368299. Retrieved
2011-08-23.
[4] Ice Age Animals. Illinois State Museum
[5] Evans, A. R.; et al. (2012-01-30). The maximum rate of mammal evolution.
PNAS 109.
doi:10.1073/pnas.1120774109.
Retrieved 2011-0211.
[6] Smith, F. A.; Boyer, A. G.; Brown, J. H.; Costa, D. P.;
Dayan, T.; Ernest, S. K. M.; Evans, A. R.; Fortelius,
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24

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CHAPTER 2. MEGAFAUNA

2.9 External links

Megafauna First Victims of the Human-Caused
Extinction

Chapter 3

Pleistocene
before the present, as opposed to the currently accepted
2.588 million years BP: publications from the preceding
years may use either denition of the period.

3.1 Dating
Earth during the Pleistocene epoch.

The Pleistocene /plastsin/ (symbol PS[1] ) is the geological epoch which lasted from about 2,588,000 to
11,700 years ago, spanning the worlds recent period of
repeated glaciations.
Charles Lyell introduced this term in 1839 to describe
strata in Sicily that had at least 70% of their molluscan fauna still living today. This distinguished it
from the older Pliocene Epoch, which Lyell had originally thought to be the youngest fossil rock layer. He
constructed the name Pleistocene (Most New or
Newest) from the Greek , plestos, most,
and , kains (latinized as cnus), new";[2] this
contrasting with the immediately preceding Pleiocene
(More New or Newer, from , plen, more,
and kains; usual spelling: Pliocene), and the immediately subsequent Holocene (wholly new or entirely
new, from , hlos, whole, and kains) epoch,
which extends to the present time.

The Pleistocene has been dated from 2.588 million

(.005) to 11,700 years before present (BP), with the end
date expressed in radiocarbon years as 10,000 carbon-14
years BP.[4] It covers most of the latest period of repeated
glaciation, up to and including the Younger Dryas cold
spell. The end of the Younger Dryas has been dated to
about 9640 BC (11,654 calendar years BP). It was not
until after the development of radiocarbon dating, however, that Pleistocene archaeological excavations shifted
to stratied caves and rock-shelters as opposed to openair river-terrace sites.[5]
In 2009 the International Union of Geological Sciences
(IUGS) conrmed a change in time period for the Pleistocene, changing the start date from 1.806 to 2.588 million years BP, and accepted the base of the Gelasian as
the base of the Pleistocene, namely the base of the Monte
San Nicola GSSP.[6] The IUGS has yet to approve a type
section, Global Boundary Stratotype Section and Point
(GSSP), for the upper Pleistocene/Holocene boundary
(i.e. the upper boundary). The proposed section is the
North Greenland Ice Core Project ice core 75 06' N 42
18' W.[7] The lower boundary of the Pleistocene Series
is formally dened magnetostratigraphically as the base
of the Matuyama (C2r) chronozone, isotopic stage 103.
Above this point there are notable extinctions of the calcareous nanofossils: Discoaster pentaradiatus and Discoaster surculus.[8][9]

The Pleistocene is the rst epoch of the Quaternary

Period or sixth epoch of the Cenozoic Era.[3] The end of
the Pleistocene corresponds with the end of the last glacial
period. It also corresponds with the end of the Paleolithic
age used in archaeology. In the ICS timescale, the Pleistocene is divided into four stages or ages, the Gelasian,
Calabrian, Ionian and Tarantian. All of these stages were
dened in southern Europe. In addition to this international subdivision, various regional subdivisions are often The Pleistocene covers the recent period of repeated
used.
glaciations. The name Plio-Pleistocene has in the past
Before a change nally conrmed in 2009 by the been used to mean the last ice age. The revised deniInternational Union of Geological Sciences, the time tion of the Quaternary, by pushing back the start date of
boundary between the Pleistocene and the preceding the Pleistocene to 2.58 Ma, results in the inclusion of all
Pliocene was regarded as being at 1.806 million years the recent repeated glaciations within the Pleistocene.
27

28

CHAPTER 3. PLEISTOCENE
mean annual temperature at the edge of the ice was 6
C (21 F); at the edge of the permafrost, 0 C (32 F).
Each glacial advance tied up huge volumes of water in
continental ice sheets 1,500 to 3,000 metres (4,900
9,800 ft) thick, resulting in temporary sea-level drops of
100 metres (300 ft) or more over the entire surface of
the Earth. During interglacial times, such as at present,
drowned coastlines were common, mitigated by isostatic
or other emergent motion of some regions.
The eects of glaciation were global. Antarctica was icebound throughout the Pleistocene as well as the preceding
Pliocene. The Andes were covered in the south by the
Patagonian ice cap. There were glaciers in New Zealand
and Tasmania. The current decaying glaciers of Mount
Kenya, Mount Kilimanjaro, and the Ruwenzori Range in
east and central Africa were larger. Glaciers existed in
the mountains of Ethiopia and to the west in the Atlas
mountains.
In the northern hemisphere, many glaciers fused into one.
The Cordilleran ice sheet covered the North American
northwest; the east was covered by the Laurentide. The
Fenno-Scandian ice sheet rested on northern Europe, including Great Britain; the Alpine ice sheet on the Alps.
Scattered domes stretched across Siberia and the Arctic
shelf. The northern seas were ice-covered.

The maximum extent of glacial ice in the north polar area during
the Pleistocene period.

3.2 Paleogeography and climate

The modern continents were essentially at their present
positions during the Pleistocene, the plates upon which
they sit probably having moved no more than 100 km relative to each other since the beginning of the period.

South of the ice sheets large lakes accumulated because

outlets were blocked and the cooler air slowed evaporation. When the Laurentide ice sheet retreated, north central North America was totally covered by Lake Agassiz.
Over a hundred basins, now dry or nearly so, were overowing in the North American west. Lake Bonneville,
for example, stood where Great Salt Lake now does. In
Eurasia, large lakes developed as a result of the runo
from the glaciers. Rivers were larger, had a more copious
ow, and were braided. African lakes were fuller, apparently from decreased evaporation. Deserts on the other
hand were drier and more extensive. Rainfall was lower
because of the decrease in oceanic and other evaporation.

According to Mark Lynas (through collected data), the

Pleistocenes overall climate could be characterized as
3.2.2 Major events
a continuous El Nio with trade winds in the south
Pacic weakening or heading east, warm air rising near
Further information: Timeline of glaciation
Peru, warm water spreading from the west Pacic and
Over 11 major glacial events have been identied, as
the Indian Ocean to the east Pacic, and other El Nio
well as many minor glacial events.[11] A major glacial
[10]
markers.
event is a general glacial excursion, termed a glacial.
Glacials are separated by interglacials. During a glacial,
the glacier experiences minor advances and retreats. The
3.2.1 Glacial features
minor excursion is a stadial"; times between stadials are
Pleistocene climate was marked by repeated glacial cycles interstadials.
in which continental glaciers pushed to the 40th parallel
in some places. It is estimated that, at maximum glacial
extent, 30% of the Earths surface was covered by ice. In
addition, a zone of permafrost stretched southward from
the edge of the glacial sheet, a few hundred kilometres
in North America, and several hundred in Eurasia. The

These events are dened dierently in dierent regions

of the glacial range, which have their own glacial history
depending on latitude, terrain and climate. There is a general correspondence between glacials in dierent regions.
Investigators often interchange the names if the glacial
geology of a region is in the process of being dened.

3.2. PALEOGEOGRAPHY AND CLIMATE

29
comes from the underlying cyclical motions of the planet,
which eventually drag all the transients into harmony with
them. The repeated glaciations of the Pleistocene were
caused by the same factors.
Milankovitch cycles
Main article: Milankovitch cycles

Glaciation in the Pleistocene was a series of glacials and

interglacials, stadials and interstadials, mirroring periodic
changes in climate. The main factor at work in climate
Ice ages as reected in atmospheric CO2 , stored in bubbles from cycling is now believed to be Milankovitch cycles. These
are periodic variations in regional and planetary solar raglacial ice of Antarctica.
diation reaching the Earth caused by several repeating
changes in the Earths motion.
However, it is generally incorrect to apply the name of a
Milankovitch cycles cannot be the sole factor responsiglacial in one region to another.
ble for the variations in climate since they explain neither
For most of the 20th century only a few regions had been the long term cooling trend over the Plio-Pleistocene, nor
studied and the names were relatively few. Today the ge- the millennial variations in the Greenland Ice Cores. Miologists of dierent nations are taking more of an interest lankovitch pacing seems to best explain glaciation events
in Pleistocene glaciology. As a consequence, the num- with periodicity of 100,000, 40,000, and 20,000 years.
ber of names is expanding rapidly and will continue to Such a pattern seems to t the information on climate
expand. Many of the advances and stadials remain un- change found in oxygen isotope cores. The timing of
named. Also, the terrestrial evidence for some of them our present interglacial interval (known as the Holocene,
has been erased or obscured by larger ones, but evidence Postglacial, or the Present Interglacial) to that of the preremains from the study of cyclical climate changes.
vious interglacial, beginning about 130,000 years ago
The glacials in the following tables show historical us- (The Eemian Interglacial), suggests that the next glacial
ages, are a simplication of a much more complex cycle would likely begin in about 3,000 years.
of variation in climate and terrain, and are generally no
longer used. These names have been abandoned in favor
Oxygen isotope ratio cycles
of numeric data because many of the correlations were
found to be either inexact or incorrect and more than four
Main article: Oxygen isotope ratio cycle
major glacials have been recognized since the historical
terminology was established.[11][12][13]
In oxygen isotope ratio analysis, variations in the ratio of
Corresponding to the terms glacial and interglacial, the
18
terms pluvial and interpluvial are in use (Latin: pluvia,
O to 16
rain). A pluvial is a warmer period of increased rainfall;
O (two isotopes of oxygen) by mass (measured by a mass
an interpluvial, of decreased rainfall. Formerly a pluvial
spectrometer) present in the calcite of oceanic core samwas thought to correspond to a glacial in regions not iced,
ples is used as a diagnostic of ancient ocean temperature
and in some cases it does. Rainfall is cyclical also. Pluchange and therefore of climate change. Cold oceans are
vials and interpluvials are widespread.
richer in 18
There is no systematic correspondence of pluvials to O, which is included in the tests of the microorganisms
glacials, however. Moreover, regional pluvials do not cor- (foraminifera) contributing the calcite.
respond to each other globally. For example, some have
A more recent version of the sampling process makes use
used the term Riss pluvial in Egyptian contexts. Any
of modern glacial ice cores. Although less rich in 18
coincidence is an accident of regional factors. Only a few
O than sea water, the snow that fell on the glacier year by
of the names for pluvials in restricted regions have been
year nevertheless contained 18
strategraphically dened.
O and 16
O in a ratio that depended on the mean annual temperature.
3.2.3 Palaeocycles
Temperature and climate change are cyclical when plotThe sum of transient factors acting at the Earths surface ted on a graph of temperature versus time. Temperature
is cyclical: climate, ocean currents and other movements, coordinates are given in the form of a deviation from towind currents, temperature, etc. The waveform response days annual mean temperature, taken as zero. This sort

30

CHAPTER 3. PLEISTOCENE

of graph is based on another of isotope ratio versus time.

Ratios are converted to a percentage dierence from the
ratio found in standard mean ocean water (SMOW).
The graph in either form appears as a waveform with
overtones. One half of a period is a Marine isotopic stage
(MIS). It indicates a glacial (below zero) or an interglacial
(above zero). Overtones are stadials or interstadials.
According to this evidence, Earth experienced 102 MIS
stages beginning at about 2.588 Ma BP in the Early Pleistocene Gelasian. Early Pleistocene stages were shallow
and frequent. The latest were the most intense and most
widely spaced.
By convention, stages are numbered from the Holocene,
which is MIS1. Glacials receive an even number; interglacials, odd. The rst major glacial was MIS2-4 at about
8511 ka BP. The largest glacials were 2, 6, 12, and 16;
the warmest interglacials, 1, 5, 9 and 11. For matching
of MIS numbers to named stages, see under the articles
for those names.

Pleistocene of South America showing Megatherium and two

Glyptodon.

had replaced the megafauna and migrated north.

The extinctions were especially severe in North America
where native horses and camels were eliminated.

3.3 Fauna

Asian land mammal ages (ALMA) include

Zhoukoudianian, Nihewanian, and Yushean.

See also: Quaternary extinction event

European land mammal ages (ELMA) include

Gelasian (2.51.8 Ma).

Both marine and continental faunas were essentially modern and many animals, specically, mammals were much
larger in body form than their modern relatives .

North American land mammal ages (NALMA) include Blancan (4.751.8), Irvingtonian (1.80.24)
and Rancholabrean (0.240.01) in millions of years.
The Blancan extends signicantly back into the
Pliocene.
South American land mammal ages (SALMA) include Uquian (2.51.5), Ensenadan (1.50.3) and
Lujanian (0.30.01) in millions of years. The
Uquian previously extended signicantly back into
the Pliocene, although the new denition places it
entirely within the Pleistocene.

Pleistocene of Northern Spain showing woolly mammoth, cave

lions eating a reindeer, tarpans, and woolly rhinoceros.

The severe climatic changes during the ice age had major impacts on the fauna and ora. With each advance of
the ice, large areas of the continents became totally depopulated, and plants and animals retreating southward
in front of the advancing glacier faced tremendous stress.
The most severe stress resulted from drastic climatic
changes, reduced living space, and curtailed food supply.
A major extinction event of large mammals (megafauna),
which included mammoths, mastodons, saber-toothed
cats, glyptodons, ground sloths, Irish elk, cave bears, and
short-faced bears, began late in the Pleistocene and continued into the Holocene. Neanderthals also became extinct during this period. At the end of the last ice age,
cold-blooded animals, smaller mammals like wood mice,
migratory birds, and swifter animals like whitetail deer

3.4 Humans
Main articles: Human evolution, Paleolithic and Models
of migration to the New World
Scientic evidence[14] indicates that humans evolved into
their present form during the Pleistocene.[15] In the beginning of the Pleistocene Paranthropus species are still
present, as well as early human ancestors, but during the
lower Palaeolithic they disappeared, and the only hominid
species found in fossilic records is Homo erectus for much
of the Pleistocene. The Middle and late Palaeolithic saw
the appearance of new types of humans, as well as the
development of more elaborate tools than found in previous eras. According to mitochondrial timing techniques,
modern humans migrated from Africa after the Riss
glaciation in the middle Palaeolithic during the Eemian

3.7. REFERENCES
Stage, spreading all over the ice-free world during the late
Pleistocene.[16][17][18] A 2005 study posits that humans
in this migration interbred with archaic human forms already outside of Africa by the late Pleistocene, incorporating archaic human genetic material into the modern
human gene pool.[19]

3.5 Deposits

31

[7] Svensson, A., S. W. Nielsen, S. Kipfstuhl, S. J. Johnsen,

J. P. Steensen, M. Bigler, U. Ruth, and R. Rthlisberger
(2005) Visual stratigraphy of the North Greenland Ice
Core Project (NorthGRIP) ice core during the last glacial
period Journal of Geophysical Research 110: (D02108)
[8] Gradstein, Felix M.; Ogg, James G. and Smith, A. Gilbert
(eds.) (2005) A Geologic Time Scale 2004 Cambridge
University Press, Cambridge, UK, p. 28, ISBN 0-52178142-6
[9] Rio, D.; Sprovieri, R.; Castradori, D. and Di Stefano, E.

Pleistocene non-marine sediments are found primarily

(1998) The Gelasian Stage (Upper Pliocene): a new unit
in uvial deposits, lakebeds, slope and loess deposits as
of the global standard chronostratigraphic scale Episodes
well as in the large amounts of material moved about by
21: pp. 82-87
glaciers. Less common are cave deposits, travertines and
volcanic deposits (lavas, ashes). Pleistocene marine de- [10] National Geographic Channel, Six Degrees Could Change
posits are found primarily in shallow marine basins mostly
The World, Mark Lynas interview. Retrieved February
14, 2008.
(but with important exceptions) in areas within a few tens
of kilometers of the modern shoreline. In a few geologically active areas such as the Southern California coast, [11] Richmond, G.M. and D.S. Fullerton, 1986, Summation
of Quaternary glaciations in the United States of America.
Pleistocene marine deposits may be found at elevations
Quaternary Science Reviews. vol. 5, pp. 183-196.
of several hundred meters.

3.6 See also

Climate state
Geologic time scale
Pleistocene megafauna
Timeline of glaciation

3.7 References
[1] Geologic Age Symbol Font (StratagemAge)". USGS.
99-430. Retrieved 2011-06-22.
[2] Pleistocene. Online Etymology Dictionary.
[3] Gibbard, P. and van Kolfschoten, T. (2004) The Pleistocene and Holocene Epochs Chapter 22 PDF (3.1 MB)
In Gradstein, F. M., Ogg, James G., and Smith, A. Gilbert
(eds.), A Geologic Time Scale 2004 Cambridge University
Press, Cambridge, ISBN 0-521-78142-6
[4] For the top of the series, see: Lourens, L., Hilgen, F.,
Shackleton, N.J., Laskar, J., Wilson, D., (2004) The
Neogene Period. In: Gradstein, F., Ogg, J., Smith, A.G.
(Eds.), A Geologic Time Scale 2004. Cambridge: Cambridge University Press.

[12] Roy, M., P.U. Clark, R.W. Barendregt, J.R., Glasmann,

and R.J. Enkin, 2004, Glacial stratigraphy and paleomagnetism of late Cenozoic deposits of the north-central United
States, PDF version, 1.2 MB. Geological Society of America Bulletin.116(1-2): pp. 30-41; doi:10.1130/B25325.1
[13] Aber, J.S. (1991) Glaciations of Kansas Boreas 20(4):
pp. 297-314 - (contains a summary of how and why the
Nebraskan, Aftonian, Kansan, and Yarmouthian stages
were abandoned by modern stratigraphers).
[14] Rogers, A.R. and Jorde, L.B. (1995) Genetic evidence
on modern human origins Human Biology 67: pp. 136
[15] Wall, J.D. and Przeworski, M. (2000) When did the
human population start increasing?" Genetics 155: pp.
18651874
[16] Cann, R.L.; Stoneking, M. and Wilson, A.C.(1987) Mitochondrial DNA and human evolution Nature 325: pp.
3136
[17] Stringer, C.B. (1992) Evolution of early modern humans In: Jones, Steve; Martin, R. and Pilbeam, David
R. (eds.) (1992) The Cambridge encyclopedia of human
evolution Cambridge University Press, Cambridge, ISBN
0-521-32370-3, pp. 241251.
[18] Templeton, A. (2002) Out of Africa again and again Nature 416: p. 45

[5] Moore, Mark; Brumm (2007). Stone artifacts and hominins in island Southeast Asia: New insights from Flores, eastern Indonesia. Journal of Human Evolution 52:
88. Retrieved 10 April 2014.

[19] Eswarana, Vinayak; Harpendingb, Henry and Rogers,

Alan R. (2005) Genomics refutes an exclusively African
origin of humans Journal of Human Evolution 49(1): pp.
118 Abstract

[6] Riccardi, Alberto C. (30 June 2009) IUGS ratied ICS

Recommendation on redenition of Pleistocene and formal denition of base of Quaternary International Union
of Geological Sciences

Ogg, Jim; June, 2004, Overview of Global

Boundary Stratotype Sections and Points (GSSPs,
Stratigraphy.org, Accessed April 30, 2006.

32

CHAPTER 3. PLEISTOCENE

3.8 External links

The SMU-in-Taos Research Publications digital
collection of anthropological and archaeological
monographs contains Late Pleistocene environments of the southern high plains.
Pleistocene Microfossils:
Foraminifera

50+

images

of

Chapter 4

Prehistoric mammal
4.2 See also
Cynodont
List of extinct mammals
Mammaliaformes
Megamammals
Pleistocene extinctions
Pleistocene megafauna
Synapsid
Therapsid
An early drawing depicting prehistoric mammals

Prehistoric mammals are groups of mammals that became extinct before humans developed writing. 164
million years ago, in the Jurassic period, Castorocauda
lutrasimilis, a mammaliaform (mammal-shaped) animal
weighing about 500 grams (1.1 lb), had a full mammalian
pelt, with guard hairs and underfur, webbed feet, and
scales on the tail like a modern beaver, as well as teeth
specialized for catching sh.
Later, about 130 million years ago in the Cretaceous,
there existed larger mammals; a fossil of Repenomamus
giganticus indicates that the animal was about 1 meter (3 ft) long. In the stomach of a smaller cousin,
Repenomamus robustus at 52 cm (20 in), the remains
of a juvenile dinosaur have been preserved.
The lineages of many varieties continued through the
Cenozoic era where some reached very large sizes. Most
of the very large mammals became extinct in the last ice
age, but have smaller descendants.

4.1 List of prehistoric mammals

Main article: List of prehistoric mammals
Prehistoric mammals include:
33

Chapter 5

Stone Age
For other uses, see Stone Age (disambiguation).
The Stone Age is a broad prehistoric period during

5.1 Historical signicance

The Stone Age is contemporaneous with the evolution of
the genus Homo, the only exception possibly being at the
very beginning, when species prior to Homo may have
manufactured tools. According to the age and location
of the current evidence, the cradle of the genus is the
East African Rift System, especially toward the north in
Ethiopia, where it is bordered by grasslands. The closest
relative among the other living Primates, the genus Pan,
represents a branch that continued on in the deep forest,
where the primates evolved. The rift served as a conduit
for movement into southern Africa and also north down
the Nile into North Africa and through the continuation
of the rift in the Levant to the vast grasslands of Asia.
Starting from about 3 million years ago (mya) a single
biome established itself from South Africa through the
rift, North Africa, and across Asia to modern China,
which has been called transcontinental 'savannahstan'"
recently.[2] Starting in the grasslands of the rift, Homo
erectus, the predecessor of modern humans, found an
ecological niche as a tool-maker and developed a dependence on it, becoming a tool equipped savanna
dweller.[3]

Modern Awash River, Ethiopia, descendant of the PalaeoAwash, source of the sediments in which the oldest Stone Age
tools have been found

which stone was widely used to make implements with

a sharp edge, a point, or a percussion surface. The period
lasted roughly 3.4 million years, and ended between 6000
BCE and 2000 BCE with the advent of metalworking.[1]
Stone Age artifacts include tools used by humans and by
their predecessor species in the genus Homo, as well as the
earlier partly contemporaneous genera Australopithecus
and Paranthropus. Bone tools were used during this period as well but are rarely preserved in the archaeological
record. The Stone Age is further subdivided by the types
of stone tools in use.

5.2 The Stone Age in archaeology

5.2.1 Beginning of the Stone Age

During 2010, fossilised animal bones bearing marks from

stone tools were found in the Lower Awash Valley in
Ethiopia. Discovered by an international team led by
Shannon McPherron, at 3.4 million years old they are the
oldest evidence of stone tool use ever found anywhere in
The Stone Age is the rst of the three-age system the world.[1]
of archaeology, which divides human technological
The oldest known stone tools have been excavated from
prehistory into three periods:
several sites at Gona, Ethiopia, on the sediments of the
paleo-Awash River, which serve to date them. All the
The Stone Age
tools come from the Busidama Formation, which lies
above a disconformity, or missing layer, which would
The Bronze Age
have been from 2.9 to 2.7 mya. The oldest sites contain The Iron Age
ing tools are dated to 2.62.55 mya.[4] One of the most
34

5.2. THE STONE AGE IN ARCHAEOLOGY

35
the initial period of the Bronze Age and is unquestionably
part of the Age of Metals. The Bronze Age was followed
by the Iron Age. During this entire time stone remained
in use in parallel with the metals for some objects, including those also used in the Neolithic, such as stone pottery.
The transition out of the Stone Age occurred between
6000 BCE and 2500 BCE for much of humanity living in
North Africa and Eurasia. The rst evidence of human
metallurgy dates to between the 5th and 6th millennium
BCE in the archaeological sites of Majdanpek, Yarmovac
and Plonik (a copper axe from 5500 BCE belonging to
the Vinca culture), though not conventionally considered
part of the Chalcolithic or Copper Age, this provides
the earliest known example of copper metallurgy.[8] and
the Rudna Glava mine in Serbia. tzi the Iceman, a
mummy from about 3300 BCE carried with him a copper
axe and a int knife.

In regions such as Subsaharan Africa, the Stone Age was

followed directly by the Iron Age. The Middle East and
southeastern Asian regions progressed past Stone Age
technology around 6000 BCE. Europe, and the rest of
Asia became postStone Age societies by about 4000
BCE. The proto-Inca cultures of South America continued at a Stone Age level until around 2000 BCE, when
gold, copper and silver made their entrance, the rest folObsidian projectile point
lowing later. Australia remained in the Stone Age until
the 17th century. Stone tool manufacture continued. In
striking circumstances about these sites is that they are Europe and North America, millstones were in use until
from the Late Pliocene, where previous to their discovery well into the 20th century, and still are in many parts of
tools were thought to have evolved only in the Pleistocene. the world.
Rogers and Semaw, excavators at the locality, point out
that:[5]

5.2.3 The concept of Stone Age

"...the earliest stone tool makers were skilled
intknappers .... The possible reasons behind
this seeming abrupt transition from the absence
of stone tools to the presence thereof include ...
gaps in the geological record.

The terms was never meant to suggest that advancement

and time periods in prehistory are only measured by the
type of tool material, rather than, for example, social organization, food sources exploited, adaptation to climate,
adoption of agriculture, cooking, settlement and religion.
Like pottery, the typology of the stone tools combined
with the relative sequence of the types in various regions
provide a chronological framework for the evolution of
man and society. They serve as diagnostics of date, rather
than characterizing the people or the society.

The excavators are condent that more tools will be

found elsewhere from 2.9 mya. The species who
made the Pliocene tools remains unknown. Fragments
of Australopithecus garhi, Australopithecus aethiopicus[6]
and Homo, possibly Homo habilis, have been found in
sites near the age of the oldest tools.[7]
Lithic analysis is a major and specialised form of archaeological investigation. It involves the measurement
of the stone tools to determine their typology, function
5.2.2 End of the Stone Age
and the technology involved. It includes scientic study
Innovation of the technique of smelting ore ended the of the lithic reduction of the raw materials, examining
Stone Age and began the Bronze Age. The rst most how the artifacts were made. Much of this study takes
signicant metal manufactured was bronze, an alloy of place in the laboratory in the presence of various specialcopper and tin, each of which was smelted separately. ists. In experimental archaeology, researchers attempt to
The transition from the Stone Age to the Bronze Age was create replica tools, to understand how they were made.
a period during which modern people could smelt cop- Flintknappers are craftsmen who use sharp tools to reper, but did not yet manufacture bronze, a time known duce intstone to int tool.
as the Copper Age, or more technically the Chalcolithic, In addition to lithic analysis, the eld prehistorian uticopper-stone age. The Chalcolithic by convention is lizes a wide range of techniques derived from multiple

36

CHAPTER 5. STONE AGE

a negative characterisation. For them, 'primitive' denotes irrational use of resources and absence of the intellectual and moral standards of
'civilised' human societies.... From the standpoint of anthropological knowledge, both these
views are equally one-sided and simplistic.

5.2.4 The three-stage system

A variety of stone tools

elds. The work of the archaeologist in determining the

paleocontext and relative sequence of the layers is supplemented by the eorts of the geologic specialist in identifying layers of rock over geologic time, of the paleontological specialist in identifying bones and animals, of the
palynologist in discovering and identifying plant species,
of the physicist and chemist in laboratories determining
dates by the carbon-14, potassium-argon and other methods. Study of the Stone Age has never been mainly about
stone tools and archaeology, which are only one form of
evidence. The chief focus has always been on the society
and the physical people who belonged to it.
Useful as it has been, the concept of the Stone Age has its
limitations. The date range of this period is ambiguous,
disputed, and variable according to the region in question. While it is possible to speak of a general 'stone
age' period for the whole of humanity, some groups never
developed metal-smelting technology, so remained in a
'stone age' until they encountered technologically developed cultures. The term was innovated to describe the
archaeological cultures of Europe. It may not always be
the best in relation to regions such as some parts of the
Indies and Oceania, where farmers or hunter-gatherers
used stone for tools until European colonisation began.

In the 1920s, South African archaeologists organizing the

stone tool collections of that country observed that they
did not t the newly detailed Three-Age System. In the
words of J. Desmond Clark,[10]
It was early realized that the threefold division of culture into Stone, Bronze and Iron
Ages adopted in the nineteenth century for Europe had no validity in Africa outside the Nile
valley.
Consequently they proposed a new system for Africa, the
Three-stage System. Clark regarded the Three-age System as valid for North Africa; in sub-Saharan Africa, the
Three-stage System was best.[11] In practice, the failure
of African archaeologists either to keep this distinction
in mind, or to explain which one they mean, contributes
to the considerable equivocation already present in the literature. There are in eect two Stone Ages, one part of
the Three-age and the other constituting the Three-stage.
They refer to one and the same artifacts and the same
technologies, but vary by locality and time.
The Three-stage System was proposed in 1929 by Astley
John Hilary Goodwin, a professional archaeologist, and
Clarence van Riet Lowe, a civil engineer and amateur archaeologist, in an article titled Stone Age Cultures of
South Africa in the journal Annals of the South African
Museum. By then, the dates of the Early Stone Age, or
Paleolithic, and Late Stone Age, or Neolithic (neo = new),
were fairly solid and were regarded by Goodwin as absolute. He therefore proposed a relative chronology of periods with oating dates, to be called the Earlier and Later
Stone Age. The Middle Stone Age would not change its
name, but it would not mean Mesolithic.[12]

The archaeologists of the late 19th and early 20th centuries CE, who adapted the three-age system to their
ideas, hoped to combine cultural anthropology and archaeology in such a way that a specic contemporaneous
tribe can be used to illustrate the way of life and beliefs
of the people exercising a specic Stone-Age technology.
As a description of people living today, the term stone age The duo thus reinvented the Stone Age. In Sub-Saharan
is controversial. The Association of Social Anthropolo- Africa, however, it was ended by the intrusion of the Iron
gists discourages this use, asserting:[9]
Age from the north. The Neolithic and the Bronze Age
never occurred. Moreover, the technologies included in
To describe any living group as 'primitive'
those 'stages, as Goodwin called them, were not exactly
or 'Stone Age' inevitably implies that they are
the same. Since then, the original relative terms have beliving representatives of some earlier stage of
come identied with the technologies of the Paleolithic
human development that the majority of huand Mesolithic, so that they are no longer relative. Moremankind has left behind. For some, this could
over, there has been a tendency to drop the comparative
be a positive description, implying, for examdegree in favor of the positive: resulting in two sets of
ple, that such groups live in greater harmony
Early, Middle and Late Stone Ages of quite dierent conwith nature .... For others, ... 'primitive' is
tent and chronologies.

5.3. CHRONOLOGY
By voluntary agreement, archaeologists respect the decisions of the Pan-African Congress of Prehistory, which
meets every four years to resolve archaeological business
brought before it. Delegates are actually international; the
organization takes its name from the topic. Louis Leakey
hosted the rst one in Nairobi in 1947. It adopted Goodwin and Lowes 3-stage system at that time, the stages to
be called Early, Middle and Later.

5.2.5

The problem of the transitions

37
but were ocially rejected in 1965 (again on an advisory basis) by Burg Wartenstein Conference #29, Systematic Investigation of the African Later Tertiary and
Quaternary,[15] a conference in anthropology held by the
Wenner-Gren Foundation, at Burg Wartenstein Castle,
which it then owned in Austria, attended by the same
scholars that attended the Pan African Congress, including Louis Leakey and Mary Leakey, who was delivering
a pilot presentation of her typological analysis of Early
Stone Age tools, to be included in her 1971 contribution
to Olduvai Gorge, Excavations in Beds I and II, 1960
1963.[16]

18

Equivalent
Vostok T (C)

O Benthic
Carbonate (per mil)

The problem of the transitions in archaeology is a branch However, although the intermediate periods were gone,
of the general philosophic continuity problem, which the search for the transitions continued.
examines how discrete objects of any sort that are
contiguous in any way can be presumed to have a relationship of any sort. In archaeology, the relationship is
5.3 Chronology
one of causality. If Period B can be presumed to descend
from Period A, there must be a boundary between A and
B, the AB boundary. The problem is in the nature of
2
2
41 kyr cycle
100 kyr cycle
this boundary. If there is no distinct boundary, then the
2.5
0
-2
population of A suddenly stopped using the customs char3
Five Million Years of
-4
acteristic of A and suddenly started using those of B, an
3.5
Climate Change
-6
unlikely scenario in the process of evolution. More real4
-8 From Sediment Cores
istically, a distinct border period, the A/B transition, ex4.5
Millions of Years Ago
isted, in which the customs of A were gradually dropped
and those of B acquired. If transitions do not exist, then
Time series plot of temperature over the previous 5 million years
there is no proof of any continuity between A and B.
The Stone Age of Europe is characteristically in decit
of known transitions. The 19th and early 20th-century
innovators of the modern three-age system recognized
the problem of the initial transition, the gap between
the Paleolithic and the Neolithic. Louis Leakey provided
something of an answer by proving that man evolved in
Africa. The Stone Age must have begun there to be carried repeatedly to Europe by migrant populations. The
dierent phases of the Stone Age thus could appear there
without transitions. The burden on African archaeologists became all the greater, because now they must nd
the missing transitions in Africa. The problem is dicult
and ongoing.

In 1859 Jens Jacob Worsaae rst proposed a division of

the Stone Age into older and younger parts based on his
work with Danish kitchen middens that began in 1851.[17]
In the subsequent decades this simple distinction developed into the archaeological periods of today. The major
subdivisions of the Three-age Stone Age cross two epoch
boundaries on the geologic time scale:

After its adoption by the First Pan African Congress in

1947, the Three-Stage Chronology was amended by the
Third Congress in 1955 to include a First Intermediate Period between Early and Middle, to encompass the
Fauresmith and Sangoan technologies, and the Second Intermediate Period between Middle and Later, to encompass the Magosian technology and others. The chronologic basis for denition was entirely relative. With the
arrival of scientic means of nding an absolute chronology, the two intermediates turned out to be will-of-thewisps. They were in fact Middle and Lower Paleolithic.
Fauresmith is now considered to be a facies of Acheulean,
while Sangoan is a facies of Lupemban.[13] Magosian is
an articial mix of two dierent periods.[14]

The geologic PleistoceneHolocene boundary

(modern climate)

The geologic PliocenePleistocene

(highly glaciated climate)

boundary

The Paleolithic period of archaeology

Mesolithic or Epipaleolithic period of archaeology

Neolithic period of archaeology
The succession of these phases varies enormously from
one region (and culture) to another.

5.3.1 Three-age chronology

Main articles: Paleolithic, Human evolution and ThreeOnce seriously questioned, the intermediates did not wait age system
for the next Pan African Congress two years hence,

38

CHAPTER 5. STONE AGE

The Paleolithic or Palaeolithic (from Greek: ,

palaios, old"; and , lithos, stone lit. old stone,
coined by archaeologist John Lubbock and published in
1865) is the earliest division of the Stone Age. It covers the greatest portion of humanitys time (roughly 99%
of human technological history,[18] where human and
humanity are interpreted to mean the genus Homo),
extending from 2.5 or 2.6 million years ago, with the
rst documented use of stone tools by hominans such as
Homo habilis, to the end of the Pleistocene around 10,000
BCE.[18] The Paleolithic era ended with the Mesolithic,
or in areas with an early neolithisation, the Epipaleolithic.
Lower Paleolithic
Main article: Lower Paleolithic
At sites dating from the Lower Paleolithic Period (about
2,500,000 to 200,000 years ago), simple pebble tools
have been found in association with the remains of what
may have been the earliest human ancestors. A somewhat
more sophisticated Lower Paleolithic tradition, known as
the Chopper chopping-tool industry, is widely distributed
in the Eastern Hemisphere. This tradition is thought
to have been the work of the hominin species named
Homo erectus. Although no such fossil tools have yet
been found, it is believed that H. erectus probably made
tools of wood and bone as well as stone. About 700,000
years ago, a new Lower Paleolithic tool, the hand ax, appeared. The earliest European hand axes are assigned
to the Abbevillian industry, which developed in northern
France in the valley of the Somme River; a later, more rened hand-ax tradition is seen in the Acheulian industry,
evidence of which has been found in Europe, Africa, the
Middle East, and Asia. Some of the earliest known hand
axes were found at Olduvai Gorge (Tanzania) in association with remains of H. erectus. Alongside the handax tradition there developed a distinct and very dierent stone-tool industry, based on akes of stone: special
tools were made from worked (carefully shaped) akes of
int. In Europe, the Clactonian industry is one example
of a ake tradition. The early ake industries probably
contributed to the development of the Middle Paleolithic
ake tools of the Mousterian industry, which is associated
with the remains of Neanderthal man.[19]
Oldowan in Africa Main article: Oldowan
The earliest documented stone tools were found in East
Africa, manufacturers unknown. They belonged to an
industry now known as Oldowan, after the type site of
Olduvai Gorge in Tanzania; however, sites in Ethiopia
later proved to be older.

This is a Mode 1, or Oldowan, stone tool from the western Sahara.

nal stone is called a core; the resultant pieces, akes. Typically, but not necessarily, small pieces are detached from
a larger piece, in which case the larger piece may be called
the core and the smaller pieces the akes. The prevalent
usage, however, is to call all the results akes, which can
be confusing. A split in half is called bipolar aking.
Consequently the method is often called core-andake. More recently, the tradition has been called small
ake since the akes were small compared to subsequent
Acheulean tools.[20]
The essence of the Oldowan is the making
and often immediate use of small akes.
Another naming scheme is Pebble Core Technology
(PBC)":[21]
Pebble cores are ... artifacts that have
been shaped by varying amounts of hardhammer percussion.
Various renements in the shape have been called choppers, discoids, polyhedrons, subspheroid, etc. To date no
reasons for the variants have been ascertained:[22]
From a functional standpoint, pebble
cores seem designed for no specic purpose.
However, they would not have been manufactured for no
purpose:[22]
Pebble cores can be useful in many cutting, scraping or chopping tasks, but ... they
are not particularly more ecient in such tasks
than a sharp-edged rock ....

The tools were formed by knocking pieces o a river peb- The whole point of their utility is that each is a sharpble, or stones like it, with a hammerstone to obtain large edged rock in locations where nature has not provided
and small pieces with one or more sharp edges. The origi- any. There is additional evidence that Oldowan, or Mode

5.3. CHRONOLOGY
1, tools were utilized in percussion technology"; that is,
they were designed to be gripped at the blunt end and
strike something with the edge, from which use they were
given the name of choppers. Modern science has been
able to detect mammalian blood cells on Mode 1 tools
at Sterkfontein, Member 5 East, in South Africa. As the
blood must have come from a fresh kill, the tool users
are likely to have done the killing and used the tools for
butchering. Plant residues bonded to the silicon of some
tools conrm the use to chop plants.[23]
Although the exact species authoring the tools remains
unknown, Mode 1 tools in Africa were manufactured and
used predominantly by Homo habilis. They cannot be said
to have developed these tools or to have contributed the
tradition to technology. They continued a tradition of yet
unknown origin. As chimpanzees sometimes naturally
use percussion to extract or prepare food in the wild, and
may use either unmodied stones or stones that they have
split, creating an Oldowan tool, the tradition may well be
far older than its current record.

39
temporaneously in the same regions H. habilis inherited
the tools around 2.3 mya. At about 1.9 mya H. erectus
came on stage and lived contemporaneously with the others. Mode 1 was now being shared by a number of Hominans over the same ranges, presumably subsisting in different niches, but the archaeology is not precise enough
to say which.
Oldowan out of Africa Tools of the Oldowan tradition rst came to archaeological attention in Europe,
where, being intrusive and not well dened, compared
to the Acheulean, they were puzzling to archaeologists.
The mystery would be elucidated by African archaeology at Olduvai, but meanwhile, in the early 20th century,
the term Pre-Acheulean came into use in climatology.
C.E.P, Brooks, a British climatologist working in the
United States, used the term to describe a chalky boulder
clay underlying a layer of gravel at Hoxne, central England, where Acheulean tools had been found.[31] Whether
any tools would be found in it and what type was not
known. Hugo Obermaier, a contemporary German archaeologist working in Spain, quipped:

Towards the end of Oldowan in Africa a new species appeared over the range of Homo habilis: Homo erectus.
The earliest unambiguous evidence is a whole cranium,
KNM-ER 3733 (a nd identier) from Koobi Fora in
Unfortunately, the stage of human indusKenya, dated to 1.78 mya.[24] An early skull fragment,
try which corresponds to these deposits cannot
KNM-ER 2598, dated to 1.9 mya, is considered a good
be positively identied. All we can say is that
candidate also.[25] Transitions in paleoanthropology are
it is pre-Acheulean....
always hard to nd, if not impossible, but based on the
long-legged limb morphology shared by H. habilis and
H. rudolfensis in East Africa, an evolution from one of This uncertainty was claried by the subsequent excavations at Olduvai; nevertheless, the term is still in use for
those two has been suggested.[26]
pre-Acheulean contexts, mainly across Eurasia, that are
The most immediate cause of the new adjustments apyet unspecied or uncertain but with the understanding
pears to have been an increasing aridity in the region and
that they are or will turn out to be pebble-tool.[32]
consequent contraction of parkland savanna, interspersed
with trees and groves, in favor of open grassland, dated There are ample associations of Mode 2 with H. erectus in
1.81.7 mya. During that transitional period the percent- Eurasia. H. erectus Mode 1 associations are scantier but
age of grazers among the fossil species increased from they do exist, especially in the Far East. One strong piece
1525% to 45%, dispersing the food supply and requir- of evidence prevents the conclusion that only H. erectus
ing a facility among the hunters to travel longer distances reached Eurasia: at Yiron, Israel, Mode 1 tools have been
[33]
comfortably, which H. erectus obviously had.[27] The ul- found dating to 2.4 mya, about 0.5 my earlier than the
timate proof is the dispersal of H. erectus across much known H. erectus nds. If the date is correct, either anof Africa and Asia, substantially before the development other Hominan preceded H. erectus out of Africa or the
of the Mode 2 technology and use of re ....[26] H. erectus earliest H. erectus has yet to be found.
carried Mode 1 tools over Eurasia.
After the initial appearance at Gona in Ethiopia at 2.7
According to the current evidence (which may change at
any time) Mode 1 tools are documented from about 2.6
mya to about 1.5 mya in Africa,[28] and to 0.5 mya outside
of it.[29] The genus Homo is known from H. habilis and
H. rudolfensis from 2.3 to 2.0 mya, with the latest habilis
being an upper jaw from Koobi Fora, Kenya, from 1.4
mya. H. erectus is dated 1.80.6 mya.[30]
According to this chronology Mode 1 was inherited by Homo from unknown Hominans, probably
Australopithecus and Paranthropus, who must have continued on with Mode 1 and then with Mode 2 until their
extinction no later than 1.1 mya. Meanwhile living con-

mya, pebble tools date from 2.0 mya at Sterkfontein,

Member 5, South Africa, and from 1.8 mya at El Kherba,
Algeria, North Africa. The manufacturers had already
left pebble tools at Yiron, Israel, at 2.4 mya, Riwat, Pakistan, at 2.0 mya, and Renzidong, South China, at over 2
mya.[34] The identication of a fossil skull at Mojokerta,
Pernung Peninsula on Java, dated to 1.8 mya, as H. erectus, suggests that the African nds are not the earliest to
be found in Africa, or that, in fact, erectus did not originate in Africa after all but on the plains of Asia.[26] The
outcome of the issue waits for more substantial evidence.
Erectus was found also at Dmanisi, Georgia, from 1.75
mya in association with pebble tools.

40
Pebble tools are found the latest rst in southern Europe
and then in northern. They begin in the open areas of Italy
and Spain, the earliest dated to 1.6 mya at Pirro Nord,
Italy. The mountains of Italy are rising at a rapid rate
in the framework of geologic time; at 1.6 mya they were
lower and covered with grassland (as much of the highlands still are). Europe was otherwise mountainous and
covered over with dense forest, a formidable terrain for
warm-weather savanna dwellers. Similarly there is no evidence that the Mediterranean was passable at Gibraltar
or anywhere else to H. erectus or earlier hominans. They
might have reached Italy and Spain along the coasts.
In northern Europe pebble tools are found earliest at
Happisburgh, United Kingdom, from 0.8 mya. The
last traces are from Kents Cavern, dated 0.5 mya. By
that time H. erectus is regarded as having been extinct; however, a more modern version apparently had
evolved, Homo heidelbergensis, who must have inherited
the tools.[35] He also explains the last of the Acheulean in
Germany at 0.4 mya.
In the late 19th and early 20th centuries archaeologists
worked on the assumptions that a succession of Hominans and cultures prevailed, that one replaced another.
Today the presence of multiple hominans living contemporaneously near each other for long periods is accepted
as proved true; moreover, by the time the previously assumed earliest culture arrived in northern Europe, the
rest of Africa and Eurasia had progressed to the Middle and Upper Palaeolithic, so that across the earth all
three were for a time contemporaneous. In any given region there was a progression from Oldowan to Acheulean,
Lower to Upper, no doubt.

CHAPTER 5. STONE AGE

at Kokiselei, West Turkana, Kenya.[25] At Sterkfontein,
South Africa, they are in Member 5 West, 1.71.4
mya.[23] The 1.7 is a fairly certain, fairly standard date.
Mode 2 is often found in association with H. erectus. It
makes sense that the most advanced tools should have
been innovated by the most advanced Hominan; consequently, they are typically given credit for the innovation.
A Mode 2 tool is a biface consisting of two concave
surfaces intersecting to form a cutting edge all the way
around, except in the case of tools intended to feature a
point. More work and planning go into the manufacture
of a Mode 2 tool. The manufacturer hits a slab o a larger
rock to use as a blank. Then large akes are struck o the
blank and worked them into bifaces by hard-hammer percussion on an anvil stone. Finally the edge is retouched:
small akes are hit o with a bone or wood soft hammer to sharpen or resharpen it. The core can be either
the blank or another ake. Blanks are ported for manufacturing supply in places where nature has provided no
suitable stone.
Although most Mode 2 tools are easily distinguished from
Mode 1, there is a close similarity of some Oldowan and
some Acheulean, which can lead to confusion. Some
Oldowan tools are more carefully prepared to form a
more regular edge. One distinguishing criterion is the
size of the akes. In contrast to the Oldowan small ake
tradition, Acheulean is large ake:" The primary technological distinction remaining between Oldowan and
the Acheulean is the preference for large akes (>10
cm) as blanks for making large cutting tools (handaxes
and cleavers) in the Acheulean.[36] Large Cutting Tool
(LCT)" has become part of the standard terminology as
well.[22]

In North Africa, the presence of Mode 2 remains a mysAcheulean in Africa Main article: Acheulean
The end of Oldowan in Africa was brought on by the tery, as the oldest nds are from Thomas Quarry in
Morocco at 0.9 mya.[34] Archaeological attention, however, shifts to the Jordan Rift Valley, an extension of
the East African Rift Valley (the east bank of the Jordan is slowly sliding northward as East Africa is thrust
away from Africa). Evidence of use of the Nile Valley
is in decit, but Hominans could easily have reached the
palaeo-Jordan river from Ethiopia along the shores of the
Red Sea, one side or the other. A crossing would not have
been necessary, but it is more likely there than over a theoretical but unproven land bridge through either Gibraltar
or Sicily.
Meanwhile Acheulean went on in Africa past the 1.0 mya
mark and also past the extinction of H. erectus there. The
last Acheulean in East Africa is at Olorgesailie, Kenya,
dated to about 0.9 mya. Its owner was still H. erectus,[34]
but in South Africa, Acheulean at Elandsfontein, 1.0
0.6 mya, is associated with Saldanha man, classied as
H. heidelbergensis, a more advanced, but not yet modAn Acheulean tool, not worked over the entire surface
ern, descendant most likely of H. erectus. The Thoman
Quarry Hominans in Morocco similarly are most likely
appearance of Acheulean, or Mode 2, stone tools. The Homo rhodesiensis,[37] in the same evolutionary status as
earliest known instances are in the 1.71.6 mya layer

5.3. CHRONOLOGY

41

H. heidelbergensis.

itation of archeology, but after 1 mya evidence not available to Movius indicates the prevalence of Acheulean.
For example, the Acheulean site at Bose, China, is dated
Acheulean out of Africa Mode 2 is rst known out 0.8033K mya.[46] The authors of this chronologically
of Africa at 'Ubeidiya, Israel, a site now on the Jordan later East Asian Acheulean remain unknown, as does
River, then frequented over the long term (hundreds of whether it evolved in the region or was brought in.
thousands of years) by Homo on the shore of a variableThere is no named boundary line between Mode 1 and
level palaeo-lake, long since vanished. The geology was
Mode 2 on the west; nevertheless, Mode 2 is equally late
created by successive transgression and regression of
in Europe as it is in the Far East. The earliest comes
the lake[38] resulting in four cycles of layers. The tools
from a rock shelter at Estrecho de Qupar in Spain, dated
are located in the rst two, Cycles Li (Limnic Inferior)
to greater than 0.9 mya. Teeth from an undetermined
and Fi (Fluviatile Inferior), but mostly in Fi. The cyHominan were found there also.[47] The last Mode 2 in
cles represent dierent ecologies and therefore dierent
Southern Europe is from a deposit at Fontana Ranuccio
cross-sections of fauna, which makes it possible to date
near Anagni in Italy dated to 0.45 mya, which is generally
them. They appear to be the same faunal assemblages as
linked to Homo cepranensis, a late variant of H. erectus,
the Ferenta Faunal Unit in Italy, known from excavations
a fragment of whose skull was found at Ceprano nearby,
at Selvella and Pietertta, dated to 1.61.2 mya.[39]
dated 0.46 mya.[48]
At 'Ubeidiya the marks on the bones of the animal species
found there indicate that the manufacturers of the tools
butchered the kills of large predators, an activity that has Middle Paleolithic
been termed scavenging.[40] There are no living oors,
nor did they process bones to obtain the marrow. These Main article: Middle Paleolithic
activities cannot be understood therefore as the only or
even the typical economic activity of Hominans. Their
This period is best known as the era during which the
interests were selective: they were primarily harvesting
Neanderthals lived in Europe and the Near East (c.
the meat of Cervids,[41] which is estimated to have been
300,00028,000 years ago). Their technology is mainly
available without spoiling for up to four days after the kill.
the Mousterian, but Neanderthal physical characterisThe majority of the animals at the site were of tics have been found also in ambiguous association with
Palaearctic biogeographic origin.[42] However, these the more recent Chtelperronian archeological culture
overlapped in range on 3060% of African biogeo- in Western Europe and several local industries like the
graphic origin.[43] The biome was Mediterranean, not Szeletian in Eastern Europe/Eurasia. There is no evisavanna. The animals were not passing through; there dence for Neanderthals in Africa, Australia or the Amerwas simply an overlap of normal ranges. Of the Homi- icas.
nans, H. erectus left several cranial fragments. Teeth of
Neanderthals nursed their elderly and practised ritual
undetermined species may have been H. ergaster.[44] The
burial indicating an organised society. The earliest evitools are classied as Lower Acheulean and Developed
dence (Mungo Man) of settlement in Australia dates to
Oldowan. The latter is a disputed classication created
around 40,000 years ago when modern humans likely
by Mary Leakey to describe an Acheulean-like tradition
crossed from Asia by island-hopping. Evidence for symin Bed II at Olduvai. It is dated 1.531.27 mya. The date
bolic behavior such as body ornamentation and burial is
of the tools therefore probably does not exceed 1.5 mya;
ambiguous for the Middle Paleolithic and still subject to
1.4 is often given as a date. This chronology, which is
debate. The Bhimbetka rock shelters exhibit the earliest
denitely later than in Kenya, supports the out of Africa
traces of human life in India, some of which are approxhypothesis for Acheulean, if not for the Hominans.
imately 30,000 years old.
From Southwest Asia, as the Levant is now called, the
Acheulean extended itself more slowly eastward, arriving
at Isampur, India, about 1.2 mya. It does not appear in Upper Paleolithic
China and Korea until after 1mya and not at all in Indonesia. There is a discernible boundary marking the furthest Main article: Upper Paleolithic
extent of the Acheulean eastward before 1 mya, called the
Movius Line, after its proposer, Hallam L. Movius. On From 50,000 to 10,000 years ago in Europe, the Upthe east side of the line the small ake tradition contin- per Paleolithic ends with the end of the Pleistocene and
ues, but the tools are additionally worked Mode 1, with onset of the Holocene era (the end of the last ice age).
aking down the sides. In Athirampakkam at Chennai in Modern humans spread out further across the Earth durTamil Nadu the Acheulean age started at 1.51 mya and it ing the period known as the Upper Paleolithic. The Upis also prior than North India and Europe.[45]
per Paleolithic is marked by a relatively rapid succession
The cause of the Movius Line remains speculative, of often complex stone artifact technologies and a large
whether it represents a real change in technology or a lim- increase in the creation of art and personal ornaments.

42

CHAPTER 5. STONE AGE

During period between 35 and 10 kya evolved: from 38

to 30 kya Chtelperronian, 4028 Aurignacian, 2822
Gravettian, 2217 Solutrean, and 1810 Magdalenian.
All of these industries except the Chtelperronian are associated with anatomically modern humans. Authorship
of the Chtelperronian is still the subject of much debate.
The Americas were colonised via the Bering land bridge
which was exposed during this period by lower sea levels.
These people are called the Paleo-Indians, and the earliest accepted dates are those of the Clovis culture sites,
some 13,500 years ago. Globally, societies were huntergatherers but evidence of regional identities begins to appear in the wide variety of stone tool types being devel- gantija temples, Gozo. Some of the worlds oldest free-standing
oped to suit very dierent environments.
structures.
Epipaleolithic/Mesolithic
Main articles: Epipaleolithic, Mesolithic
The period starting from the end of the last ice age,
10,000 years ago, to around 6,000 years ago was characterized by rising sea levels and a need to adapt to a changing environment and nd new food sources. The development of Mode 5 (microlith) tools began in response to
these changes. They were derived from the previous Paleolithic tools, hence the term Epipaleolithic, or were intermediate between the Paleolithic and the Neolithic, hence
the term Mesolithic (Middle Stone Age). The choice of
a word depends on exact circumstances and the inclination of the archaeologists excavating the site. Microliths
were used in the manufacture of more ecient composite
tools, resulting in an intensication of hunting and shing and with increasing social activity the development
of more complex settlements, such as Lepenski Vir. Domestication of the dog as a hunting companion probably
dates to this period.
The earliest known battle occurred during the Mesolithic
period at a site in Egypt known as Cemetery 117.
Neolithic
Main article: Neolithic
The Neolithic, New Stone Age, was approximately characterized by the adoption of agriculture, the shift from
food gathering to food producing in itself is one of the
most revolutionary changes in human history so-called
Neolithic Revolution, the development of pottery, polished stone tools and more complex, larger settlements
such as atal Hyk and Jericho. Some of these features
began in certain localities even earlier, in the transitional
Mesolithic. The rst Neolithic cultures started around
7000 BCE in the fertile crescent and spread concentrically to other areas of the world; however, the Near East
was probably not the only nucleus of agriculture, the cultivation of maize in Meso-America and of rice in the Far
East being others.

Skara Brae, Scotland. Europes most complete Neolithic village

Due to the increased need to harvest and process plants,

ground stone and polished stone artifacts became much
more widespread, including tools for grinding, cutting,
and chopping. Skara Brae located on Orkney island
o Scotland is one of Europes best examples of a Neolithic village. The community contains stone beds,
shelves and even an indoor toilet linked to a stream. The
rst large-scale constructions were built, including settlement towers and walls, e.g., Jericho and ceremonial
sites, e.g.: Stonehenge. The gantija temples of Gozo
in the Maltese archipelago are the oldest surviving free
standing structures in the world, erected c. 36002500
BCE. The earliest evidence for established trade exists in
the Neolithic with newly settled people importing exotic
goods over distances of many hundreds of miles.
These facts show that there were sucient resources and
co-operation to enable large groups to work on these
projects. To what extent this was a basis for the development of elites and social hierarchies is a matter of ongoing debate.[49] Although some late Neolithic societies
formed complex stratied chiefdoms similar to Polynesian societies such as the Ancient Hawaiians, based on the
societies of modern tribesmen at an equivalent technological level, most Neolithic societies were relatively simple
and egalitarian.[50] A comparison of art in the two ages
leads some theorists to conclude that Neolithic cultures

5.4. MATERIAL CULTURE

43

were noticeably more hierarchical than the Paleolithic years ago.[53] It is considered as an equivalent of Eurocultures that preceded them.[51]
pean Middle Paleolithic.[54] It is associated with anatomically modern or almost modern Homo sapiens. Early
physical evidence comes from Omo [55] and Herto,[56]
both in Ethiopia and dated respectively at c. 195 ka and
5.3.2 Three-stage chronology
at c. 160 ka.
The Earlier or Early Stone Age (ESA)
The Later Stone Age (LSA)
Main articles: Paleolithic and Lower Paleolithic
This period is not to be identied with Old Stone Age,
Main article: Later Stone Age
The Later Stone Age (LSA, sometimes also called the
Late Stone Age) refers to a period in African prehistory.
Its beginnings are roughly contemporaneous with the European Upper Paleolithic. It lasts until historical times
and this includes cultures corresponding to Mesolithic
and Neolithic in other regions.

5.4 Material culture

5.4.1 Tools

Acheulean biface from Lake Langano area, Ethiopia.

a translation of Paleolithic, or with Paleolithic, or with

the Earlier Stone Age that originally meant what became the Paleolithic and Mesolithic. In the initial decades
of its denition by the Pan-African Congress of Prehistory, it was parallel in Africa to the Upper and Middle
Paleolithic. However, since then Radiocarbon dating has
shown that the Middle Stone Age is in fact contemporaneous with the Middle Paleolithic.[52] The Early Stone
Age therefore is contemporaneous with the Lower Paleolithic and happens to include the same main technologies, Oldowan and Acheulean, which produced Mode 1
and Mode 2 stone tools respectively. A distinct regional
term is warranted, however, by the location and chronology of the sites and the exact typology.

Stone tools were made from a variety of stone. For example, int and chert were shaped (or chipped) for use
as cutting tools and weapons, while basalt and sandstone
were used for ground stone tools, such as quern-stones.
Wood, bone, shell, antler (deer) and other materials were
widely used, as well. During the most recent part of
the period, sediments (such as clay) were used to make
pottery. Agriculture was developed and certain animals
were domesticated.
Some species of non-Primates are able to use stone tools,
such as the Sea Otter, which breaks Abalone shells with
them. Primates can both use and manufacture stone tools.
This combination of abilities is more marked in apes and
men, but only men, or more generally Hominans, depend
on tool use for survival.[57] The key anatomical and behavioral features required for tool manufacture, which are
possessed only by Hominans, are the larger thumb and the
ability to hold by means of an assortment of grips.[58]

5.4.2 Food and drink

Main articles: Paleolithic diet and Paleolithic diet and
nutrition

Food sources of the Palaeolithic hunter-gatherers were

wild plants and animals harvested from the environment.
They liked animal organ meats, including the livers,
Main article: Middle Stone Age
kidneys and brains. Large seeded legumes were part
of the human diet long before the agricultural revoluThe Middle Stone Age was a period of African prehis- tion, as is evident from archaeobotanical nds from the
tory between Early Stone Age and Late Stone Age. It be- Mousterian layers of Kebara Cave, in Israel.[59] Moregan around 300,000 years ago and ended around 50,000 over, recent evidence indicates that humans processed
The Middle Stone Age (MSA)

44

CHAPTER 5. STONE AGE

and consumed wild cereal grains as far back as 23,000 Petroglyphs

years ago in the Upper Paleolithic.[60]
Near the end of the Wisconsin glaciation, 15,000 to 9,000 Main article: Petroglyph
years ago, mass extinction of Megafauna such as the
Wooly mammoth occurred in Asia, Europe, North America and Australia. This was the rst Holocene extinction event. It possibly forced modication in the dietary
habits of the humans of that age and with the emergence
of agricultural practices, plant-based foods also became
a regular part of the diet. A number of factors have been
suggested for the extinction: certainly over-hunting, but
also deforestation and climate change.[61] The net eect
was to fragment the vast ranges required by the large animals and extinguish them piecemeal in each fragment.

Petroglyphs appeared in the Neolithic. A Petroglyph is an

intaglio abstract or symbolic image engraved on natural
stone by various methods, usually by prehistoric peoples.
They were a dominant form of pre-writing symbols. Petroglyphs have been discovered in dierent parts of the
world, including Asia (Bhimbetka, India), North America (Death Valley National Park), South America (Cumbe
Mayo, Peru), and Europe (Finnmark, Norway).
Rock paintings

5.4.3

Shelter and habitat

Around 2 million years ago, Homo habilis is believed

to have constructed the rst man-made structure in East
Africa, consisting of simple arrangements of stones to
hold branches of trees in position. A similar stone circular arrangement believed to be around 380,000 years
old was discovered at Terra Amata, near Nice, France.
(Concerns about the dating have been raised, see Terra
Amata). Several human habitats dating back to the Stone
Age have been discovered around the globe, including:
A tent-like structure inside a cave near the Grotte du
Lazaret, Nice, France.

Rock painting at Bhimbetka, India, a World heritage site

Main article: Cave painting

A structure with a roof supported with timber, disIn paleolithic times, mostly animals were painted, in thecovered in Dolni Vestonice, The Czech Republic,
ory ones that were used as food or represented strength,
dates to around 23,000 BCE. The walls were made
such as the rhinoceros or large cats (as in the Chauvet
of packed clay blocks and stones.
Cave). Signs such as dots were sometimes drawn.
Rare human representations include handprints and half Many huts made of mammoth bones were found in human/half-animal gures. The Cave of Chauvet in the
Eastern Europe and Siberia. The people who made Ardche dpartement, France, contains the most importhese huts were expert mammoth hunters. Exam- tant cave paintings of the paleolithic era, dating from
ples have been found along the Dniepr river valley about 31,000 BCE. The Altamira cave paintings in Spain
of Ukraine, including near Chernihiv, in Moravia, were done 14,000 to 12,000 BCE and show, among othCzech Republic and in southern Poland.
ers, bisons. The hall of bulls in Lascaux, Dordogne,
France, dates from about 15,000 to 10,000 BCE.
An animal hide tent dated to around 15000 to 10000 The meaning of many of these paintings remains unBCE, in the Magdalenian, was discovered at Plateau known. They may have been used for seasonal rituParain, France.
als. The animals are accompanied by signs that suggest

5.4.4

Art

Prehistoric art is visible in the artifacts. Prehistoric music

is inferred from found instruments, while parietal art can
be found on rocks of any kind. The latter are petroglyphs
and rock paintings. The art may or may not have had a
religious function.

a possible magic use. Arrow-like symbols in Lascaux are

sometimes interpreted as calendar or almanac use, but the
evidence remains interpretative.[62]
Some scenes of the Mesolithic, however, can be typed
and therefore, judging from their various modications,
are fairly clear. One of these is the battle scene between
organized bands of archers. For example, the marching
Warriors, a rock painting at Cingle de la Mola, Castelln
in Spain, dated to about 7,0004,000 BCE, depicts about

5.5. MODERN POPULAR CULTURE AND THE STONE AGE

50 bowmen in two groups marching or running in step
toward each other, each man carrying a bow in one hand
and a stful of arrows in the other. A le of ve men leads
one band, one of whom is a gure with a high crowned
hat. In other scenes elsewhere, the men wear headdresses and knee ornaments but otherwise ght nude.
Some scenes depict the dead and wounded, bristling with
arrows.[63] One is reminded of tzi the Iceman, a Copper
Age mummy revealed by an Alpine melting glacier, who
collapsed from loss of blood due to an arrow wound in
the back.

5.4.5

45

Megalithic tombs, multichambered, and dolmens,

single-chambered, were graves with a huge stone
slab stacked over other similarly large stone slabs;
they have been discovered all across Europe and
Asia and were built in the Neolithic and the Bronze
Age.

5.5 Modern popular culture and

the Stone Age

Stone Age rituals and beliefs

Main articles: Paleolithic religion, Prehistoric religion

and Mother goddess
Modern studies and the in-depth analysis of nds dating

Imaginative depiction of the Stone Age, by Viktor Vasnetsov

Poulnabrone dolmen in County Clare, Ireland

The image of the caveman is commonly associated with

the Stone Age. For example, the 2003 documentary series showing the evolution of humans through the Stone
Age was called Walking with Cavemen, although only the
last programme showed humans living in caves. While
the idea that human beings and dinosaurs coexisted is
sometimes portrayed in popular culture in cartoons, lms
and computer games, such as The Flintstones, One Million
Years B.C. and Chuck Rock, the notion of hominids and
non-avian dinosaurs co-existing is not supported by any
scientic evidence.
Other depictions of the Stone Age include the best-selling
Earths Children series of books by Jean M. Auel, which
are set in the Paleolithic and are loosely based on archaeological and anthropological ndings. The 1981 lm
Quest for Fire by Jean-Jacques Annaud tells the story of
a group of neanderthals searching for their lost re. A
twenty rst century series, Chronicles of Ancient Darkness by Michelle Paver tells of two New Stone Age children ghting to full a prophecy and save their clan.

Monte Bubbonia dolmen (single-chambered tomb), Sicily[64]

from the Stone Age indicate certain rituals and beliefs of

the people in those prehistoric times. It is now believed
that activities of the Stone Age humans went beyond the
immediate requirements of procuring food, body coverings, and shelters. Specic rites relating to death and
burial were practiced, though certainly diering in style
and execution between cultures.

5.6 See also

Megalith
Prehistoric warfare
Ice Age
Pleistocene

46

CHAPTER 5. STONE AGE

Homo

[21] Shea 2010, p. 49

Timeline of the Stone Age

[22] Shea 2010, p. 50

[23] Barham & Mitchell 2008, p. 132

5.7 Notes
[1] http://www.nhm.ac.uk/about-us/news/2010/august/
oldest-tool-use-and-meat-eating-revealed75831.html

[24] Barham & Mitchell 2008, pp. 126127.

[25] Barham & Mitchell 2008, p. 128
[26] Barham & Mitchell 2008, p. 145

[2] Barham & Mitchell 2008, p. 106

[27] Barham & Mitchell 2008, p. 146.

[3] Barham & Mitchell 2008, p. 147

[28] Barham & Mitchell 2008, p. 112

[4] Rogers & Semaw 2009, pp. 162163

[29] Shea 2010, p. 57

[5] Rogers & Semaw 2009, p. 155

[30] Barham & Mitchell 2008, p. 73

[6] As to whether aethiopicus is the genus Australopithecus or

the genus Paranthropus, broken out to include the more
robust forms, anthropological opinion is divided and both
usages occur in the professional sources.

[31] Brooks, Charles E.P. (1919), The Correlation of the

Quaternary Deposits of the British Isles with Those of
the Continent of Europe, Annual Report of the Board of
Regents of the Smithsonian Institution 1917, Washington:
Government Pronting Oce, p. 277

[7] Rogers & Semaw 2009, p. 164

[8] Neolithic Vinca was a metallurgical culture. Archaeo
News. Reuters. 17 November 2007. Retrieved 25 January 2011.

[32] Hugo Obermaier; Christine Matthew; Henry Osborne

(1924). Fossil Man in Spain. New Haven: Yale University
Press for the Hispanic Society of America. p. 272.
[33] Barham & Mitchell 2008, pp. 106107

[9] ASA Statement on the use of 'primitive' as a descriptor of

contemporary human groups. ASA News (Association of
Social Anthropologists of the UK and Commonwealth).
27 August 2007.

[34] Shea 2010, pp. 5557

[35] Barham & Mitchell 2008, p. 24

[10] Clark 1970, p. 22

[36] Barham & Mitchell 2008, p. 130

[11] Clark 1970, pp. 1819

[37] Jean-Paul Raynal; et al. (2010). Hominid Cave at

Thomas Quarry I (Casablanca, Morocco): Recent ndings and their context. Quaternary International (223
224): 369382.

[12] Deacon & Deacon 1999, pp. 56

[13] Isaac, Glynn (1982). The Earliest Archaeological
Traces. In Clark, J. Desmond. The Cambridge History
of Africa. Volume. I: From the Earliest Times to C. 500
BC. Cambridge: Cambridge University Press. p. 246.
[14] Willoughby, Pamela R. (2007). The evolution of modern
humans in Africa: a comprehensive guide. Lanham, MD:
AltaMira Press. p. 54.
[15] Barham & Mitchell 2008, p. 477
[16] History: Systematic Investigation of the African Later
Tertiary and Quaternary. The Wenner-Gren Foundation.
Retrieved 3 March 2011.
[17] Worsaae, Jens Jacob Asmussen. Encyclopdia Britannica.
[18] Toth, Nicholas; Schick, Kathy (2007). 21 Overview
of Paleolithic Archaeology. In Henke, H.C. Winfried;
Hardt, Thorolf; Tattersall, Ian. Handbook of Paleoanthropology. Volume 3. Berlin; Heidelberg; New York:
Springer-Verlag. p. 1944. ISBN 978-3-540-32474-4
[19] http://www.britannica.com/EBchecked/topic/439507/
Paleolithic-Period
[20] Barham & Mitchell 2008, p. 130.

[38] Belmaker 2006, p. 9

[39] Belmaker 2006, pp. 119120
[40] Belmaker 2006, p. 149
[41] Belmaker 2006, p. 147
[42] Belmaker 2006, p. 67
[43] Belmaker 2006, p. 21
[44] Belmaker 2006, p. 20
[45] Acheulian stone tools discovered near Chennai. The
Hindu.
[46] Bose, China. What Does It Mean to be Human?. Smithsonian National Museum of Natural History.
[47] Dalton, Rex (2 September 2009).
Europes
oldest axes discovered.
Nature News (Nature).
doi:10.1038/news.2009.878.
[48] Giovanni Muttoni; et al. (2009). Pleistocene magnetochronology of early hominid sites at Ceprano and
Fontana Ranuccio, Italy. Earth and Planetary Science
Letters 286: 255268. doi:10.1016/j.epsl.2009.06.032.

5.8. REFERENCES

[49] Kuijt, Ian (2000). Chapter 13: Near Eastern Neolithic

Research: Directions and Trends. In Kuijt, Ian. Life
in Neolithic Farming Communities: Social Organization,
Identity, and dierentiation. Fundamental Issues in Archaeology. New York: Kluwer Academic/Plenum Publishers. p. 317
[50] Boehm, Christopher (2000). The Origin of Morality as
Social Control. In Katz, Leonard D. Evolutionary Origins of Morality: Cross-disciplinary Perspectives. Journal
of Consciousness Studies Volume 7. Thorverton: Imprint
Academic. p. 158. ISBN 0-7190-5612-8

47

5.8 References
Barham, Lawrence; Mitchell, Peter (2008). The
First Africans: African Archaeology from the Earliest Toolmakers to Most Recent Foragers. Cambridge
World Archaeology. Oxford: Oxford University
Press.
Belmaker, Miriam (March 2006). Community
Structure through Time: 'Ubeidiya, a Lower Pleistocene Site as a Case Study (Thesis). Paleoanthropology Society.

[51] Guthrie, Russell Dale (2005). The Nature of Paleolithic

Art. Chicago: University of Chicago Press. pp. 419420.
ISBN 978-0-226-31126-5.

Clark, J. Desmond (1970). The Prehistory of Africa.

Ancient People and Places, Volume 72. New York;
Washington: Praeger Publishers.

[52] Clark, J. Desmond (1982). The Culture of the Middle

Paleolithic/MIddle Stone Age. In Clark, J. Desmond.
The Cambridge History of Africa. Volume. I: From the
Earliest Times to C. 500 BC. Cambridge: Cambridge
University Press. p. 248.

Deacon, Hilary John; Deacon, Janette (1999). Human beginnings in South Africa: uncovering the secrets of the Stone Age. Walnut Creek, Calif. [u.a.]:
Altamira Press.

[53] McBrearty and Brooks 2000

[54] Biological origins of modern humans
[55] McDougall et al. 2005
[56] White et al. 2003
[57] Barham & Mitchell 2008, p. 74
[58] Barham & Mitchell 2008, p. 108
[59] Efraim Lev; Mordechai E. Kislev; Ofer Bar-Yosef (March
2005). Mousterian vegetal food in Kebara Cave, Mt.
Carmel. Journal of Archaeological Science 32 (3): 475
484. doi:10.1016/j.jas.2004.11.006.
[60] Dolores R. Piperno; Ehud Weiss; Irene Holst; Dani Nadel
(5 August 2004). Processing of wild cereal grains in
the Upper Palaeolithic revealed by starch grain analysis.
Nature 430 (7000): 6703. doi:10.1038/nature02734.
PMID 15295598.
[61] Turvey, Samuel T. (2009). Chapter 2: In the shadow of
the megafauna: prehistoric mammal and bird extinctions
across the Holocene. In Turvey, Samuel T. Holocene Extinctions. Oxford Biology. Oxford: Oxford University
Press. pp. 1617
[62] Aczel, Amir D. (2000). The Cave and the Cathedral: How
a Real-Life Indiana Jones and a Research Scholar Decoded the Ancient Art of Man. Hoboken: John Wiley &
Sons Inc. pp. 157158.
[63] Martnez, Antonio Beltrn (1982) [1979]. Rock art of the
Spanish Levant. The Imprint of Man. Cambridge: Cambridge University Press. pp. 4851.
[64] Salvatore Piccolo, Ancient Stones: The Prehistoric Dolmens of Sicily. Abingdon, 2013.

Piccolo, Salvatore (2013). Ancient Stones: The Prehistoric Dolmens of Sicily. Abingdon (UK): Brazen
Head Publishing.
Rogers, Michael J.; Semaw, Sileshi (2009). From
Nothing to Something: The Appearance and Context of the Earliest Archaeological Record. In
Camps i Calbet, Marta; Chauhan, Parth R. Sourcebook of paleolithic transitions: methods, theories,
and interpretations. New York: Springer.
Schick, Kathy D.; Nicholas Toth (1993). Making Silent Stones Speak: Human Evolution and the
Dawn of Technology. New York: Simon & Schuster. ISBN 0-671-69371-9.
Shea, John J. (2010). Stone Age Visiting Cards Revisited: a Strategic Perspective on the Lithic Technology of Early Hominin Dispersal. In Fleagle,
John G.; Shea, John J.; Grine, Frederick E.; Boden,
Andrea L.; Leakey, Richard E,. Out of Africa I: the
First Hominin Colonization of Eurasia. Dordrecht;
Heidelberg; London; New York: Springer. pp. 47
64.

5.9 Further reading

Scarre, Christopher (ed.) (1988). Past Worlds: The
Times Atlas of Archaeology. London: Times Books.
ISBN 0-7230-0306-8.

5.10 External links

Giusepi, Robert A. (2000). The Stone Age. History World International. Retrieved 22 February
2011.

48

CHAPTER 5. STONE AGE

Kowalski, D.R. Stone Age Hand-axes. AerobiologicalEngineering.com. Retrieved 22 February

2011.
Kowalski, D.R. Stone Age Habitats. AerobiologicalEngineering.com. Retrieved 22 February 2011.
PanAfrican Archaeological Association.
trieved 28 February 2011.

Re-

Society of Africanist Archaeologists. Retrieved 3

March 2011.
The ASA. Association of Social Anthropologists
of the UK and Commonwealth.

Chapter 6

Woolly mammoth
The woolly mammoth (Mammuthus primigenius) was a
species of mammoth, the common name for the extinct
elephant genus Mammuthus. The woolly mammoth was
one of the last in a line of mammoth species, beginning
with Mammuthus subplanifrons in the early Pliocene. M.
primigenius diverged from the steppe mammoth, M. trogontherii, about 200,000 years ago in eastern Asia. Its
closest extant relative is the Asian elephant.

change and consequent shrinkage of its habitat, hunting

by humans, or a combination of the two. Isolated populations survived on St. Paul Island until 6,400 years ago
and Wrangel Island until 4,000 years ago. After its extinction, humans continued using its ivory as a raw material, a tradition that continues today. It has been proposed
the species could be recreated through cloning, but this
method is as yet infeasible because of the degraded state
The appearance and behaviour of this species are among of the remaining genetic material.
the best studied of any prehistoric animal because of the
discovery of frozen carcasses in Siberia and Alaska, as
well as skeletons, teeth, stomach contents, dung, and de- 6.1 Taxonomy
piction from life in prehistoric cave paintings. Mammoth
remains had long been known in Asia before they became known to Europeans in the 17th century. The origin
of these remains was long a matter of debate, and often
explained as being remains of legendary creatures. The
mammoth was identied as an extinct species of elephant
by Georges Cuvier in 1796.
The woolly mammoth was roughly the same size as modern African elephants. Males reached shoulder heights
between 2.7 and 3.4 m (9 and 11 ft) and weighed up to
6 tonnes (6.6 short tons). Females averaged 2.62.9 metres (8.59.5 ft) in height and weighed up to 4 tonnes (4.4
short tons). A newborn calf weighed about 90 kilograms
(200 lb). The woolly mammoth was well adapted to the
cold environment during the last ice age. It was covered
in fur, with an outer covering of long guard hairs and a
shorter undercoat. The colour of the coat varied from
dark to light. The ears and tail were short to minimise
frostbite and heat loss. It had long, curved tusks and four
molars, which were replaced six times during the lifetime of an individual. Its behaviour was similar to that
of modern elephants, and it used its tusks and trunk for
manipulating objects, ghting, and foraging. The diet of
the woolly mammoth was mainly grass and sedges. Individuals could probably reach the age of 60. Its habitat
was the mammoth steppe, which stretched across northern Eurasia and North America.
The woolly mammoth coexisted with early humans, who
used its bones and tusks for making art, tools, and
dwellings, and the species was also hunted for food.[1]
It disappeared from its mainland range at the end of the
Pleistocene 10,000 years ago, most likely through climate

A mammoth tusk with Inuit carvings of scenes on the Yukon

River, 19th century.

Remains of various extinct elephants were known by

Europeans for centuries, but were generally interpreted,
based on biblical accounts, as the remains of legendary
creatures such as behemoths or giants. It was also theorised that they were remains of modern elephants that
had been brought to Europe during the Roman Republic, for example the war elephants of Hannibal the Great
and Pyrrhus of Epirus, or animals that had wandered
north.[2] The rst woolly mammoth remains studied by
European scientists were examined by Hans Sloane in
1728 and consisted of fossilised teeth and tusks from
Siberia. Sloane was the rst to recognise that the remains
belonged to elephants.[3] Sloane turned to another biblical explanation for the presence of elephants in the Arctic, asserting that they had been buried during the Great
Flood, and that Siberia had previously been tropical prior
to a drastic climate change.[4] Others interpreted Sloanes
conclusion slightly dierently, arguing the ood had carried elephants from the Tropics to the Arctic. Sloanes

49

50

CHAPTER 6. WOOLLY MAMMOTH

paper was based on travellers descriptions and a few scattered bones collected in Siberia and Britain. He discussed
the question of whether or not the remains were from elephants, but drew no conclusions.[5]
In 1738, Johann Philipp Breyne argued that mammoth
fossils represented some kind of elephant. He could not
explain why a tropical animal would be found in such a
cold area as Siberia, and suggested that they might have
been transported there by the Great Flood.[6] In 1796,
French anatomist Georges Cuvier was the rst to identify
the woolly mammoth remains not as modern elephants
transported to the Arctic, but as an entirely new species.
He argued this species had gone extinct and no longer
existed, a concept that was not widely accepted at the
time.[2][7]

6.1.1

Etymology

the species, since holotype designation was not practised

in Blumenbachs time.[8] In 1828 Joshua Brookes recognised the species was distinct enough to warrant a new
genus, and reclassied it as Mammuthus primigenius.[9]
It is unclear where and how the word mammoth originated. According to the Oxford English Dictionary, it
comes from an old Vogul word mmot 'earth-horn'. [10]
It may be a version of mehemot, the Arabic version of
the biblical word behemoth. Another possible origin is
Estonian, where maa means earth, and mutt means mole.
The word was rst used in Europe during the early 17th
century, when referring to maimanto tusks discovered in
Siberia.[11] Thomas Jeerson, who had a keen interest
in palaeontology, is partially responsible for transforming the word mammoth from a noun describing the prehistoric elephant to an adjective describing anything of
surprisingly large size. The rst recorded use of the word
as an adjective was in a description of a wheel of cheese
(the "Cheshire Mammoth Cheese") given to Jeerson in
1802.[12]

6.1.2 Evolution
The earliest known proboscideans, the clade which contains elephants, existed about 55 million years ago around
the Tethys Sea. The closest known relatives of the Proboscidea are the sirenians and the hyraxes. The family Elephantidae existed six million years ago in Africa
and includes the modern elephants and the mammoths.
Among many now extinct clades, the mastodon is only a
distant relative of the mammoths, and part of the separate
Mammutidae family, which diverged 25 million years before the mammoths evolved.[13] The following cladogram
shows the placement of the genus Mammuthus among
other proboscideans, based on hyoid characteristics:[14]

Comparison between a woolly mammoth (L) and an American

mastodon (R).

Cuviers 1796 comparison between the mandible of a woolly

mammoth and an Indian elephant.

Following Cuviers identication, Johann Friedrich Blumenbach gave the woolly mammoth its scientic name,
Elephas primigenius, in 1799, placing it in the same genus
as the Asian elephant. This name means the rstborn
elephant. Henry Faireld Osborn chose a molar from
Blumenbachs collection as the lectotype specimen for

In 2005, researchers assembled a complete mitochondrial

genome prole of the woolly mammoth, which allowed
them to trace the close evolutionary relationship between
mammoths and Asian elephants, Elephas maximus.[15]
African elephants, Loxodonta africana, branched away
from this clade around 6 million years ago, close to
the time of the similar split between chimpanzees and
humans. Before the publication of the Neanderthal
genome, many researchers expected the rst fully sequenced nuclear genome of an extinct species would be
that of the mammoth.[16] A 2010 study conrmed these
relationships, and suggested the mammoth and Asian ele-

6.2. DESCRIPTION

51

phant lineages diverged 5.87.8 million years ago, while

African elephants diverged from an earlier common ancestor 6.68.8 million years ago.[17] In 2008, much of
the woolly mammoths chromosomal DNA was mapped.
The analysis showed that the woolly mammoth and the
African elephant are 98.55% to 99.40% identical.[18] The
team mapped the woolly mammoths nuclear genome sequence by extracting DNA from the hair follicles of both
a 20,000-year-old mammoth retrieved from permafrost,
and another that died 60,000 years ago.[19] In 2012,
proteins were condently identied for the rst time, collected from a 43,000-year-old woolly mammoth.[20]

Cast of an intermediate form between M. trogontherii and M.

primigenius, M. p. fraasi

entered North America. A 2011 genetic study showed

that two examined specimens of the Columbian mammoth were grouped within a subclade of woolly mammoths. This suggests that the two populations interbred
and produced fertile ospring. A North American form
M. jeersonii, a possible hybrid between Columbian and woolly known as M. jeersonii may be a hybrid between the two
species.[23]
mammoths.
Since many remains of each species of mammoth are
known from several localities, it is possible to reconstruct the evolutionary history of the genus through
morphological studies. Mammoth species can be identied from the number of enamel ridges on their molars;
primitive species had few ridges, and the number increased gradually as new species evolved and replaced
the preceding ones. The crowns of the teeth lengthened
and the skulls became taller to accommodate this. At the
same time, the skulls became shorter from front to back
to minimise the weight.[21] These adaptations were acquired gradually as mammoths turned to more abrasive
food items.[22]
The rst known members of the genus Mammuthus are
the African species M. subplanifrons from the Pliocene,
and M. africanavus from the Pleistocene. The former is
thought to be the ancestor of later forms. Mammoths
entered Europe around 3 million years ago. The earliest type known from there has been named M. rumanus,
which spread across Europe and China. Only its molars
are known, which show that it had 810 enamel ridges.
A population evolved 1214 ridges, splitting o from and
replacing the earlier type, becoming M. meridionalis. In
turn, this species was replaced by the steppe mammoth,
M. trogontherii, with 1820 ridges, which evolved in eastern Asia c. 1 million years ago. Mammoths derived from
M. trogontherii evolved molars with 26 ridges 200,000
years ago in Siberia and became the woolly mammoth, M.
primigenius.[21] The Columbian mammoth, M. columbi,
evolved from a population of M. trogontherii that had

Individuals and populations showing transitional morphologies between each of the mammoth species are
known, and primitive and derived species coexisted as
well until the former disappeared. The dierent species
and their intermediate forms therefore can be termed
"chronospecies". Many intermediate subspecies have
also been proposed, but their validity is uncertain; depending on author, they are either considered primitive forms of an advanced species or advanced forms of
a primitive species.[21] Regional and intermediate subspecies such as M. p. primigenius, M. p. jatzkovi, M.
p. sibiricus, and M. p. fraasi have been proposed.[24]
The St. Paul Island and Wrangel Island populations
were described as dwarf varieties, much smaller than the
mainland woolly mammoth; the Wrangel Island population was also proposed to be a new subspecies, M. p.
vrangeliensis.[25][26] The Wrangel mammoths were isolated for 5000 years, but experienced only a slight loss
of genetic variation.[27]

6.2 Description
The appearance of the woolly mammoth is probably the
best known of any prehistoric animal due to the many
frozen specimens with preserved soft tissue and depictions by contemporary humans in their art. Fully grown
males reached shoulder heights between 2.7 and 3.4 m (9
and 11 ft) and weighed up to 6 tonnes (6.6 short tons).
This is almost as large as extant male African elephants,

52

CHAPTER 6. WOOLLY MAMMOTH

Model at the Royal BC Museum

Cave art from Les Combarelles, France

dle Kolyma mammoth, which was preserved with a complete trunk tip. Unlike the trunk lobes of modern elephants, the upper nger at the tip of the trunk had a long
pointed lobe and was 10 cm (3.9 in) long, while the lower
thumb was 5 cm (2.0 in) and was broader. The trunk
of Dima was 76 cm (2.49 ft) long, whereas the trunk
of the adult Liakhov mammoth was 2 metres (6.6 ft)
long.[31] Few frozen specimens have preserved genitals,
so the gender is usually determined through examination
of the skeleton. Males were generally larger and had more
robust skeletons and tusks. The best indication of sex is
the size of the pelvic girdle, as the birth canal is always
wider in females than in males.[33]

which commonly reach 33.4 m (9.811.2 ft), and is

less than the size of the earlier mammoth species M.
meridionalis and M. trogontherii, and the contemporary
M. columbi. The reason for the smaller size is unknown.
Female woolly mammoths averaged 2.62.9 m (8.59.5
ft) in height and were built more lightly than males, weighing up to 4 tonnes (4.4 short tons). A newborn calf would
have weighed about 90 kg (200 lb). These sizes are deduced from comparison with modern elephants of similar size.[28] Though the mammoths on Wrangel Island
were smaller than those of the mainland, their size varied, and they were not small enough to be considered
dwarves.[29] It has been claimed that the last woolly
mammoth populations decreased in size and increased 6.2.1
their sexual dimorphism, but this was dismissed in a 2012
study.[30]
Woolly mammoths had several adaptations to the cold,
most noticeably the layer of fur covering all parts of the
body. Other adaptations to cold weather include ears that
are far smaller than those of modern elephants; they were
about 38 cm (15 in) long and 1828 cm (7.111.0 in)
across, and the ear of the 612 month old frozen calf
Dima was under 13 cm (5.1 in) long. The small ears reduced heat loss and frostbite, and the tail was short for the
same reason, only 36 cm (14 in) long in the Berezovka
mammoth. The tail contained 21 vertebrae, whereas the
tails of modern elephants contain 2833. Their skin was
no thicker than that of present-day elephants, between
1.25 and 2.5 cm (0.49 and 0.98 in). They had a layer of
fat up to 10 cm (3.9 in) thick under the skin, which helped
to keep them warm. Woolly mammoths had broad aps
of skin under their tails which covered the anus; this is
also seen in modern elephants.[31]
Other characteristic features depicted in cave paintings
include a large, high, single-domed head and a sloping
back with a high shoulder hump, resulting from long
spinous processes on the neck vertebrae. These features were not present in juveniles, which had concave
backs like African elephants.[32] Another feature shown
in cave paintings was conrmed by the discovery of a
frozen specimen in 1924, an adult nicknamed the Mid-

Coat

Fur in Naturhistorisches Museum, Vienna

The coat consisted of an outer layer of long, coarse guard

hair, which was 30 cm (12 in) on the upper part of the
body, up to 90 cm (35 in) in length on the anks and underside, and 0.5 mm (0.020 in) in diameter, and a denser
inner layer of shorter, slightly curly under-wool, up to 8
cm (3.1 in) long and 0.05 mm (0.0020 in) in diameter.
The hairs on the upper leg were up to 38 cm (15 in) long,
and those of the feet were 15 cm (5.9 in) long, reaching
the toes. The hairs on the head were relatively short, but
longer on the underside and the sides of the trunk. The
tail was extended by coarse hairs up to 60 cm (24 in) long,

6.2. DESCRIPTION

53

which were thicker than the guard hairs. It is likely that

the woolly mammoth moulted seasonally, and that the
heaviest fur was shed during spring. Since mammoth carcasses were more likely to be preserved during autumn,
it is possible that only the winter coat has been preserved
in frozen specimens. Modern elephants have much less
hair, though juveniles have a more extensive covering of
hair than adults.[34] Comparison between the over-hairs
of woolly mammoths and extant elephants show that they
did not dier much in overall morphology.[35] Woolly
mammoths had numerous sebaceous glands in their skin,
which secreted oils into their hair; this would have improved the wools insulation, repelled water, and given the
fur a glossy sheen.[36]
Preserved woolly mammoth fur is orange-brown, but this
is believed to be an artefact from the bleaching of pigment during burial. The amount of pigmentation varied from hair to hair and also within each hair.[31] A
2006 study sequenced the Mc1r gene (which inuences
hair colour in mammals) from woolly mammoth bones.
Two alleles were found: a dominant (fully active) and
a recessive (partially active) one. In mammals, recessive Mc1r alleles results in light hair. Mammoths born Specimen with asymmetrical tusks, the Smithsonian Museum
with at least one copy of the dominant allele would have
had dark coats, while those with two copies of the recessive allele would have had light coats.[37] A 2011 study tusks, but there is no fossil evidence that any adult woolly
showed that light individuals would have been rare.[38] A mammoths lacked them.[40]
2014 study instead indicated that the colouration of an
individual varied from non-pigmented on the overhairs,
bi coloured, non-pigmented and mixed red-brown guard
hairs, and non-pigmented underhairs, which would give a
light overall appearance.[39]

6.2.2

Dentition

Woolly mammoths had very long tusks, which were more

curved than those of modern elephants. The largest
known male tusk is 4.2 m (14 ft) long and weighs 91 kg
(201 lb), but 2.42.7 m (7.98.9 ft) and 45 kg (99 lb) was
a more typical size. Female tusks averaged at 1.51.8 m
(4.95.9 ft) and weighed 9 kg (20 lb). About a quarter
of the length was inside the sockets. The tusks grew spirally in opposite directions from the base and continued
in a curve until the tips pointed towards each other. In
this way, most of the weight would have been close to the
skull, and there would be less torque than with straight
tusks. The tusks were usually asymmetrical and showed
considerable variation, with some tusks curving down instead of outwards and some being shorter due to breakage. Calves developed small milk tusks a few centimetres
long at six months old, which were replaced by permanent tusks a year later. Tusk growth continued throughout
life but became slower as the animal reached adulthood.
The tusks grew by 2.515 cm (0.985.91 in) each year.
Some cave paintings show woolly mammoths with small
or no tusks, but it is unknown whether this reected reality or was artistic license. Female Asian elephants have no

Molar from font de Champdamoy, France

Woolly mammoths had four functional molar teeth at a

time, two in the upper jaw and two in the lower. 23 cm
(9.1 in) of the crown was within the jaw, and 2.5 cm (1 in)
was above. The crown was continually pushed forwards
and up as it wore down, comparable to a conveyor belt.
The teeth had up to 26 separated ridges of enamel, which
were themselves covered in prisms that were directed
towards the chewing surface. These were quite wear resistant and kept together by cementum and dentine. A
mammoth had six sets of molars throughout a lifetime,
which were replaced ve times, though a few specimens
with a seventh set are known. The latter condition could
extend the lifespan of the individual, unless the tooth only
consisted of a few plates. The rst molars were about
the size of those of a human, 1.3 cm (0.51 in), the third
were 15 cm (6 in) 15 cm (5.9 in) long, and the sixth were
about 30 cm (1 ft) long and weighed 1.8 kg (4 lb). The
molars grew larger and contained more ridges with each

54
replacement.[41]
Distortion in the molars is the most common health problem found in woolly mammoth fossils. Sometimes the
replacement was disrupted, and the molars were pushed
into abnormal positions, but some animals are known
to have survived this. Teeth from Britain showed that
2% of specimens had periodontal disease, with half of
these containing caries. The teeth also sometimes had
cancerous growths.[42]

6.3 Palaeobiology

CHAPTER 6. WOOLLY MAMMOTH

supported by fossil assemblages and cave paintings showing groups. It is therefore probable that most of their
other social behaviour was similar to those of modern
elephants. Accumulations of modern elephant remains
have been termed "elephants graveyards", as these sites
were erroneously thought to be where old elephants went
to die. Similar accumulations of woolly mammoth bones
have been found; it is thought these are the result of individuals dying near or in the rivers over thousands of
years, and their bones eventually being brought together
by the streams, or due to animals being mired in mud.
Some accumulations are also thought to be the remains
of herds that died together at the same time, perhaps due
to ooding.[45]

Restoration of a herd walking near the Somme River by Charles

R. Knight

Adult woolly mammoths could eectively defend themselves from predators with their tusks, trunks and size, but
juveniles and weakened adults were vulnerable to pack
hunters such as wolves, cave hyenas and large felines. The
tusks may also have been used in intra-species ghting,
such as territorial ghts or ghts over mates. Because
of their curvature, the tusks were not suitable for stabbing, but may have been used for hitting, as indicated
by injuries to some fossil shoulder blades. As in modern elephants, the sensitive and muscular trunk worked
as a limb-like organ with many functions. It was used
for manipulating objects, and in social interactions. The
very long hairs on the tail probably compensated for the
shortness of the tail, enabling its use as a yswatter, similar to the tail on modern elephants.[43] As in reindeer and
musk oxen, the haemoglobin of the woolly mammoth was
adapted to the cold, with three mutations to improve oxygen delivery around the body and prevent freezing. This
feature may have helped the mammoths to live in high
latitudes.[44]

Leg and foot of the Yukagir mammoth

Trackways made by a woolly mammoth herd 11,300

11,000 years ago, have been found in the St. Mary Reservoir in Canada, showing that there were in this case almost equal numbers of adults, sub-adults and juveniles.
The adults had a stride of 2 m (6.6 ft), and the juveniles ran to keep up.[46] The well-preserved foot of the
adult male "Yukagir mammoth" shows that the soles of
the feet contained many cracks that would have helped in
gripping surfaces during locomotion. Like modern elephants, woolly mammoths walked on their toes and had
large, eshy pads behind the toes.[31]
Evidence of several dierent bone diseases has been
found in woolly mammoths. The most common of
these diseases was osteoarthritis, found in 2% of specimens. One specimen from Switzerland had several fused
vertebrae as a result of this condition. The Yukagir
mammoth had suered from spondylitis in two vertebrae, and osteomyelitis is also known from some specimens. Several specimens have healed bone fractures,
showing that the animals had survived these injuries.[47]
Parasitic ies and protozoa were identied in the gut of
the calf Dima.[48]

6.3.1 Diet
Food at various stages of digestion has been found in the
intestines of several woolly mammoths, giving a good picture of their diet. Woolly mammoths sustained themLike modern elephants, woolly mammoths were likely selves on plant food, mainly grass and sedges, which were
very social and lived in matriarchal family groups. This is supplemented with herbaceous plants, owering plants,
Mounted family group

6.3. PALAEOBIOLOGY

55

Mandibles at various growth stages, Naturalis, Leiden

wards while opening, which made the sharp enamel ridges

cut across each other and grind the food. The ridges were
wear-resistant to enable the animal to chew large quantities of food, which often contained grit. Woolly mammoths may have used their tusks as shovels to clear snow
from the ground and reach the vegetation buried below,
and to break ice to drink. This is indicated on many preThe frozen calf Lyuba which still had food in its stomach
served tusks by at, polished sections up to 30 centimetres (12 in) long on the part of the surface that would have
reached the ground. The tusks were also used for obshrubs, mosses, and tree matter. The composition and taining food in other ways, such as digging up plants and
[54]
exact varieties diered from location to location. Woolly stripping o bark.
mammoths needed a varied diet to support their growth,
like modern elephants. An adult of six tonnes would need
to eat 180 kg (397 lb) daily, and may have foraged as 6.3.2 Growth and reproduction
long as twenty hours every day. The two-ngered tip of
the trunk was probably adapted for picking up the short
grasses of the last ice age (Quaternary glaciation, 2.58
million years ago to present) by wrapping around them,
whereas modern elephants curl their trunks around the
longer grass of their tropical environments. The trunk
could also be used for pulling o large grass tufts, delicately picking buds and owers, and tearing o leaves
and branches where trees and shrubs were present. The
Yukagir mammoth had ingested plant matter that contained spores of dung fungus.[49] Isotope analysis shows
that woolly mammoths fed mainly on C3 plants, unlike
horses and rhinos.[50]
Scientists identied milk in the stomach and faecal matter in the intestines of the mammoth calf "Lyuba".[51]
The faecal matter may have been eaten by Lyuba to
promote development of the intestinal microbes necessary for digestion of vegetation, as is the case in modern
elephants.[52] An isotope analysis of woolly mammoths
from Yukon, Canada, showed that the young nursed for at
least three years, and were weaned and gradually changed
to a diet of plants when they were two to three years old.
This is later than in modern elephants and may be due to
a higher risk of predator attack or diculty in obtaining
food during the long periods of winter darkness in high
latitudes.[53]
The molars were adapted to their diet of coarse tundra
grasses, with more enamel plates and a higher crown than
their earlier, southern relatives. The woolly mammoth
chewed its food by using its powerful jaw muscles to move
the mandible forwards and close the mouth, then back-

Cross sectioned tusk with growth rings

The age of a woolly mammoth can be determined by

counting the growth rings of its tusks when viewed in
cross section. Each major line represents a year, and
weekly and daily ones can be found in between. Dark
bands correspond to summers, and it is therefore possible to determine the season in which a mammoth died.
The growth of the tusks slowed when it became harder to
forage, for example during disease, when a male was banished from the herd, and during periods of severe glaciation. Woolly mammoths continued growing past adulthood, like other elephants. Unfused limb bones show that
males grew until they reached the age of 40, and females
grew until they were 25. The frozen calf Dima was 90
cm (35 in) tall when it died at the age of 612 months.

56
At this age, the second set of molars would be in the process of erupting, and the rst set would be worn out at
18 months of age. The third set of molars lasted for ten
years, and this process was repeated until the nal, sixth
set emerged when the animal was 30 years old. A woolly
mammoth could probably reach the age of 60, like modern elephants of the same size. By then the last set of
molars would be worn out, the animal would be unable to
chew and feed, and it would die of starvation.[55]

CHAPTER 6. WOOLLY MAMMOTH

stretched across northern Asia, many parts of Europe,
and the northern part of North America during the last ice
age. It was similar to the grassy steppes of modern Russia, but the ora was more diverse, abundant, and grew
faster. Grasses, sedges, shrubs, and herbaceous plants
were present, and scattered trees were mainly found in
southern regions. This habitat was not dominated by ice
and snow, as is popularly believed, since these regions
are thought to have been high-pressure areas at the time.
The habitat of the woolly mammoth also supported other
grazing herbivores such as the woolly rhinoceros, wild
horses and bison.[60] A 2014 study concluded that forbs (a
group of herbaceous plants) were more important in the
steppe-tundra than previously acknowledged, and that it
was a primary food source for the ice-age megafauna.[61]

Head of the Yukagir mammoth

The best preserved head of a frozen adult specimen, that

of a male nicknamed the Yukagir mammoth, shows
that woolly mammoths had temporal glands between the
ear and the eye.[56] This feature indicates that, like bull
elephants, male woolly mammoths also entered "musth",
a period of heightened aggressiveness. The glands are
used especially by males to produce an oily substance with
a strong smell called temporin. Their fur may have helped
in spreading the scent further.[57]
Examination of preserved calves show that they were
all born during spring and summer, and since modern
elephants have gestation periods of 2122 months, it
is probable that the mating season was from summer
to autumn.[58] 15N isotopic analysis of the teeth of
Lyuba has demonstrated their prenatal development,
and indicates its gestation period was similar to that of
a modern elephant, and that it was born in spring.[59]

6.4 Distribution and habitat

Possible distribution during the last glacial period, based on locations of fossil nds

The habitat of the woolly mammoth is known as

"mammoth steppe" or tundra steppe. This environment

Restoration of a group in late Pleistocene northern Spain, by

Mauricio Antn

The southernmost woolly mammoth specimen known is

from the Shandong province of China, and is 33,000
years old.[62] The southernmost European remains are
from the Depression of Granada in Spain and are of
roughly the same age.[63] DNA studies have helped determine the phylogeography of the woolly mammoth. A
2008 DNA study showed there were two distinct groups
of woolly mammoths: one that went extinct 45,000 years
ago and another one that went extinct 12,000 years ago.
The two groups are speculated to be divergent enough
to be characterised as subspecies. The group that went
extinct earlier stayed in the middle of the high Arctic,
while the group with the later extinction had a much
wider range.[64] Recent stable isotope studies of Siberian
and New World mammoths have shown there were also
dierences in climatic conditions on either side of the
Bering land bridge, with Siberia being more uniformly
cold and dry throughout the Late Pleistocene.[65] During the Younger Dryas age, woolly mammoths briey
expanded into north-east Europe, whereafter the mainland populations became extinct.[66] A 2008 genetic study
showed that some of the woolly mammoths that entered
North America through the Bering land bridge from Asia
migrated back about 300,000 years ago and had replaced
the previous Asian population by about 40,000 years ago,
not long before the entire species went extinct.[67] Woolly
mammoths have been found in the same locations as those
of the Columbian mammoth in North America, but it is
unknown whether the two species were sympatric. The
woolly mammoth may have entered these southern ar-

6.5. RELATIONSHIP WITH HUMANS

57

eas during times when Columbian mammoth populations day, more than ve hundred depictions of woolly mamwere absent.[68]
moths are known, in media ranging from cave paintings
and engravings on the walls of 46 caves in Russia, France
and Spain to engravings and sculptures (termed "portable
art") made from ivory, antler, stone and bone. Cave paint6.5 Relationship with humans
ings of woolly mammoths exist in several styles and sizes.
The French Rougnac cave has most depictions, 159,
and some of the drawings are more than 2 metres (6.5
ft) in length. Other notable caves with mammoth depictions are the Chauvet Cave, Les Combarelles Cave,
and Font-de-Gaume.[70] A depiction in the Cave of El
Castillo may instead show Palaeoloxodon, the straighttusked elephant.[71]

Woolly mammoth carved in ivory, discovered by douard Lartet

in 1864

Portable art can be more accurately dated than cave art

since it is found in the same deposits as tools and other
ice age artefacts. The largest collection of portable mammoth art, consisting of 62 depictions on 47 plaques, was
found in the 1960s at an excavated open-air camp near
Gnnersdorf in Germany. There does not seem to be a
correlation between the number of mammoths depicted
and the species that were most often hunted, since reindeer bones are the most frequently found animal remains
at the site. Two spear throwers shaped as woolly mammoths have also been found in France.[70] Some portable
mammoth depictions may not have been produced where
they were discovered, but could have moved around by
ancient trading.[71]

Modern humans coexisted with woolly mammoths during the Upper Palaeolithic period when they entered Europe from Africa between 30,000 and 40,000 years ago.
Prior to this, Neanderthals had coexisted with mammoths
during the Middle Palaeolithic. Woolly mammoths were
very important to ice age humans, and human survival
may have depended on the mammoth in some areas. Evidence for such coexistence was not recognised until the
19th century. William Buckland published his discovery of the Red Lady of Paviland skeleton in 1823, which
was found in a cave alongside woolly mammoth bones, 6.5.1
but he mistakenly denied that these were contemporaries.
In 1864, douard Lartet found an engraving of a woolly
mammoth on a piece of mammoth ivory in the Abri de
la Madeleine cave in Dordogne, France. This was the
rst widely accepted evidence for the coexistence of humans with prehistoric extinct animals and is the rst contemporary depiction of such a creature known to modern
science.[69]

Exploitation

Reconstructed bone hut, based on nds in Mezhyrich

Cro-Magnon artists painting mammoths in Font-de-Gaume, by

Charles R. Knight

The woolly mammoth is the third most depicted animal in

ice age art, after horses and bison, and these images were
produced between 35,000 and 11,500 years ago. To-

Woolly mammoth bones were used as construction material for dwellings by both Neanderthals and modern humans during the ice age. More than 70 such dwellings
are known, mainly from the Russian Plain. The bases of
the huts were circular, and ranged from 8 to 24 square
metres (86 to 258 sq ft). The arrangement of dwellings
varied, and ranged from 1 m (3.3 ft) to 20 m (66 ft) apart,
depending on location. Large bones were used as foundations for the huts, tusks for the entrances, and the roofs
were probably skins held in place by bones or tusks. Some
huts had oors that extended 40 cm (16 in) below ground.
Some huts included replaces, which used bones as fuel,
probably because wood was scarce. It is possible that

58

CHAPTER 6. WOOLLY MAMMOTH

some of the bones used for materials came from mam- hunted intensively, but perhaps mainly when ivory was
moths killed by humans, but the state of the bones, and needed.[77]
the fact that bones used to build a single dwelling varied
by several thousands of years in age, suggests that they
were collected remains of long-dead animals. Woolly 6.6 Extinction
mammoth bones were also made into various tools, furniture, and musical instruments. Large bones, such as
shoulder blades, were also used to cover dead human bodies during burial.[72]

16,500 year old mammoth spear thrower from France

The Venus of Brassempouy

Woolly mammoth ivory was used to create art objects.

Several Venus gurines, including the Venus of Brassempouy and the Venus of Lespugue, were made from this
material. Weapons made from ivory, such as daggers,
spears, and a boomerang, are also known. To be able
to process the ivory, the large tusks had to be chopped,
chiselled and split into smaller, more manageable pieces.
Some ivory artefacts show that tusks had been straightened, and it is unknown how this was achieved.[73]
Several woolly mammoth specimens show evidence of
being butchered by humans, which is indicated by breaks,
cut-marks, and associated stone tools. It is not known
how much prehistoric humans relied on woolly mammoth
meat, since there were many other large herbivores available. Many mammoth carcasses may have been scavenged by humans rather than hunted. Some cave paintings show woolly mammoths in structures interpreted as
pitfall traps. Few specimens show direct, unambiguous
evidence of having been hunted by humans. A Siberian
specimen with a spearhead embedded in its shoulder
blade shows that a spear had been thrown at it with great
force.[74] A specimen from the Mousterian age of Italy
shows evidence of spear hunting by Neanderthals.[75] The
juvenile specimen nicknamed Yuka is the rst frozen
mammoth with evidence of human interaction. It shows
evidence of having been killed by a large predator, and of
having been scavenged by humans shortly after. Some of
its bones had been removed, and were found nearby.[76]
A site near the Yana River in Siberia has revealed several specimens with evidence of human hunting, but the
nds were interpreted to show that the animals were not

Most woolly mammoth populations disappeared during

the late Pleistocene and early Holocene, alongside most of
the Pleistocene megafauna, during the Quaternary extinction event.[78] Scientists are divided over whether hunting or climate change, which led to the shrinkage of its
habitat, was the main factor that contributed to the extinction of the woolly mammoth, or whether it was due
to a combination of the two. Whatever the cause, large
mammals are generally more vulnerable than smaller ones
due to their smaller population size and low reproduction rates. Dierent woolly mammoth populations did
not die out simultaneously across their range, but gradually went extinct over time. The last mainland population existed in the Kyttyk Peninsula of Siberia 9,650
years ago.[79] A small population of woolly mammoths
survived on St. Paul Island, Alaska, until 6,400 years
ago.[26][80][81] The last known population remained on
Wrangel Island in the Arctic Ocean until 4,000 years
ago.[82][83][84] Genetic evidence implies the extinction of
this nal population was sudden, rather than the culmination of a gradual decline;[84] the disappearance coincides roughly in time with the rst evidence for humans on
the island.[85] The woolly mammoths of eastern Beringia
(modern Alaska and Yukon) had similarly died out about
13,300 years ago, soon (roughly 1000 years) after the rst
appearance of humans in the area, which parallels the fate
of all the other late Pleistocene proboscids (mammoths,
gomphotheres and mastodons), as well as most of the rest
of the megafauna, of the Americas.[86] In contrast, the
St. Paul Island mammoth population apparently died out
prior to human arrival due to habitat shrinkage resulting
from the post-ice age sea level rise.[86]
A 2008 study estimated that changes in climate
shrank suitable mammoth habitat from 7,700,000 km2
(3,000,000 sq mi) 42,000 years ago to 800,000 km2
(310,000 sq mi) 6,000 years ago.[87][88] Woolly mammoths survived an even greater loss of habitat at the end
of the Saale glaciation 125,000 years ago, and it is likely

6.7. FROZEN SPECIMENS

Cast of the Hebior Mammoth specimen, which bears

tool/butcher marks

that humans hunted the remaining populations to extinction at the end of the last glacial period.[89][90] Studies
of an 11,30011,000 year old trackway in southwestern
Canada showed that M. primigenius was in decline while
coexisting with humans, since far fewer tracks of juveniles were identied than would be expected in a normal
herd.[46]

59
fewer nds in the latter. Such remains are mostly found
above the Arctic Circle, in permafrost. It appears that
soft tissue was less likely to be preserved between 30,000
and 15,000 years ago, perhaps because the climate was
milder during that period. Most specimens have partially
degraded prior to discovery, due to exposure or to being scavenged. This "natural mummication" required
the animal to have been buried rapidly in liquid or semisolids such as silt, mud and icy water, which then froze.[92]
The presence of undigested food in the stomach and seed
pods still in the mouth of many of the specimens suggests neither starvation nor exposure are likely. The maturity of this ingested vegetation places the time of death
in autumn rather than in spring, when owers would be
expected.[93] The animals may have fallen through ice
into small ponds or potholes, entombing them. Many
are certainly known to have been killed in rivers, perhaps
through being swept away by oods. In one location, by
the Berelekh River in Yakutia in Siberia, more than 8,000
bones from at least 140 mammoths have been found in
a single spot, apparently having been swept there by the
current.[94]

A 2010 study suggests that the decline of the woolly

mammoth could have increased temperatures by up to
0.2 C (0.36 F) at high latitudes in the northern hemisphere. Mammoths frequently ate birch trees, creating a
grassland habitat. With the disappearance of mammoths,
birch forests, which absorb more sunlight than grasslands,
expanded, leading to regional warming.[91]

6.7 Frozen specimens

Illustration of the Adams mammoth skeleton with outward
curving tusks, 1815

Early 19th century interpretation of the "Adams mammoth" carcass prior to excavation

Woolly mammoth fossils have been found in many dierent types of deposits, including former rivers and lakes,
and also in "Doggerland" in the North Sea, which was
dry at times during the ice age. Such fossils are usually fragmentary and contain no soft tissue. Apart from
frozen remains, the only soft tissue known is from a specimen that was preserved in a petroleum seep in Starunia,
Poland. Frozen remains of woolly mammoths have been
found in the northern parts of Siberia and Alaska, with far

Between 1692 and 1806, only four descriptions of frozen

mammoths were published in Europe. None of the remains of those ve were preserved, and no complete
skeleton was recovered during that time.[95] While frozen
woolly mammoth carcasses had been excavated by Europeans as early as 1728, the rst fully documented specimen was discovered near the delta of the Lena River
in 1799 by Ossip Schumachov, a Siberian hunter.[96]
Schumachov let it thaw until he could retrieve the tusks
for sale to the ivory trade. While in Yakutsk in 1806,
Michael Friedrich Adams heard about the frozen mammoth. Upon arrival at the location, Adams discovered
that wild animals had eaten most of the organs and esh
of the mammoth, including the trunk. He examined the
carcass and realised what was left would still be the most
complete mammoth recovered by that time. Adams recovered the entire skeleton, apart from the tusks, which
Shumachov had already sold, and one foreleg, most of
the skin, and nearly 18 kg (40 lb) of hair. During his return voyage he purchased a pair of tusks that he believed

60

CHAPTER 6. WOOLLY MAMMOTH

were the ones that Shumachov had sold. Adams brought

it to the Zoological Museum of the Zoological Institute of
the Russian Academy of Sciences, and the task of mounting the skeleton was given to Wilhelm Gottlieb Tilesius.[5]
The Kunstkamera, the museum established by Peter the
Great, contained the skeleton of an Indian elephant that
could be used as reference.[97] This was one of the rst attempts at reconstructing the skeleton of an extinct animal.
Most of the reconstruction is correct, but Tilesius placed
each tusk in the opposite socket, so that they curved outward instead of inward. The error was not corrected until
1899, and the correct placement of mammoth tusks was
still a matter of debate into the twentieth century.[98][99]
Dima, a frozen calf about seven months old

A third of this model is covered with the skin of the Berezovka

mammoth, Museum of Zoology, St. Petersburg

The 1901 excavation of the Berezovka mammoth is the

best documented of the early nds. It was discovered by
the Berezovka River, and the Russian authorities nanced
its excavation. Its head was exposed, and the esh had
been scavenged. The animal still had grass between its
teeth and on the tongue, showing that it had died suddenly. The entire expedition took 10 months, and the
specimen had to be cut to pieces before it could be transported to St. Petersburg. It was identied as a 3540
year old male, which had died 35,000 years ago. One of
its shoulder blades was broken, which may have happened
when it fell into a crevasse.[93]
By 1929, the remains of 34 mammoths with frozen soft
tissues (skin, esh, or organs) had been documented.
Only four of them were relatively complete. Since then,
about that many more have been found. In most cases, the
esh showed signs of decay before its freezing and later
desiccation.[100] Since 1860, Russian authorities have offered rewards of up to .1000 for nds of frozen woolly
mammoth carcasses. Often such nds were kept secret
due to superstition. Several carcasses have been lost because they were not reported, and one was fed to dogs.
In more recent years, scientic expeditions have been
devoted to nding carcasses instead of relying solely on
chance encounters.[101]

the Kolyma River in northeastern Siberia. This specimen weighed approximately 100 kg (220 lb) at death
and was 104 cm (41 in) high and 115 cm (45 in) long.
Radiocarbon dating determined that Dima died about
40,000 years ago. Its internal organs are similar to those
of modern elephants, but its ears are only one-tenth the
size of those of an African elephant of similar age. A less
complete juvenile, nicknamed Mascha, was found on
the Yamal Peninsula in 1988. It was 34 months old, and
a laceration on its right foot may have been the cause of
death. It is the westernmost frozen mammoth found.[102]
In 1997, a piece of mammoth tusk was discovered protruding from the tundra of the Taymyr Peninsula in
Siberia, Russia. In 1999, this 20,380 year old carcass and
25 tons of surrounding sediment were transported by an
Mi-26 heavy lift helicopter to an ice cave in Khatanga.
The specimen was nicknamed the Jarkov mammoth.
In October 2000, the careful defrosting operations in this
cave began with the use of hairdryers to keep the hair and
other soft tissues intact.[103][104]

Frozen calf nicknamed Mascha

In 2007, the carcass of a female calf nicknamed Lyuba

was discovered near the Yuribey River, where it had
been buried for 41,800 years.[52][105] By cutting a section
through a molar and analysing its growth lines, they found
that the animal had died at the age of one month.[59] The
mummied calf weighed 50 kg (110 lb), was 85 cm (33
In 1977, the well-preserved carcass of a seven- to eight- in) high and 130 cm (51 in) in length.[106][107] At the time
month-old woolly mammoth calf named Dima was dis- of discovery, its eyes and trunk were intact and some fur
covered. This carcass was recovered near a tributary of remained on its body. Its organs and skin are very well

6.7. FROZEN SPECIMENS

61

preserved.[108] Lyuba is believed to have been suocated by mud in a river that its herd was crossing.[52][109]
After death, its body may have been colonised by bacteria that produce lactic acid, which pickled it, preserving
the mammoth in a nearly pristine state.[52]

Skeleton cast of a calf, North American Museum of Ancient Life

Skull and mandible of the frozen calf Yuka

In 2012, a juvenile was found in Siberia, which had manmade cut marks. Scientists estimated its age at death to
be 2.5 years, and nicknamed it Yuka. Its skull and
pelvis had been removed prior to discovery, but were
found nearby.[110][76] Another mammoth discovery was
reported in October 2012, when it was excavated on the
Taymyr Peninsula. It was dated to 30,000 years old. It
was named Zhenya after the boy who found it.[111]

elephantmammoth hybrid, and the process would have

to be repeated so more hybrids could be used in breeding.
After several generations of cross-breeding these hybrids,
an almost pure woolly mammoth would be produced. The
fact that sperm cells of modern mammals are potent for
15 years at most after deep-freezing is a hindrance to
this method.[113] In one case, an Asian elephant and an
African elephant produced a live calf named Motty, but
In 2013, a well preserved carcass was found on Maly
it died of defects at less than two weeks old.[114]
Lyakhovsky Island, one of the islands in the New Siberian
Islands archipelago, a female between 50 and 60 years old In 2008, a Japanese team found usable DNA in the
at the time of death. The carcass contained well preserved brains of mice that had been frozen for 16 years. They
muscular tissue. When it was extracted from the ice, liq- hope to use similar methods to nd usable mammoth
uid blood spilled from the abdominal cavity. The nd- DNA.[115] In 2009, the Pyrenean Ibex (a subspecies of
ers interpreted this as indicating woolly mammoth blood the Spanish ibex) was the rst extinct animal to be cloned
back to life; the clone lived for only seven minutes bepossessed anti-freezing properties.[112]
fore dying of lung defects.[116] As the woolly mammoth
genome has been mapped, it may be possible to recreate a complete set of woolly mammoth chromosomes in
6.7.1 Recreating the species
the future by adding mammoth-only sequences to eleThe existence of preserved soft tissue remains and DNA phant chromosomes.[117] If the process is ever successful,
of woolly mammoths has led to the idea that the species there are plans to introduce cloned woolly mammoths to
could be recreated by scientic means. Two methods Pleistocene Park, a wildlife reserve in Siberia.[118]
have been proposed to achieve this. The rst is cloning, Mammoth expert Adrian Lister questions the ethics of
which would involve removal of the DNA-containing such recreation attempts. In addition to the technical
nucleus of the egg cell of a female elephant, and replace- problems, he notes that there is not much habitat left that
ment with a nucleus from woolly mammoth tissue. The would be suitable for woolly mammoths. Because the
cell would then be stimulated into dividing, and inserted species was social and gregarious, creating a few specback into a female elephant. The resulting calf would imens would not be ideal. He also notes that the time
have the genes of the woolly mammoth, although its fetal and resources required would be enormous, and that the
environment would be dierent. To date, even the most scientic benets would be unclear; he suggests these reintact mammoths have had little usable DNA because of sources should instead be used to preserve extant elephant
their conditions of preservation. There is not enough to species which are endangered.[113] A 2014 article about
guide the production of an embryo.[113]
potential cloning also questioned the ethics of using eleThe second method involves articially inseminating an phants as surrogate mothers, as most embryos would not
elephant egg cell with sperm cells from a frozen woolly survive, and noted that it would be impossible to know
mammoth carcass. The resulting ospring would be an the exact needs of a resurrected calf.[119]

62

6.8 Cultural signicance

CHAPTER 6. WOOLLY MAMMOTH

to get it through customs. Mammoth ivory looks similar to elephant ivory, but the former is browner and the
Schreger lines are coarser in texture.[121] In the 21st century, global warming has made access to Siberian tusks
easier, since the permafrost thaws more quickly, exposing the mammoths embedded within it.[122]

Peter III of Russia carved in mammoth ivory

The woolly mammoth has remained culturally signicant

long after its extinction. Indigenous peoples of Siberia
had long found what are now known to be woolly mammoth remains, collecting their tusks for the ivory trade.
Native Siberians believed woolly mammoth remains to be
those of giant mole-like animals that lived underground
and died when burrowing to the surface.[6] Woolly mammoth tusks had been articles of trade in Asia long before Europeans became acquainted with them. Gyk,
the 13th-century Khan of the Mongols, is reputed to have
sat on a throne made from mammoth ivory.[95] Siberian
mammoth ivory is reported to have been exported to Russia and Europe in the 10th century. The rst Siberian
ivory to reach western Europe was brought to London in
1611.[120]

Le Mammouth by Paul Jamin, 1885

Stories abound about frozen woolly mammoth meat that

was consumed once defrosted, especially that of the
Berezovka mammoth, but most of these are considered
dubious. The carcasses were in most cases decayed, and
the stench so unbearable that only wild scavengers and
the dogs accompanying the nders showed any interest
in the esh. It appears that such meat was once recommended against illness in China, and Siberian natives have
occasionally cooked the meat of frozen carcasses they
When Russia occupied Siberia, the ivory trade grew discovered.[123]
and became a widely exported commodity, with huge
amounts being excavated for this market. From the 19th
century and onwards, woolly mammoth ivory became a 6.8.1 Cryptozoology
highly prized commodity, used as raw material for many
dierent products. Today it is still in great demand as a There have been occasional claims that the woolly mamreplacement for the now-banned export of elephant ivory, moth is not extinct, and that small isolated herds might
and has been referred to as white gold. Local dealers survive in the vast and sparsely inhabited tundra of the
estimate that there are 10 million mammoths still frozen Northern Hemisphere. In the 19th century, several rein Siberia, and conservationists have suggested that this ports of large shaggy beasts were passed on to the Ruscould help save the living species of elephants from ex- sian authorities by Siberian tribesmen, but no scientic
tinction. Elephants are hunted by poachers for their ivory, proof ever surfaced. A French charg d'aaires workbut if this could instead be supplied by the already ex- ing in Vladivostok, M. Gallon, said in 1946 that in 1920
tinct mammoths, the demand could instead be met by he had met a Russian fur-trapper who claimed to have
these. Trade in elephant ivory has been forbidden in seen living giant, furry elephants deep into the taiga.
most places following the 1989 Lausanne Conference, but Gallon added that the fur-trapper had not heard of mamdealers have been known to label it as mammoth ivory moths before.[124] Due to the large area of Siberia, it can-

6.9. REFERENCES
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Mammoth?".
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[90] Burney, D.; Flannery, T. (2005).
Fifty millennia of catastrophic extinctions after human contact.
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[93] Pzenmayer, E. W. (1939). Siberian Man and Mammoth.
London: Blackie and Son. pp. 4661.
[94] Vereshchagin, N. K. (2009). The mammoth cemeteries of north-east Siberia. Polar Record 17 (106): 3.
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[95] Tolmacho, I. P. (1929). The carcasses of the mammoth and rhinoceros found in the frozen ground of
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[96] Adams, M. (1808). Some Account of a Journey to the
Frozen-Sea, and of the Discovery of the Remains of a
Mammoth. The Philadelphia Medical and Physical Journal 3: 120137.
[97] Tilesio, W. G. (1815). De skeleto mammonteo Sibirico
ad maris glacialis littora anno 1807 eosso, cui praemissae
Elephantini generis specierum distinctiones. Mmoires
de l'Acadmie Impriale des Sciences de St. Ptersbourg
(in Latin) 5: 406514.
[98] Cohen, C. (2002). The Fate of the Mammoth: Fossils,
Myth, and History. University of Chicago Press. p. 113.
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[99] Pzenmayer, E. (1907). A Contribution to the Morphology of the Mammoth, Elephas Primigenius Blumenbach;
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[86] Fiedel, Stuart (2009). Sudden Deaths: The Chronology of Terminal Pleistocene Megafaunal Extinction.
In Haynes, Gary. American Megafaunal Extinctions at
the End of the Pleistocene. Springer. pp. 2137. [100] Farrand, W. R. (1961).
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8792-9.
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[87] Nogus-Bravo, D.; Rodrguez, J. S.; Hortal, J. N.;
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729735.
Batra, P.; Arajo, M. B. (2008). Barnosky, Andoi:10.1126/science.133.3455.729. PMID 17777646.
thony, ed. Climate Change, Humans, and the Extinction of the Woolly Mammoth. PLoS Biology 6 (4): [101] Lister, 2007. p. 59
e79. doi:10.1371/journal.pbio.0060079. PMC 2276529.
PMID 18384234.
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[103] Mol, D. et al. (2001). The Jarkov Mammoth: 20,000- [118] Zimov, S. A. (2005). ESSAYS ON SCIENCE AND
Year-Old carcass of a Siberian woolly mammoth MamSOCIETY: Pleistocene Park: Return of the Mammuthus primigenius (Blumenbach, 1799). The World of
moths Ecosystem. Science 308 (5723): 796798.
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doi:10.1126/science.1113442.
( 1620 October 2001, Rome): 305309. Full text pdf
[119] Loi, Pasqualino; Saragusty, Joseph; Ptak, Grazyna
(2014). Cloning the Mammoth: A Complicated Task
[104] Debruyne, R. G.; Barriel, V. R.; Tassy, P. (2003).
or Just a Dream?" 753. pp. 489502. doi:10.1007/978Mitochondrial cytochrome b of the Lyakhov mam1-4939-0820-2_19. PMID 25091921.
moth (Proboscidea, Mammalia): New data and phylogenetic analyses of Elephantidae. Molecular Phylogenetics and Evolution 26 (3): 421434. doi:10.1016/S1055- [120] Lister, 2007. p. 137
7903(02)00292-0. PMID 12644401.
[121] Lister, 2007. pp. 137139
[105] Kosintsev, P. A.; Lapteva, E. G.; Tromova, S. S.; Zan- [122] Of Mammoths and Men April 2013 National Geographic
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(magazine)
Environmental reconstruction inferred from the intestinal contents of the Yamal baby mammoth Lyuba (Mam- [123] Lister, 2007. p. 54
muthus primigenius Blumenbach, 1799)". Quaternary International 255: 231. Bibcode:2012QuInt.255..231K. [124] Sjgren, B. (1962). Farliga djur och djur som inte nns (in
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[106] Rincon, P. (10 July 2007). Baby mammoth discovery un- [125] Lister, 2007. p. 55
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[108] Smith, O. (21 April 2009). Baby mammoth Lyuba,
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[127] Lankford, G. E. (1980). Pleistocene Animals in Folk

Memory. The Journal of American Folklore 93 (369):
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[109] Fisher, Daniel C. (2014). X-ray computed tomography [128] Mayor, A. (2005). Fossil Legends of the First Americans.
Princeton: Princeton University Press. p. 97. ISBN 0of two mammoth calf mummies. Journal of Paleontol691-11345-9.
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[110] Mashchenko, E. N.; Protopopov, A. V.; Plotnikov, V. V.; [129] Murray, M. (1960). Henry Tukeman: Mammoths Roar
was Heard All The Way to the Smithsonian. Tacoma
Pavlov, I. S. (2013). Specic characters of the mamPublic Library. Retrieved 17 January 2008.
moth calf (Mammuthus primigenius) from the Khroma
River (Yakutia)". Biology Bulletin 40 (7): 626641.
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[111] Child nds 30,000 year old mammoth in North Russia.
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[114] Stone, R. (1999). Cloning the Woolly Mammoth. Discover Magazine.
[115] Cloned Mammoths Made More Likely by Frozen Mice.
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6.10 Bibliography

Data related to Mammuthus primigenius at Wikispecies

Media related to Mammuthus primigenius at Wikimedia Commons
Lister, A.; Bahn, P. (2007). Mammoths - Giants of
the Ice Age (3 ed.). London: Frances Lincoln. ISBN
978-0-520-26160-0.

Chapter 7

Woolly rhinoceros
The woolly rhinoceros (Coelodonta antiquitatis) is an
extinct species of rhinoceros that was common throughout Europe and northern Asia[1] during the Pleistocene
epoch and survived the last glacial period. The genus
name Coelodonta means cavity tooth. The woolly
rhinoceros was a member of the Pleistocene megafauna.

7.1 Evolution

Cast of the mummied Starunia specimen, Natural History Museum, London

an estimated weight of around 2,7213,175 kg (5,999

7,000 lb).[1] The woolly rhinoceros could grow to be 2
m (6.6 ft) tall;[1] the body size was thus comparable, or
slightly larger than, the extant White rhinoceros.[4] Two
horns on the skull were made of keratin, the anterior horn
being 61 cm (24 in) in length,[5] with a smaller horn between its eyes.[6] It had thick, long fur, small ears, short,
Chauvet cave art depicting a woolly rhino
thick legs, and a stocky body. Cave paintings suggest
a wide dark band between the front and hind legs, but
As the last and most derived member of the Pleistocene
the feature is not universal, and identication of pictured
rhinoceros lineage, the woolly rhinoceros was well
rhinoceroses as woolly rhinoceros is uncertain.
adapted to its environment. Stocky limbs and thick
woolly pelage made it well suited to the steppe-tundra Its shape was known only from prehistoric cave drawings
environment prevalent across the Palearctic ecozone dur- until a completely preserved specimen (missing only the
ing the Pleistocene glaciations. Like the vast majority fur and hooves) was discovered in a tar pit in Starunia,
of rhinoceroses, the body plan of the woolly rhinoceros Poland. The specimen, an adult female, is now on display
adhered to a conservative morphology, like the rst in the Polish Academy of Sciences' Museum of Natural
History in Krakw. Several frozen specimens have also
rhinoceroses seen in the late Eocene.
been found in Siberia, the latest in 2007.[7]
A study of 40-70.000 year old DNA samples showed its
closest extant relative is the Sumatran rhinoceros.[2]

7.3 Behavior and habitat

7.2 Description
The external appearance of woolly rhinos is known from
mummied individuals from Siberia as well as cave
paintings.[3] An adult woolly rhinoceros was typically
around 3 to 3.8 metres (10 to 12.5 feet) in length, with

The woolly rhinoceros used its horns for defensive purposes and to attract mates. During Greenland Stadial 2
(the Last Glacial Maximum[8] ) the North Sea retreated
northward, as sea levels were up to 125 metres (410 ft)
lower than today. The woolly rhinoceros roamed the ex-

68

7.4. EXTINCTION

69
constructed using several lines of evidence. Climatic reconstructions indicate the preferred environment to have
been cold and arid steppe-tundra, with large herbivores
forming an important part of the feedback cycle. Pollen
analysis shows a prevalence of grasses and sedges within
a more complicated vegetation mosaic.

Restoration by Charles R. Knight

posed Doggerland and much of Northern Europe and

was common in the cold, arid desert that is southern
England[9] and the North Sea today. Its geographical range expanded and contracted with the alternating
cold and warm cycles, forcing populations to migrate as
glaciers receded. The woolly rhinoceros co-existed with
woolly mammoths and several other extinct larger mammals of the Pleistocene megafauna. A close relative,
Elasmotherium, had a more southern range.

A strain vector biomechanical investigation of the skull,

mandible and teeth of a well-preserved last cold stage individual recovered from Whitemoor Haye, Staordshire,
revealed musculature and dental characteristics that support a grazing feeding preference. In particular, the enlargement of the temporalis and neck muscles is consistent with that required to resist the large tugging forces
generated when taking large mouthfuls of fodder from
the ground. The presence of a large diastema supports
this theory.
Comparisons with extant perissodactyls conrm that
Coelodonta was a hindgut fermentor with a single stomach, and as such would have grazed upon cellulose-rich,
protein-poor fodder. This method of digestion would
have required a large throughput of food and thus links
the large mouthful size to the low nutritive content of the
chosen grasses and sedges.[12]

Recent evidence suggests that woolly rhinos alive in the

Arctic during the Last Glacial Maximum consumed
In 2011, the oldest known woolly rhinoceros fossil was approximately equal volumes of forbs, such as Artemisia,
discovered from 3.6 million years in the Himalayas on the and graminoids.[13]
cold Tibetan Plateau, suggesting it existed there during a
period of general climate warmth around the earth. It is
believed that they migrated from there to northern Asia
and Europe when the Ice Age began.[10]
Females gave birth to one or two calves.[11]

7.3.1

Diet

7.4 Extinction

Main article: Pleistocene megafauna

Many species of Pleistocene megafauna, like the woolly

Woolly rhinoceros and other Ice Age mammals in late Pleistocene

northern Spain, by Mauricio Antn

rhinoceros, became extinct around the same time period.

Human and Neanderthal hunting is often cited as one
cause.[14] Other theories for the cause of the extinctions
are climate change associated with the receding Ice age
Frozen head, leg and horn from Siberia, 1849
and the hyperdisease hypothesis (q.v. Quaternary extinc[15]
Controversy has long surrounded the precise dietary pref- tion event).
erence of Coelodonta as past investigations have found Recent radiocarbon dating indicates that populations surboth grazing and browsing modes of life to be plausi- vived as recently as 8,000 BC in western Siberia. Howble. The palaeodiet of the woolly rhinoceros has been re- ever, the accuracy of this date is uncertain, as several ra-

70
diocarbon plateaus exist around this time. The extinction
does not coincide with the end of the last ice age but does
coincide with a minor yet severe climatic reversal that
lasted for about 1,0001,250 years, the Younger Dryas
(GS1 - Greenland Stadial 1), characterized by glacial
readvances and severe cooling globally, a brief interlude
in the continuing warming subsequent to the termination
of the last major ice age (GS2), thought to have been due
to a shutdown of the thermohaline circulation in the ocean
due to huge inuxes of cold fresh water from the preceding sustained glacial melting during the warmer Interstadial (GI1 - Greenland Interstadial 1 - ca. 16,00011,450
14
C years B.P.).
The Pinhole Cave Man is a late Paleolithic gure of a man
engraved on a rib bone of the Woolly rhinoceros, found
at Creswell Crags in England.[16]

7.5 See also

7.6 References
[1] Woolly Rhino (Coelodonta antiquitatis)". International
Rhino Foundation. Retrieved October 30, 2011.
[2] Orlando, L.; Leonard, J. A.; Thenot, A. L.; Laudet,
V.; Guerin, C.; Hnni, C. (2003). Ancient DNA
analysis reveals woolly rhino evolutionary relationships. Molecular Phylogenetics and Evolution 28 (3):
485499. doi:10.1016/S1055-7903(03)00023-X. PMID
12927133.

CHAPTER 7. WOOLLY RHINOCEROS

[9] Ian Rolfe, W. D. (1966). Woolly rhinoceros from the

Scottish Pleistocene. Scottish Journal of Geology 2 (3):
253. doi:10.1144/sjg02030253.
[10] Ice Age giants may have evolved in Tibet. CNN. 1
September 2011. Retrieved 2 September 2011.
[11] Walker, Matt (6 December 2012). Prehistoric rhino reveals secrets. BBC News.
[12] SAS Bulletin, Volume 26, number 3/4, Winter 2003 from
the Society for Archaeological Sciences
[13] Willerslev E, Davison J, Moora M, Zobel M, Coissac E,
Edwards ME, Lorenzen ED, Vestergrd M, Gussarova
G, Haile J, Craine J, Gielly L, Boessenkool S, Epp LS,
Pearman PB, Cheddadi R, Murray D, Brthen KA, Yoccoz N, Binney H, Cruaud C, Wincker P, Goslar T, Alsos IG, Bellemain E, Brysting AK, Elven R, Snsteb JH,
Murton J, Sher A, Rasmussen M, Rnn R, Mourier T,
Cooper A, Austin J, Mller P, Froese D, Zazula G, Pompanon F, Rioux D, Niderkorn V, Tikhonov A, Savvinov
G, Roberts RG, MacPhee RD, Gilbert MT, Kjr KH,
Orlando L, Brochmann C, Taberlet P. (2014). Fifty
thousand years of Arctic vegetation and megafaunal diet.
Nature 506 (7486): 4751. doi:10.1038/nature12921.
PMID 24499916.
[14] Diamond, Jared (1997). Guns, Germs and Steel. New
York: Vintage. ISBN 0-09-930278-0.
[15] Grayson, D. K.; Meltzer, D. J. (2003). A requiem
for North American overkill. Journal of Archaeological Science 30 (5): 585593. doi:10.1016/S03054403(02)00205-4.
[16] engraved bone/antler. British Museum.

[3] Frozen Fauna of the Mammoth Steppe.

[4] Krause, Hans (2011). HKHPE 07 02. hanskrause.de.
Retrieved July 12, 2012.
[5] Haines, Tim; Chambers, Paul (2005). Coelodonta. The
complete guide to prehistoric life (First ed.). Bualo, N.Y.:
Firey Books. p. 203. ISBN 978-1-55407-181-4.

Parker, Steve. Dinosaurus: The Complete Guide to

Dinosaurs. Firey Books Inc, 2003. Pg. 422.

7.7 External links

[6] Fortelius, Mikael (1983). The morphology and paleobiological signicance of the horns ofCoelodonta
antiquitatis(Mammalia:
Rhinocerotidae)".
Journal of Vertebrate Paleontology 3 (2):
125135.
doi:10.1080/02724634.1983.10011964.
ISSN 02724634.

More pictures of the fully preserved tar pit wholly

rhinoceros that was found in Poland (text in Polish)

[7] Boeskorov, G. G. (2012).

Some specic morphological and ecological features of the fossil
woolly rhinoceros (Coelodonta antiquitatis Blumenbach 1799)".
Biology Bulletin 39 (8): 692707.
doi:10.1134/S106235901208002X.

International Rhino Foundation: Woolly Rhino

[8] Jacobi, Roger M.; Rose, James; MacLeod, Alison;

Higham, Thomas F.G. (2009). Revised radiocarbon ages on woolly rhinoceros (Coelodonta antiquitatis) from western central Scotland: signicance for
timing the extinction of woolly rhinoceros in Britain
and the onset of the LGM in central Scotland.
Quaternary Science Reviews 28 (2526): 25512556.
doi:10.1016/j.quascirev.2009.08.010. ISSN 0277-3791.

Fossil skull of a woolly rhinoceros from Belgium

Fossil skull of a woolly rhinoceros from Germany

7.8. TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

71

7.8 Text and image sources, contributors, and licenses

7.8.1

Text

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Anome, Tarquin, Manning Bartlett, Jeronimo, Ap, Andre Engels, Rgamble, Rmhermen, Roadrunner, SimonP, Jaknouse, Paul Barlow,
Lexor, Menchi, Ixfd64, Minesweeper, Ahoerstemeier, Stan Shebs, Jimfbleak, William M. Connolley, Den fjttrade ankan, Glenn, Michael
Shields, Silthor, Nikai, Andres, Lee M, The Tom, Charles Matthews, Timwi, Steinsky, Hao2lian, Dragons ight, Tempshill, SEWilco,
Toreau, Joy, Wetman, Chrisjj, Pakaran, Pollinator, Chuunen Baka, Robbot, ChrisO, Jenmoa, Moondyne, Naddy, Chris Roy, Babbage,
Yosri, Sverdrup, Ojigiri, Meelar, Aetheling, Unfree, Terjepetersen, Hermes, Giftlite, DocWatson42, Jyril, SamE, Meursault2004, Zigger, Wwoods, Everyking, Bkonrad, Hoho, Maha ts, Curps, Alison, Leonard G., VampWillow, RayTomes, Wmahan, Mackeriv, Pgan002,
Andycjp, Gdr, SarekOfVulcan, Zeimusu, Antandrus, Beland, Jossi, Karl-Henner, Sam Hocevar, Huaiwei, Joyous!, Kareeser, Syvanen,
Trevor MacInnis, Lacrimosus, Atrian, R, Freakofnurture, Imroy, DanielCD, Discospinster, Rich Farmbrough, Guanabot, Wclark, Wrp103,
Vsmith, ArnoldReinhold, JimR, Xezbeth, GregBenson, Bender235, Violetriga, RJHall, Shanes, RoyBoy, Bobo192, Nigelj, NetBot, Jolomo,
BillCook, Martg76, Obradovic Goran, Hintha, Haham hanuka, Jonathunder, Hooperbloob, Nsaa, Jason One, Alansohn, Gary, Anthony Appleyard, Shadikka, Arthena, Jeltz, Lightdarkness, Bart133, Snowolf, Wtmitchell, Velella, EvenT, Yuckfoo, Tony Sidaway, TenOfAllTrades,
Mikeo, Redvers, Phi beta, Adrian.benko, WilliamKF, Zanaq, Boothy443, MagicBez, Tyz, Merlinme, Before My Ken, Ruud Koot, MGTom, Tckma, MONGO, Kelisi, Dzordzm, GregorB, Jugger90, Rotten, Prothonotar, Kmontgom, Marudubshinki, Dysepsion, Lusitana, Graham87, Magister Mathematicae, Cuchullain, Kbdank71, Demonuk, Mana Excalibur, Rjwilmsi, CyberGhostface, Ittiz, Ikh, Seraphimblade,
ErikHaugen, Nneonneo, Dolphonia, Mohawkjohn, Williamborg, Nguyen Thanh Quang, Yamamoto Ichiro, Leo44, FlaBot, Ian Pitchford,
SchuminWeb, Nihiltres, Nivix, RexNL, Gurch, Ayla, Neofelis Nebulosa, IlGreven, Bmicomp, Tedder, Zotel, King of Hearts, Bornhj,
DVdm, Gwernol, Wavelength, TexasAndroid, RobotE, Sceptre, Cyferx, RussBot, DMahalko, Sarranduin, Anonymous editor, Splash, SpuriousQ, RadioFan2 (usurped), Stephenb, Wimt, Big Brother 1984, NawlinWiki, ENeville, Anchjo, Wiki alf, Chick Bowen, NickBush24,
DavidH, Robchurch, Retired username, Hogne, Inselpeter, Nick C, Semperf, Syrthiss, DGJM, Mditto, Lockesdonkey, DeadEyeArrow,
Jpeob, Leptictidium, WAS 4.250, FF2010, Lt-wiki-bot, Theda, Sarefo, JoanneB, Slehar, Hayden120, Whobot, Staxringold, AlexD, Whouk,
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Pilotguy, Charivari, Ace ETP, Autopilot, SashatoBot, Swatjester, Dr. Sunglasses, Kuru, John, J 1982, Normalityrelief, Locutus Borg,
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Elliott Small, Courcelles, Brianjohn, RobinDaniel, Tawkerbot2, VinceB, Kaze0010, Friendly Neighbour, CmdrObot, Sir Vicious, Unionhawk, Bons, Rwammang, Fitzaubrey, Shizane, Yopienso, Funnyfarmofdoom, Sopoforic, Cydebot, Michaelas10, Gogo Dodo, Was a bee,
Kallerdis, Tkynerd, Odie5533, DumbBOT, Chrislk02, Dferrantino, Epanalepsis, Romon, JamesAM, Malleus Fatuorum, Epbr123, Nowimnthing, KimDabelsteinPetersen, Daniel, Andyjsmith, Callmarcus, A3RO, James086, JustAGal, Dfrg.msc, Dgies, Sean William, Escarbot,
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DennyColt, Sanfranman59, 2marcus, Rjm at sleepers, Littlebrandon, Webster girl, Robert1947, Shouriki, C-M, Ftjrwrites, Wenli, Billgordon1099, Meters, Srcfoo, Graymornings, Falcon8765, Insanity Incarnate, Rau J, NHRHS2010, Djmatch3000, SieBot, Cmiller.design,
Tresiden, Euryalus, Matthewhunter1234, Dawn Bard, Astrovega, Yintan, The way, the truth, and the light, Twirling, Calabraxthis, Chris
Light, Keilana, Oda Mari, Pierre-tienne Messier, Wilson44691, Aly89, Cameron3, Oxymoron83, Antonio Lopez, Lightmouse, Blackeydp,
Limbo466, Alex.muller, Lynntoniolondon, Venomousdolphin6, Hatmatbbat10, Spartan-James, Sugarcubez, Anchor Link Bot, Tesi1700,
Abmcdonald, WikiLaurent, Lololo85, Denisarona, Escape Orbit, Velvetron, Karent82, Webridge, Zerueu, ClueBot, The Thing That Should
Not Be, 0nullbinary0, Swedish fusilier, Stinkyface5, Ndenison, Wysprgr2005, Arakunem, Minging19994, Candystorebut, Werockhard,
Blanchardb, Awickert, Ktr101, Excirial, Bob 1232345324644, Canis Lupus, Jusdafax, Simon D M, Tyler, JamieS93, Norandav, Jo Weber,
Tnxman307, Razorame, Ppwu, Noosentaal, Dekisugi, Audaciter, Ottawa4ever, Thehelpfulone, Thingg, Aitias, Peekaboo1996, Versus22,
SoxBot III, Egmontaz, Wkboonec, Theking2, Calidrisalba, Dtpeck, Ost316, Feyrauth, Clive McRoderick, Quinboo42, Gimmethoseshoes,
Noctibus, WikiDao, TravisAF, SkyLined, RyanCross, HexaChord, Iranway, Jtknowles, Kajabla, Addbot, Roentgenium111, Some jerk on
the Internet, DOI bot, Felipetavira, Ronhjones, CanadianLinuxUser, CactusWriter, Cst17, Skyezx, Phatcat919, Download, Morning277,
Glane23, LinkFA-Bot, Tyw7, Charltonsmells, Craigsjones, Tide rolls, Zorrobot, Bartledan, Frehley, Legobot, Luckas-bot, ZX81, Yobot,
2D, R3D17, Cooljason1, Sanyi4, THEN WHO WAS PHONE?, Ty94ler, Ice Age Rlue, Eric-Wester, Magog the Ogre, Nakor, AnomieBOT,
SydneyLk, Message From Xenu, Jim1138, Short Brigade Harvester Boris, Piano non troppo, Collieuk, Gaugan, Kingpin13, Stickman30,
Pequod76, Userresuuser, Crecy99, Flewis, Materialscientist, Hunnjazal, Zeog, Elmmapleoakpine, The High Fin Sperm Whale, Citation
bot, OllieFury, Maxis ftw, Frankenpuppy, Neurolysis, Xqbot, Intelati, The sock that should not be, Capricorn42, Millahnna, ChildofMidnight, Tinndel, NFD9001, Anna Frodesiak, Br77rino, Kernel Corn, Didi29, RibotBOT, Franco3450, TonyHagale, John Nissen, Renwick99,
IcedNut, Hutenova, SD5, Carlog3, Joey-fca, FrescoBot, Jared852, Remotelysensed, Keelec, Pepper, Tranletuhan, Tan3157, JMS Old Al,
Vishnu2011, Chutznik, Mycocobean, Friedrich von Weech, Jamesooders, Briardew, Citation bot 1, Citation bot 4, Codyag123, Torrr torrr,
Pinethicket, I dream of horses, Arctic Night, Hyattdj, Tom.Reding, Avidmosh, Tinton5, le ottante, VinnyXY, Ice Age lover, RandomStringOfCharacters, Fumitol, Angloguy, Rannaro, Full-date unlinking bot, Camlowie, Guru42, White Shadows, Utility Monster, FoxBot,
TrickyM, SDLarsen, Vrenator, Miracle Pen, Inferior Olive, Brightbritt, Nascar1996, Stroppolo, Estien, Reach Out to the Truth, Minimac,
DARTH SIDIOUS 2, RjwilmsiBot, Bento00, Cfealtman, Hh.ezra, Chemyanda, World Lever, Giorgiogp2, Domesticenginerd, Kpuer-

72

CHAPTER 7. WOOLLY RHINOCEROS

sh, Pditlev, Jakaloke, ScottyBerg, RA0808, Poopysnoop, Heljqfy, Beatleben5, Slightsmile, TuHan-Bot, Saiguy96, Emmick4, MithrandirAgain, Alcart, Appaloosacakes, Xuiolhcarlos, 1234r00t, Lockon23, ZephyrWindOlympus, Frostyrollie, Wayne Slam, Getsagotta, JaySebastos, L Kensington, Mayur, Grammar Prof, RockMagnetist, 23Simon, Jkvc2003, DASHBotAV, Will Beback Auto, ClueBot NG,
Rich Smith, Tanmoy Panigrahi, Movses-bot, Jbrosseau1, ScottSteiner, Dream of Nyx, Ineedinfo54, MD K47, Wre111, Helpful Pixie Bot,
12kar, Calabe1992, Gob Lofa, Bibcode Bot, Lowercase sigmabot, NewsAndEventsGuy, ServiceAT, Northamerica1000, Tigona, Nyreal,
Dodshe, Sammrud, Op47, Cadiomals, Mdy66, Aranea Mortem, 16ddp, Hamish59, SBrunt7, TBrandley, Meegan98, Kellieebee, Martialartistx, ChrisGualtieri, Tulino, Joel hill, Jengibbo, Mzbob3068, Bellatrix121, Dexbot, TomoK12, Knuand, Vanshay96, Webclient101,
G.Kiruthikan, Lugia2453, Hihi1324, Everymorning, Crwaterhouse, The Wikimon, Joserial, Prokaryotes, Monkbot and Anonymous: 1200
Megafauna Source: http://en.wikipedia.org/wiki/Megafauna?oldid=636341728 Contributors: Bryan Derksen, Eclecticology, Rmhermen,
Karen Johnson, Roadrunner, Tez, Tannin, Ellywa, Habj, Charles Matthews, Timc, Populus, Wetman, Jerzy, 80.255, Dale Arnett, Moondyne, Altenmann, Sparky, Sam Spade, Clarkk, UtherSRG, Alan Liefting, Ancheta Wis, MPF, Holizz, Wiglaf, Yak, Tom Radulovich,
Golbez, Antandrus, The Singing Badger, Lesgles, Creidieki, Sparky the Seventh Chaos, DanielCD, Narsil, Bishonen, El C, Bobo192,
Polocrunch, Foobaz, MARQUIS111, Larry V, Malo, Titanium Dragon, Dinoguy2, BDD, LordAmeth, Gene Nygaard, Stemonitis, Mazca,
Pixeltoo, Miwasatoshi, Rjwilmsi, ErikHaugen, FlaBot, Zaurus, MacRusgail, TeaDrinker, Tedder, Mordicai, UkPaolo, Wavelength, Sceptre,
Hairy Dude, Jimp, Xaa, SatuSuro, Ohwilleke, Pigman, Chaser, Gaius Cornelius, CambridgeBayWeather, InterwikiLinksRule, TEB728,
Dysmorodrepanis, ExRat, Apokryltaros, Mosquitopsu, Epipelagic, Alasdair, Mhenriday, Codman, KnightRider, RupertMillard, SmackBot, Francisco Valverde, Drn8, Chris the speller, Deanmo19, J. Spencer, Jwillbur, Adrian ne, PiMaster3, GumTree, Richard001, The
PIPE, Bcasterline, Soap, Tktktk, Mgiganteus1, A. Parrot, JHunterJ, Neddyseagoon, Arctic-Editor, Kurtle, Peter Horn, Scorpios, Iridescent, Kaarel, J Milburn, CRGreathouse, Rwammang, Markhu, CuriousEric, AshLin, Joechao, Tug201, Dougweller, DumbBOT, LarryQ,
Epbr123, Marek69, Smile a While, PeterDz, Nutsuo, Escarbot, BlackOcelot, Spartaz, Canadian-Bacon, Ingolfson, Vultur, Acroterion,
WolfmanSF, Jrowle, Carom, Lucasake, JamesBWatson, PenguinJockey, Adrian J. Hunter, Inclusivedisjunction, Arnesh, NatureA16, Atarr,
Mermaid from the Baltic Sea, Jim.henderson, CommonsDelinker, AlexiusHoratius, Power level (Dragon Ball), Andy Johnston, SJP, Frickeg,
Richard New Forest, Sam Blacketer, Murderbike, Nihilrat, Je G., Philip Trueman, TXiKiBoT, MrChupon, Ethel Aardvark, MeegsC, The
way, the truth, and the light, Scarlet23, Mimihitam, Goustien, RingManX, WikiLaurent, Pavel.Riha.CB, ImageRemovalBot, Sfan00 IMG,
ClueBot, The Thing That Should Not Be, Niceguyedc, Estirabot, Amaltheus, Darkicebot, Fastily, T.carnifex, Little Mountain 5, Sleptrip,
Addbot, Guoguo12, Ronhjones, Ehrenkater, Lightbot, Laikayiu, Legobot, Luckas-bot, 2D, Azcolvin429, Againme, Kingpin13, Materialscientist, Hunnjazal, Wrelwser43, Corbon, LilHelpa, Poetaris, Meshin0, BindingArbitration, GrouchoBot, Lhuntr, FrescoBot, Bighead01753,
Tobby72, Finalius, Zencowboy27, Citation bot 2, Max moa, Citation bot 1, Jaybird vt, Googlemeister, Gaia5074Q, Jonkerz, Vrenator,
JAKEMAN5, Innotata, Tbhotch, Obsidian Soul, RjwilmsiBot, Kristian.gareau, WildBot, Mukogodo, John of Reading, Ebrambot, Peter
M. Brown, 1nv151b13.b0b, Ego White Tray, Orin04, ClueBot NG, Megaherbivore, Jonathan Fernando, Dream of Nyx, Oslep11, Helpful Pixie Bot, Plutosdogscat, Gob Lofa, Bibcode Bot, Lowercase sigmabot, MusikAnimal, Candymans, Cadiomals, CitationCleanerBot,
Tangerinehistry, Mike.BRZ, Moldovan0731, Klilidiplomus, BattyBot, Nick.mon, Khazar2, AldezD, Prettybirdie, Raptormimus456, Praemonitus, Fafnir1, Monkbot, Filedelinkerbot and Anonymous: 366
Pleistocene Source: http://en.wikipedia.org/wiki/Pleistocene?oldid=636705203 Contributors: Bryan Derksen, Zundark, Berek, Andre Engels, Vignaux, Rmhermen, Edward, Michael Hardy, Glenn, Susurrus, Evercat, Smack, Emperorbma, David Newton, Reddi, Tarosan,
The Anomebot, Dragons ight, SEWilco, Joy, Nightsky, Wetman, Frazzydee, Robbot, ChrisO, Moondyne, Altenmann, Dittaeva, Nurg,
Kamakura, Jsonitsac, Saforrest, Casito, Drew3D, GreatWhiteNortherner, Nagelfar, DocWatson42, Jao, Jyril, Hagedis, MichaelHaeckel,
Gilgamesh, Pascal666, CryptoDerk, Garth 187, Beland, Tomruen, Ta bu shi da yu, DanielCD, Vsmith, Florian Blaschke, Dbachmann,
Bender235, Swid, Eric Forste, CanisRufus, El C, Kwamikagami, Bobo192, Viriditas, Brim, JeR, Jag123, QTxVi4bEMRbrNqOorWBV,
Orangemarlin, Siim, Alansohn, Andrewpmk, Gpvos, Gene Nygaard, Bastin, Stemonitis, Kelly Martin, Jerey O. Gustafson, Woohookitty,
Carcharoth, DanHobley, Marudubshinki, Graham87, BD2412, Cmsg, Euchrid, Rjwilmsi, Swiftwindcat, SMC, Ucucha, FlaBot, Margosbot,
Maire, Zotel, Lemuel Gulliver, CJLL Wright, Chobot, Garas, Krawunsel, YurikBot, Hairy Dude, Jimp, Stephenb, Bovineone, Anomalocaris, Dysmorodrepanis, Patrick MMA Bringmans, SFC9394, Bota47, Botteville, Leptictidium, Poppy, Deville, Barryob, Pdraic MacUidhir, Sardanaphalus, Attilios, SmackBot, Enlil Ninlil, KnowledgeOfSelf, IstvanWolf, Kinhull, DHN-bot, Abyssal, AMK152, DGerman,
Khoikhoi, Noles1984, Iblardi, Paul H., Vina-iwbot, Bejnar, SashatoBot, Lambiam, Thomas keyes, Kevmin, KarlM, Mgiganteus1, Ben
Moore, Werdan7, Dicklyon, Geologyguy, Ginkgo100, P.Geol, Civil Engineer III, Brianjohn, Eluchil404, Markjoseph125, Woudloper,
AshLin, Dougweller, Kendirangu, Mattisse, HJJHolm, Epbr123, Parsa, Callmarcus, A3RO, Chrisdab, PJtP, Greg L, OrenBochman, AntiVandalBot, Luna Santin, Seaphoto, Mikenorton, Deective, Pedro, VoABot II, Fang 23, E104421, Warren Dew, Gwern, Ugajin, R'n'B,
CommonsDelinker, J.delanoy, UBeR, KenSharp, ElinWhitneySmith, Crocadog, Johnbod, Austin512, Janet1983, Rosenknospe, Soczyczi, Root7, STBotD, Tygrrr, Squids and Chips, VolkovBot, Thisisborin9, BlazeTheMovieFan, A4bot, Zybez, Cbrettin, Raven rs, Una
Smith, PDFbot, CaptainFossil, Autodidactyl, Shouriki, MCTales, Thanatos666, Ceranthor, Tom Meijer, Fanatix, SieBot, ToePeu.bot,
Cwkmail, Andrewjlockley, Chris Light, FunkMonk, Joe Gatt, DiBgd, Wilson44691, General Synopsis, PhilMacD, Lightmouse, Macy, Iceman63976, CP2002, Precious Roy, Randy Kryn, Gold1618, Anthony R. Hansen, ClueBot, Hongthay, Cygnis insignis, Arakunem, Drmies,
TheOldJacobite, CounterVandalismBot, ChandlerMapBot, Arunsingh16, Auntof6, Rockfang, Awickert, AssegaiAli, Jusdafax, Crywalt,
Isthisthingworking, Goodvac, InternetMeme, Mhese, NellieBly, Torahjerus14, Addbot, Tigerbreath13, Capouch, TutterMouse, Fluernutter, Ka Faraq Gatri, LaaknorBot, 37ophiuchi, AndersBot, Spike the Dingo, Tyw7, Tide rolls, Lightbot, , Solrac1993, Legobot,
Luckas-bot, Yobot, Ptbotgourou, MTWEmperor, Shore3, KamikazeBot, Eric-Wester, AnomieBOT, Lebanonman19, Richardlord50, Hunnjazal, The High Fin Sperm Whale, Digitaldomain, Xqbot, Poetaris, GrouchoBot, Doulos Christos, Tashka99, Moxy, Pinethicket, RedBot,
Lars Washington, December21st2012Freak, Fama Clamosa, Lotje, Vrenator, Aoidh, Stephen MUFC, Obsidian Soul, TjBot, Nossing,
Waso99, EmausBot, Chermundy, Slightsmile, AvicBot, WeijiBaikeBianji, Cobaltcigs, Morten Knudsen, TyA, NTox, TYelliot, ClueBot
NG, Joefromrandb, Vacation9, Stuartsmally, Widr, Helpful Pixie Bot, Gob Lofa, Yendor of yinn, MangoWong, MusikAnimal, Davidiad,
Cadiomals, Ornithodiez, Fjasl;d, Pseudofusulina, BattyBot, Isumbard Prince, Markomazzoni, US Jingoist, Ntra00, Hoppeduppeanut, DavidLeighEllis, Wailordwew, Prokaryotes, Param Mudgal, Animalarmageddon, Monkbot, Dhm4444, Piesquared93, TropicalCyclones243,
Marcel Hendrik and Anonymous: 265
Prehistoric mammal Source: http://en.wikipedia.org/wiki/Prehistoric%20mammal?oldid=634637497 Contributors: CesarB, UtherSRG,
Jyril, DarkFantasy, DanielCD, Dancxjo, Bobo192, Fornadan, Pol098, Phlebas, Astropithicus, Ucucha, Dracontes, Roboto de Ajvol, Wavelength, Rtkat3, CambridgeBayWeather, Welsh, EncycloPetey, Greatgavini, J. Spencer, Jerkov, Wikid77, Dragon Helm, James truong,
DuncanHill, Liverpoolpaddy, Yurei-eggtart, NatureA16, CommonsDelinker, Alataristarion, CameronPG, Rei-bot, Kcatena, Wilson44691,
ClueBot, Eriksiers, Kevjenzak, PolarYukon, Addbot, RANDREWF7777, Lightbot, Luckas-bot, Yobot, Jpdinoman3, Piano non troppo,
Poetaris, Khanhvukk, Tjmoel, Dy2007, Alph Bot, EmausBot, Peter M. Brown, Kinghistory15 and Anonymous: 56
Stone Age Source: http://en.wikipedia.org/wiki/Stone%20Age?oldid=635450035 Contributors: Vicki Rosenzweig, Bryan Derksen, Tarquin, Taw, Alex.tan, Rmhermen, SimonP, Peterlin, BryceHarrington, Olivier, Frecklefoot, Patrick, Michael Hardy, Paul Barlow, Gdarin,

7.8. TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

73

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Bellardoo, BG19bot, Smallerjim, Goddamcaptchacode, MusikAnimal, , Sowsnek, FiveColourMap, CimanyD,
Cowsgobob, Jinglearceus, Lmarcell14, Chip123456, A Timelord, Teammm, Zhaofeng Li, ChrisGualtieri, Codeh, TacticalTurtleneck, EuroCarGT, Titchybear, Verryniceguy2, Dexbot, T v shah, Lugia2453, Ak5791, Frosty, Schiltron, Darth Sitges, PizzaHutCreeper, Ruby
Murray, Tentinator, Emily2117, Ugog Nizdast, Laurenmathews123, Ginsuloft, Pratheshsum, Xspike15, Lizia7, Kimboslicee, JaconaFrere,
Dinorexcoolio, Monkbot, Supermariolink777, Braden12345, Iluhrs, Owen minns, Flingbong, HMSLavender, 115ash, PrinceSulaiman,
Oinksgiant2000, Ninafundisha, Speedytacos, Surajbryan, Jessica Simpson03,
, Guillame Heavensburg and Anonymous: 1000
Woolly mammoth Source: http://en.wikipedia.org/wiki/Woolly%20mammoth?oldid=636651581 Contributors: William Avery, Ubiquity, Jimfbleak, Raven in Orbit, Adam Bishop, Timwi, Tpbradbury, AnonMoos, Wetman, Twang, Stephan Schulz, Auric, UtherSRG,
Michael Devore, Ezhiki, Mboverload, JoJan, DragonySixtyseven, TJSwoboda, Discospinster, Rich Farmbrough, Rama, Vsmith, Wefa,
Mikkel, Xezbeth, WegianWarrior, Bender235, Malkin, Bobo192, DCEdwards1966, Hesperian, Supersexyspacemonkey, Anthony Appleyard, Buaidh, Arthena, Monado, Axl, Aranae, Super-Magician, Dinoguy2, Staeiou, Itschris, Kazvorpal, Dismas, Duke33, Zntrip,

74

CHAPTER 7. WOOLLY RHINOCEROS

Feezo, Woohookitty, WadeSimMiser, Astanhope, GregorB, Miwasatoshi, Karam.Anthony.K, Graham87, Cuvtixo, BD2412, Rjwilmsi,
Jake Wartenberg, Jehochman, DoubleBlue, Ucucha, Rui Silva, SchuminWeb, Eubot, Siv0r, Old Moonraker, Jameshsher, Michaelritchie200, DVdm, Royal Scottish, Sceptre, Hairy Dude, Rtkat3, Peter G Werner, RussBot, CambridgeBayWeather, Pseudomonas, NawlinWiki, Dysmorodrepanis, Wiki alf, Grafen, Voyevoda, Lowe4091, Joel7687, Howcheng, Apokryltaros, RFBailey, Mithunc, Musiclover,
Samir, Elkman, Nlu, Calaschysm, 21655, CapitalLetterBeginning, JRawle, Aurax, TLSuda, RG2, Elliskev, SmackBot, Esradekan, Enlil
Ninlil, KnowledgeOfSelf, Hydrogen Iodide, Pmaas, Jrockley, Unnite, Drkarthi, Edgar181, Ga, Ian Rose, Gilliam, Hmains, Chris the
speller, Deanmo19, Isaacsurh, Scwlong, Aremith, Neo139, Rrburke, Jerkov, KnowledgeLord, Krich, Ken keisel, Kevlar67, Blake-, Iblardi,
Dreadstar, TGC55, Sigma 7, Risssa, Rockpocket, TyrannoRanger, John, Dwpaul, Lazylaces, Larry E. Matthews, Jimd, MarkSutton,
Mathewignash, Noah Salzman, Mr Stephen, Jimmy Pitt, Xiaphias, Waggers, Magere Hein, Dr.K., Danilot, Peyre, MrDolomite, Hu12,
Oceanofantics, RhoOphuichi, KsprayDad, Piccor, Pithecanthropus, Chetvorno, LessHeard vanU, Thetrick, J Milburn, Ale jrb, Dycedarg,
Erik Kennedy, Dgw, CuriousEric, Ejph, Dogman15, W.F.Galway, T Houdijk, Ryan, Reywas92, Gogo Dodo, Wordbuilder, Pascal.Tesson,
Christian75, Aanhorn, DumbBOT, Satori Son, Altaileopard, JamesAM, JAF1970, Epbr123, Dark fennec, HappyInGeneral, John254, Nimakha, CielProfond, Escarbot, AntiVandalBot, The Obento Musubi, Majorly, Nathanwright, NeilEvans, Mctoomer, Gh5046, WWB, Falconleaf, Gkhan, DOSGuy, Leuko, MER-C, The Transhumanist, OhanaUnited, Gavia immer, Acroterion, Io Katai, WolfmanSF, VoABot
II, Smorg, Aka042, Kennercat, Redbo, Catgut, ClovisPt, Loonymonkey, Enquire, DerHexer, Saberclaw, Connor Behan, NatureA16, FisherQueen, MartinBot, NAHID, Anaxial, CommonsDelinker, AlexiusHoratius, Johnpacklambert, Calvinung2000, J.delanoy, Trusilver, Bogey97, StonedChipmunk, BrokenSphere, Hakufu Sonsaku, Mattximus, Chiswick Chap, Colchicum, Serge925, SmilesALot, Juliancolton,
WLRoss, Idioma-bot, Feedme4, UnicornTapestry, Je G., Dom Kaos, CameronPG, Philip Trueman, Luftschlosz, Antinice, TXiKiBoT,
Maximillion Pegasus, Malinaccier, Tameeria, Oxfordwang, FuriousGeorge160, IllaZilla, LeaveSleaves, Raymondwinn, Improve, Roland
Kaufmann, Enigmaman, Enviroboy, MrChupon, Newbyguesses, Tiddly Tom, WereSpielChequers, Jauerback, Dawn Bard, Trigaranus,
Yintan, Vanished User 8a9b4725f8376, Grundle2600, FunkMonk, Aillema, Flyer22, Oda Mari, Bsaswin, Wilson44691, Topher385, Burninator22, Aelius28, Baseball Bugs, Goustien, Mygerardromance, Beastinwith, Denisarona, Escape Orbit, Drgbvw25, Smashville, Atif.t2,
AerosmithNirvana, 31stCenturyMatt, ClueBot, The Thing That Should Not Be, Voxpuppet, Dkenyon, Hazabaza2, CounterVandalismBot,
Blanchardb, Stylteralmaldo, Rotational, Puchiko, Awickert, Excirial, Pumpmeup, Sepeople, Greiss122, Audaciter, Nutindaleftear, Torver,
Thingg, Acabashi, Tomp45673, Ivorymammoth, Horselover Frost, Wuzupdude?, Iml, Versus22, Amaltheus, NERIC-Security, Brambleberry of RiverClan, Tdslk, Fastily, Roxy the dog, Mayatheg, Ost316, Skarebo, Noctibus, Poly12, Addbot, Jojhutton, Theultimatum, Tcncv,
Ronhjones, Osfania, Adrian 1001, 064ldingla, Bastion Monk, CanadianLinuxUser, AdRem, Cst17, Download, Morning277, Crankelwitz,
Debresser, Jasper Deng, Tyw7, Peridon, Tide rolls, Lightbot, Teles, First Light, Rojypala, Laikayiu, Bartledan, Luckas-bot, ZX81, Yobot,
EdwardLane, 2D, Kilom691, KamikazeBot, Synchronism, AnomieBOT, Floquenbeam, Kingpin13, Bluerasberry, Citation bot, Maxis ftw,
Frankenpuppy, Natashavcxz, JimVC3, Captain Lamphor, Capricorn42, Yoyo20, 019045dc, Gigemag76, DSisyphBot, Anna Frodesiak,
Dancurtisthompson, Blipton, Jakouso, Ispawnxi, LevenBoy, RibotBOT, N419BH, Joaquin008, Dan6hell66, Dalekdoom, Biomanzilla, Elcinoca, Compoundinterestisboring, Hvblofdj, Gourami Watcher, Citation bot 2, Animules, Agiseb, Citation bot 1, Redrose64, Pinethicket,
Trijnstel, LittleWink, A412, Loyalist Cannons, Davidchow7, Poloratgoet, Uutela, Mjs1991, HGArvedui, Fama Clamosa, Pithecanthropus4152, Jonkerz, Sumone10154, Bryanb246, 564dude, Jrc123, Brian the Editor, Emmyanneisawesome, Innotata, McKayJohn, Alon 68,
DARTH SIDIOUS 2, Mean as custard, RjwilmsiBot, Ladyy dee, Ineverheardofhim, Howiemorenzll, Oldcrookedjaw, DASHBot, EmausBot, John of Reading, Acather96, GA bot, Immunize, Nuujinn, ScottyBerg, Faolin42, Yt95, RA0808, Angman12, TuHan-Bot, Wikipelli,
KCC10-11, H3llBot, Danblee, David J Johnson, Wingman4l7, Tolly4bolly, Kirothereaper, Donner60, Bulwersator, Orange Suede Sofa,
GermanJoe, Peter Karlsen, Wakebrdkid, ClueBot NG, AlbertBickford, KlappCK, App210, MelbourneStar, LittleJerry, Bulldog73, Hectonichus, O.Koslowski, Widr, Oslep11, Helpful Pixie Bot, Mr. Editor XVIII, Bibcode Bot, Lowercase sigmabot, Rextron, SimonBob13, Eric567, AndrePooh, Stevesg1, Dan653, Mark Arsten, Silvrous, Irishtogher, Glevum, DPL bot, Hergilfs, Hamish59, Mike.BRZ,
Moldovan0731, Glacialfox, Doc2015, Anbu121, Darylgolden, Walker000, The Illusive Man, Mediran, CrunchySkies, Johnnyboy37, AutomaticStrikeout, Hmainsbot1, Webclient101, FiverFan65, Jfgoofy, TheOwlGuy, Leptus Froggi, Frosty, Smohammed2, Vole22, WikiTyson,
Jessie can wait, Zziccardi, Kevin12xd, Doctor99268, Soccerdude0123, THEBOME2000, Epicgenius, Dead Spikes97, PMS123, VoxelBot,
IJReid, MANINPHONEBOOTH, Codonaug, SamX, Apotea, Ugog Nizdast, Glaisher, TFA Protector Bot, G Robinson263, Bokyqwer,
KhanTheDestroyer, Akdeep, Fafnir1, Monkbot, Papyrus-winged ninja Akil, Screaming Merlin, Paleolithic Man, MCDinosaurhunter, EditorMakingEdits, Yukagir, Bammie73, Mammothlover1470, Huhiu, Mdjajich7624, FlappyJenson24, Hhindman21, Rushandbentley and
Anonymous: 672
Woolly rhinoceros Source: http://en.wikipedia.org/wiki/Woolly%20rhinoceros?oldid=635394689 Contributors: Azhyd, Frecklefoot,
Earth, Tannin, Pcb21, Muriel Gottrop, GCarty, Raven in Orbit, Wetman, Eugene van der Pijll, Phil Boswell, Baldhur, Seglea, Postdlf,
UtherSRG, DocWatson42, Wiglaf, Everyking, Mike R, Gdr, The Singing Badger, Kaldari, Bumm13, DanielCD, Xezbeth, Mwng, Bender235, Kbh3rd, Leperesh, Summer Song, Adambro, Jonathan Drain, Circeus, Supersexyspacemonkey, A2Kar, Anthony Appleyard,
Demi, Mac Davis, Aranae, Dinoguy2, Netkinetic, Sin-man, Sparkit, BD2412, Pmj, The wub, Ucucha, Leithp, Vuong Ngan Ha, Eubot,
TeaDrinker, Gdrbot, YurikBot, Anonymous editor, Bsharitt, CambridgeBayWeather, Voyevoda, Neum, BirgitteSB, Apokryltaros, Moe
Epsilon, Leptictidium, Groyolo, Hiddekel, KnightRider, SmackBot, Enlil Ninlil, Pmaas, Robin Whittleton, WookieInHeat, EncycloPetey,
Punchup, J. Spencer, DHN-bot, Wikirouta, Trekphiler, Chlewbot, OrphanBot, Kevlar67, Kevmin, Mgiganteus1, Stwalkerster, Yannzgob,
Atakdoug, JayHenry, Glanthor Reviol, Pseudo-Richard, Cydebot, Fifo, DumbBOT, Thijs!bot, Escarbot, IrishPete, Mikenorton, MER-C,
Sophie means wisdom, Severo, WolfmanSF, Catgut, Anaxial, CommonsDelinker, Cuddly Panda, Johnbod, HiLo48, NewEnglandYankee,
Shiraun, DorganBot, CameronPG, Tubbienine, Sandhillcrane, ^demonBot2, UnitedStatesian, Ninjatacoshell, Insanity Incarnate, Maharishi yogi, SieBot, BotMultichill, Orna82, FunkMonk, Helioseus, Graminophile, ClueBot, The Famous White Wolf, Hazabaza3, Trivialist,
JereyW75, Bingodile, Abrech, Mammoth lover, Wkharrisjr, XLinkBot, Addbot, Ronhjones, MrOllie, Glane23, Lightbot, First Light, Zorrobot,
, Luckas-bot, Yobot, Hohenloh, AnomieBOT, Jim1138, Graywords, Hunnjazal, ImperatorExercitus, Citation bot, ArthurBot,
Xqbot, J04n, GrouchoBot, Brambleshire, Philip72, Jca123456, LucienBOT, Archaeodontosaurus, Pinethicket, Zvn, Gatormax15, EmausBot, WikitanvirBot, Tolly4bolly, ChuispastonBot, ClueBot NG, Jnorton7558, Satellizer, John P. Harrison, Antiqueight, Helpful Pixie
Bot, Picoz, Amp71, Chris the Paleontologist, Hergilei, Dexbot, FoCuSandLeArN, Hmainsbot1, Capitaneteja, Utahraptor7887, NHCLS,
Mike246, Itc editor2, Papyrus-winged ninja Akil and Anonymous: 133

7.8.2

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License: Public domain Contributors: Own work Original artist: Myself
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75

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Connolley

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CHAPTER 7. WOOLLY RHINOCEROS

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7.8.3

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