Ice Agemegafauna and time notes

Contents

1 Ice age 11.1 Origin of ice age theory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.2 Evidence for ice ages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21.3 Major ice ages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31.4 Glacials and interglacials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41.5 Positive and negative feedback in glacial periods . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

1.5.1 Positive feedback processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51.5.2 Negative feedback processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

1.6 Causes of ice ages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51.6.1 Changes in Earth’s atmosphere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61.6.2 Position of the continents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61.6.3 Fluctuations in ocean currents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71.6.4 Uplift of the Tibetan plateau and surrounding mountain areas above the snowline . . . . . . 71.6.5 Variations in Earth’s orbit (Milankovitch cycles) . . . . . . . . . . . . . . . . . . . . . . . 71.6.6 Variations in the Sun’s energy output . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81.6.7 Volcanism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

1.7 Recent glacial and interglacial phases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81.7.1 Glacial stages in North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81.7.2 Last Glacial Period in the semiarid Andes around Aconcagua and Tupungato . . . . . . . . 9

1.8 Effects of glaciation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91.9 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101.10 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101.11 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

2 Megafauna 142.1 Ecological strategy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142.2 Evolution of large body size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

2.2.1 In terrestrial mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152.2.2 In marine mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152.2.3 In flightless birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

2.3 Megafaunal mass extinctions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162.3.1 Timing and possible causes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162.3.2 Consequences of depletion of megafauna . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

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2.4 Examples . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172.5 Gallery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

2.5.1 Extinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212.5.2 Living . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

2.6 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232.7 Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232.8 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232.9 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

3 Pleistocene 273.1 Dating . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273.2 Paleogeography and climate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

3.2.1 Glacial features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283.2.2 Major events . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283.2.3 Palaeocycles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

3.3 Fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303.4 Humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303.5 Deposits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313.6 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313.7 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313.8 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

4 Prehistoric mammal 334.1 List of prehistoric mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334.2 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

5 Stone Age 345.1 Historical significance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345.2 The Stone Age in archaeology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

5.2.1 Beginning of the Stone Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345.2.2 End of the Stone Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355.2.3 The concept of Stone Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355.2.4 The three-stage system . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 365.2.5 The problem of the transitions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

5.3 Chronology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375.3.1 Three-age chronology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375.3.2 Three-stage chronology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

5.4 Material culture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435.4.1 Tools . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435.4.2 Food and drink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435.4.3 Shelter and habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 445.4.4 Art . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

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5.4.5 Stone Age rituals and beliefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 455.5 Modern popular culture and the Stone Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 455.6 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 455.7 Notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 465.8 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475.9 Further reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475.10 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

6 Woolly mammoth 496.1 Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

6.1.1 Etymology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 506.1.2 Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

6.2 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 516.2.1 Coat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 526.2.2 Dentition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

6.3 Palaeobiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 546.3.1 Diet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 546.3.2 Growth and reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

6.4 Distribution and habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 566.5 Relationship with humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

6.5.1 Exploitation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 576.6 Extinction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 586.7 Frozen specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59

6.7.1 Recreating the species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 616.8 Cultural significance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62

6.8.1 Cryptozoology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 626.9 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 636.10 Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

7 Woolly rhinoceros 687.1 Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 687.2 Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 687.3 Behavior and habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

7.3.1 Diet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697.4 Extinction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697.5 See also . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707.6 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707.7 External links . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707.8 Text and image sources, contributors, and licenses . . . . . . . . . . . . . . . . . . . . . . . . . . 71

7.8.1 Text . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 717.8.2 Images . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 747.8.3 Content license . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

Chapter 1

Ice age

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

An artist’s impression of ice age Earth at glacial maximum.Based on: Crowley, T.J. (1995). “Ice age terrestrial carbonchanges revisited”. Global Biogeochemical Cycles 9 (3): 377–389. Bibcode:1995GBioC...9..377C. doi:10.1029/95GB01107.

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

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Variations in temperature, CO2, and dust from the Vostok ice core over the last 400,000 years

temperature of Earth's surface and atmosphere, resultingin the presence or expansion of continental and polar icesheets and alpine glaciers. Within a long-term ice age,individual pulses of cold climate are termed "glacial pe-riods" (or alternatively “glacials” or “glaciations” or col-loquially as “ice age”), and intermittent warm periods arecalled "interglacials". Glaciologically, ice age implies thepresence of extensive ice sheets in the northern and south-ern hemispheres.[1] By this definition, we are in an inter-glacial period—the holocene—of the ice age that began2.6 million years ago at the start of the Pleistocene epoch,because the Greenland, Arctic, and Antarctic ice sheetsstill exist.[2]

1.1 Origin of ice age theory

In 1742 Pierre Martel (1706–1767), an engineer and ge-ographer living in Geneva, visited the valley of Chamonixin the Alps of Savoy.[3][4] Two years later he publishedan account of his journey. He reported that the inhab-itants of that valley attributed the dispersal of erraticboulders to the fact that the glaciers had once extendedmuch farther.[5][6] Later similar explanations were re-ported from other regions of the Alps. In 1815 the car-penter and chamois hunter Jean-Pierre Perraudin (1767–1858) explained erratic boulders in the Val de Bagnes inthe Swiss canton of Valais as being due to glaciers previ-

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2 CHAPTER 1. ICE AGE

ously extending further.[7] An unknown woodcutter fromMeiringen in the Bernese Oberland advocated a simi-lar idea in a discussion with the Swiss-German geologistJean de Charpentier (1786–1855) in 1834.[8] Compara-ble explanations are also known from the Val de Ferretin the Valais and the Seeland in western Switzerland[9]and in Goethe's Scientific Work.[10] Such explanationscould also be found in other parts of the world. When theBavarian naturalist Ernst von Bibra (1806–1878) visitedthe Chilean Andes in 1849–1850 the natives attributedfossil moraines to the former action of glaciers.[11]

Meanwhile, European scholars had begun to wonder whathad caused the dispersal of erratic material. From themiddle of the 18th century some discussed ice as a meansof transport. The Swedish mining expert Daniel Tilas(1712–1772) was, in 1742, the first person to suggestdrifting sea ice in order to explain the presence of erraticboulders in the Scandinavian and Baltic regions.[12] In1795, the Scottish philosopher and gentleman naturalist,James Hutton (1726–1797), explained erratic boulders inthe Alps with the action of glaciers.[13] Two decades later,in 1818, the Swedish botanist Göran Wahlenberg (1780–1851) published his theory of a glaciation of the Scandi-navian peninsula. He regarded glaciation as a regionalphenomenon.[14] Only a few years later, the Danish-Norwegian Geologist Jens Esmark (1762–1839) argueda sequence of worldwide ice ages. In a paper published in1824, Esmark proposed changes in climate as the causeof those glaciations. He attempted to show that they orig-inated from changes in Earth’s orbit.[15] During the fol-lowing years, Esmark’s ideas were discussed and takenover in parts by Swedish, Scottish and German scien-tists. At the University of Edinburgh Robert Jameson(1774–1854) seemed to be relatively open to Esmark’sideas, as reviewed by Norwegian professor of glaciol-ogy Bjørn G. Andersen (1992).[16] Jameson’s remarksabout ancient glaciers in Scotland were most probablyprompted by Esmark.[17] In Germany, Albrecht Rein-hard Bernhardi (1797–1849), a geologist and professorof forestry at an academy in Dreissigacker, since incor-porated in the southern Thuringian city of Meiningen,adopted Esmark’s theory. In a paper published in 1832,Bernhardi speculated about former polar ice caps reach-ing as far as the temperate zones of the globe.[18]

Independently of these debates, the Swiss civil engineerIgnaz Venetz (1788–1859) in 1829, explained the dis-persal of erratic boulders in the Alps, the nearby JuraMountains and the North German Plain as being dueto huge glaciers. When he read his paper before theSchweizerische Naturforschende Gesellschaft, most sci-entists remained sceptical.[19] Finally, Venetz managedto convince his friend Jean de Charpentier. De Char-pentier transformed Venetz’s idea into a theory with aglaciation limited to the Alps. His thoughts resembledWahlenberg’s theory. In fact, both men shared the samevolcanistic, or in de Charpentier’s case rather plutonisticassumptions, about the earth’s history. In 1834, de Char-

pentier presented his paper before the SchweizerischeNaturforschende Gesellschaft.[20] In the meantime, theGerman botanist Karl Friedrich Schimper (1803–1867)was studying mosses which were growing on erratic boul-ders in the alpine upland of Bavaria. He began to wonderwhere such masses of stone had come from. During thesummer of 1835 he made some excursions to the Bavar-ian Alps. Schimper came to the conclusion that ice musthave been the means of transport for the boulders in thealpine upland. In the winter of 1835 to 1836 he heldsome lectures in Munich. Schimper then assumed thatthere must have been global times of obliteration (“Verö-dungszeiten”) 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 universityfriend Louis Agassiz (1801–1873) and Jean de Charp-entier. Schimper, de Charpentier and possibly Venetzconvinced Agassiz that there had been a time of glacia-tion. DuringWinter 1836/7 Agassiz and Schimper devel-oped the theory of a sequence of glaciations. Theymainlydrew upon the preceding works of Venetz, de Charpen-tier and on their own fieldwork. There are indications thatAgassiz was already familiar with Bernhardi’s paper atthat time.[22] At the beginning of 1837 Schimper coinedthe term ice age (“Eiszeit”).[23] In July 1837 Agassiz pre-sented their synthesis before the annual meeting of theSchweizerische Naturforschende Gesellschaft at Neuchâ-tel. The audience was very critical or even opposed thenew theory because it contradicted the established opin-ions on climatic history. Most contemporary scientiststhought that the earth had been gradually cooling downsince its birth as a molten globe.[24]

In order to overcome this rejection, Agassiz embarkedon geological fieldwork. He published his book Study onglaciers ("Études sur les glaciers”) in 1840.[25] De Char-pentier was put out by this as he had also been preparinga book about the glaciation of the Alps. De Charpentierfelt that Agassiz should have given him precedence as itwas he who had introduced Agassiz to in-depth glacialresearch.[26] Besides that, Agassiz had, as a result of per-sonal quarrels, omitted any mention of Schimper in hisbook.[27]

All together, it took several decades until the ice age the-ory was fully accepted. This happened on an internationalscale in the second half of the 1870s following the workof James Croll including the publication of Climate andTime, in Their Geological Relations in 1875 which pro-vided a credible explanation for the causes of ice ages.[28]

1.2 Evidence for ice ages

There are three main types of evidence for ice ages: ge-ological, chemical, and paleontological.Geological evidence for ice ages comes in various forms,including rock scouring and scratching, glacial moraines,

1.3. MAJOR ICE AGES 3

drumlins, valley cutting, and the deposition of till ortillites and glacial erratics. Successive glaciations tend todistort and erase the geological evidence, making it diffi-cult to interpret. Furthermore, this evidence was difficultto date exactly; early theories assumed that the glacialswere short compared to the long interglacials. The ad-vent of sediment and ice cores revealed the true situation:glacials are long, interglacials short. It took some time forthe current theory to be worked out.The chemical evidence mainly consists of variations inthe ratios of isotopes in fossils present in sediments andsedimentary rocks and ocean sediment cores. For themost recent glacial periods ice cores provide climateproxies from their ice, and atmospheric samples from in-cluded bubbles of air. Because water containing heavierisotopes has a higher heat of evaporation, its proportiondecreases with colder conditions.[29] This allows a tem-perature record to be constructed. However, this evi-dence can be confounded by other factors recorded byisotope ratios.The paleontological evidence consists of changes in thegeographical distribution of fossils. During a glacial pe-riod cold-adapted organisms spread into lower latitudes,and organisms that prefer warmer conditions become ex-tinct or are squeezed into lower latitudes. This evidence isalso difficult to interpret because it requires (1) sequencesof sediments covering a long period of time, over a widerange of latitudes and which are easily correlated; (2) an-cient organisms which survive for several million yearswithout change and whose temperature preferences areeasily diagnosed; and (3) the finding of the relevant fos-sils.Despite the difficulties, analysis of ice core and oceansediment cores[30] has shown periods of glacials and in-terglacials over the past few million years. These alsoconfirm the linkage between ice ages and continentalcrust phenomena such as glacial moraines, drumlins, andglacial erratics. Hence the continental crust phenomenaare accepted as good evidence of earlier ice ages whenthey are found in layers createdmuch earlier than the timerange for which ice cores and ocean sediment cores areavailable.

1.3 Major ice ages

There have been at least five major ice ages in the earth’spast (the Huronian, Cryogenian, Andean-Saharan, KarooIce Age and the Quaternary glaciation). Outside theseages, the Earth seems to have been ice-free even in highlatitudes.[31][32]

Rocks from the earliest well established ice age, calledthe Huronian, formed around 2.4 to 2.1 Ga (billion years)ago during the early Proterozoic Eon. Several hundredsof 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: Saaleglacial at maximum (Drenthe stage); blue: Elster glacial max-imum glaciation.

Timeline of glaciations, shown in blue.

tending from near Sault Ste. Marie to Sudbury, north-east of Lake Huron, with giant layers of now-lithified tillbeds, dropstones, varves, outwash, and scoured basementrocks. Correlative Huronian deposits have been foundnearMarquette, Michigan, and correlation has beenmadewith Paleoproterozoic glacial deposits fromWestern Aus-tralia.The next well-documented ice age, and probably the mostsevere of the last billion years, occurred from 850 to 630million years ago (the Cryogenian period) and may haveproduced a Snowball Earth in which glacial ice sheetsreached the equator,[33] possibly being ended by the ac-cumulation of greenhouse gases such as CO2 produced by volcanoes. “The presence of ice on thecontinents and pack ice on the oceans would inhibit bothsilicate weathering and photosynthesis, which are the twomajor sinks for CO2 at present.”[34] It has been suggested that the end of thisice age was responsible for the subsequent Ediacaran andCambrian Explosion, though this model is recent and con-troversial.The Andean-Saharan occurred from 460 to 420 millionyears ago, during the Late Ordovician and the Silurianperiod.The evolution of land plants at the onset of the Devonianperiod caused a long term increase in planetary oxygenlevels and reduction of CO2 levels, which resulted in the Karoo Ice Age. It is namedafter the glacial tills found in the Karoo region of SouthAfrica, where evidence for this ice age was first clearlyidentified. There were extensive polar ice caps at intervals

4 CHAPTER 1. ICE AGE

from 360 to 260 million years ago in South Africa duringthe Carboniferous and early Permian Periods. Correla-tives are known from Argentina, also in the center of theancient supercontinent Gondwanaland.

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Five Million Years ofClimate Change

From Sediment Cores

Millions of Years Ago

Sediment records showing the fluctuating sequences of glacialsand interglacials during the last several million years.

The current ice age, the Pliocene-Quaternary glacia-tion, started about 2.58 million years ago during the latePliocene, when the spread of ice sheets in the NorthernHemisphere began. Since then, the world has seen cy-cles of glaciation with ice sheets advancing and retreatingon 40,000- and 100,000-year time scales called glacialperiods, glacials or glacial advances, and interglacial pe-riods, interglacials or glacial retreats. The earth is cur-rently in an interglacial, and the last glacial period endedabout 10,000 years ago. All that remains of the continen-tal ice sheets are the Greenland and Antarctic ice sheetsand smaller glaciers such as on Baffin Island.Ice ages can be further divided by location and time;for example, the names Riss (180,000–130,000 years bp)andWürm (70,000–10,000 years bp) refer specifically toglaciation in the Alpine region. The maximum extent ofthe ice is not maintained for the full interval. The scour-ing action of each glaciation tends to remove most of theevidence of prior ice sheets almost completely, except inregions where the later sheet does not achieve full cover-age.

1.4 Glacials and interglacials

See also: Glacial period and InterglacialWithin the ice ages (or at least within the current one),more temperate and more severe periods occur. Thecolder periods are called glacial periods, the warmer pe-riods interglacials, such as the Eemian Stage.Glacials are characterized by cooler and drier climatesover most of the earth and large land and sea ice massesextending outward from the poles. Mountain glaciers inotherwise unglaciated areas extend to lower elevationsdue to a lower snow line. Sea levels drop due to the re-moval of large volumes of water above sea level in theicecaps. There is evidence that ocean circulation patternsare disrupted by glaciations. Since the earth has signifi-cant continental glaciation in the Arctic and Antarctic, weare currently in a glacial minimum of a glaciation. Sucha period between glacial maxima is known as an inter-glacial. The glacials and interglacials also coincided with

Shows the pattern of temperature and ice volume changes asso-ciated with recent glacials and interglacials

Minimum (interglacial, black) and maximum (glacial, grey)glaciation of the northern hemisphere

changes in Earth’s orbit called Milankovitch cycles.The earth has been in an interglacial period known asthe Holocene for more than 11,000 years. It was con-ventional wisdom that the typical interglacial period lastsabout 12,000 years, but this has been called into ques-tion recently. For example, an article in Nature[35] ar-gues that the current interglacial might be most analo-gous to a previous interglacial that lasted 28,000 years.Predicted changes in orbital forcing suggest that the nextglacial periodwould begin at least 50,000 years from now,even in absence of human-made global warming[36] (seeMilankovitch cycles). Moreover, anthropogenic forc-ing from increased greenhouse gases might outweigh or-bital forcing for as long as intensive use of fossil fuelscontinues.[37]

1.6. CAUSES OF ICE AGES 5

Minimum (interglacial, black) and maximum (glacial, grey)glaciation of the southern hemisphere

1.5 Positive and negative feedbackin glacial periods

Each glacial period is subject to positive feedback whichmakes it more severe and negative feedback which miti-gates and (in all cases so far) eventually ends it.

1.5.1 Positive feedback processes

Ice and snow increase Earth’s albedo, i.e. they make itreflect more of the sun’s energy and absorb less. Hence,when the air temperature decreases, ice and snow fieldsgrow, and this continues until competition with a negativefeedback mechanism forces the system to an equilibrium.Also, the reduction in forests caused by the ice’s expan-sion increases albedo.Another theory proposed by Ewing and Donn in 1956[38]hypothesized that an ice-free Arctic Ocean leads to in-creased snowfall at high latitudes. When low-temperatureice covers the Arctic Ocean there is little evaporation orsublimation and the polar regions are quite dry in termsof precipitation, comparable to the amount found inmid-latitude deserts. This low precipitation allows high-latitude snowfalls to melt during the summer. An ice-free Arctic Ocean absorbs solar radiation during the longsummer days, and evaporates more water into the Arcticatmosphere. With higher precipitation, portions of thissnow may not melt during the summer and so glacial icecan form at lower altitudes and more southerly latitudes,reducing the temperatures over land by increased albedoas noted above. Furthermore, under this hypothesis thelack of oceanic pack ice allows increased exchange of wa-ters between the Arctic and the North Atlantic Oceans,warming the Arctic and cooling the North Atlantic. (Cur-

rent projected consequences of global warming includea largely ice-free Arctic Ocean within 5–20 years, seeArctic shrinkage.) Additional fresh water flowing intothe North Atlantic during a warming cycle may also re-duce the global ocean water circulation (see Shutdown ofthermohaline circulation). Such a reduction (by reduc-ing the effects of the Gulf Stream) would have a coolingeffect on northern Europe, which in turn would lead toincreased 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 toblock the Gulf Stream.

1.5.2 Negative feedback processes

Ice sheets that form during glaciations cause erosion ofthe land beneath them. After some time, this will re-duce land above sea level and thus diminish the amountof space on which ice sheets can form. This mitigates thealbedo feedback, as does the lowering in sea level thataccompanies the formation of ice sheets.Another factor is the increased aridity occurring withglacial maxima, which reduces the precipitation availableto maintain glaciation. The glacial retreat induced by thisor any other process can be amplified by similar inversepositive feedbacks as for glacial advances.According to research published in Nature Geoscience,human emissions of carbon dioxide will defer the next iceage. Researchers used data on Earth’s orbit to find the his-torical warm interglacial period that looks most like thecurrent one and from this have predicted that the next iceage would usually begin within 1,500 years. They go onto say that emissions have been so high that it will not.[39]

1.6 Causes of ice ages

The causes of ice ages are not fully understood for eitherthe large-scale ice age periods or the smaller ebb and flowof glacial–interglacial periods within an ice age. The con-sensus is that several factors are important: atmosphericcomposition, such as the concentrations of carbon diox-ide and methane (the specific levels of the previouslymentioned gases are now able to be seen with the newice core samples from EPICADome C in Antarctica overthe past 800,000 years[40] ); changes in the earth’s orbitaround the Sun known as Milankovitch cycles; the mo-tion of tectonic plates resulting in changes in the rela-tive location and amount of continental and oceanic cruston the earth’s surface, which affect wind and ocean cur-rents; variations in solar output; the orbital dynamicsof the Earth-Moon system; and the impact of relativelylarge meteorites, and volcanism including eruptions ofsupervolcanoes.Some of these factors influence each other. For exam-

6 CHAPTER 1. ICE AGE

ple, changes in Earth’s atmospheric composition (espe-cially the concentrations of greenhouse gases) may alterthe climate, while climate change itself can change theatmospheric composition (for example by changing therate at which weathering removes CO2).Maureen Raymo, William Ruddiman and others proposethat the Tibetan and Colorado Plateaus are immense CO2 “scrubbers” with a capacity to remove enough CO2 from the global atmosphere to be a significant causalfactor of the 40 million year Cenozoic Cooling trend.They further claim that approximately half of their up-lift (and CO2 “scrubbing” capacity) occurred in the past 10 millionyears.[41][42]

1.6.1 Changes in Earth’s atmosphere

There is considerable evidence that over the very recentperiod of the last 100–1000 years, the sharp increasesin human activity, especially the burning of fossil fuels,has caused the parallel sharp and accelerating increase inatmospheric greenhouse gases which trap the sun’s heat.The consensus theory of the scientific community is thatthe resulting greenhouse effect is a principal cause ofthe increase in global warming which has occurred overthe same period, and a chief contributor to the acceler-ated melting of the remaining glaciers and polar ice. A2012 investigation finds that dinosaurs released methanethrough digestion in a similar amount to humanity’s cur-rent methane release, which “could have been a key fac-tor” to the very warm climate 150 million years ago.[43]

There is evidence that greenhouse gas levels fell at thestart of ice ages and rose during the retreat of the icesheets, but it is difficult to establish cause and effect (seethe notes above on the role of weathering). Greenhousegas levels may also have been affected by other factorswhich have been proposed as causes of ice ages, such asthe movement of continents and volcanism.The Snowball Earth hypothesis maintains that the severefreezing in the late Proterozoic was ended by an increasein CO2 levels in the atmosphere, and some supporters of Snow-ball Earth argue that it was caused by a reduction in at-mospheric CO2. The hypothesis also warns of future Snowball Earths.In 2009, further evidence was provided that changes insolar insolation provide the initial trigger for the earth towarm after an Ice Age, with secondary factors like in-creases in greenhouse gases accounting for the magnitudeof the change.[44]

William Ruddiman has proposed the early anthropocenehypothesis, according to which the anthropocene era, assome people call the most recent period in the earth’s his-tory when the activities of the human species first began

to have a significant global impact on the earth’s climateand ecosystems, did not begin in the 18th century with theadvent of the Industrial Era, but dates back to 8,000 yearsago, due to intense farming activities of our early agrarianancestors. It was at that time that atmospheric greenhousegas concentrations stopped following the periodic patternof the Milankovitch cycles. In his overdue-glaciation hy-pothesis Ruddiman states that an incipient glacial wouldprobably have begun several thousand years ago, but thearrival of that scheduled glacial was forestalled by the ac-tivities of early farmers.[45]

At a meeting of the American Geophysical Union (De-cember 17, 2008), scientists detailed evidence in sup-port of the controversial idea that the introduction oflarge-scale rice agriculture in Asia, coupled with exten-sive deforestation in Europe began to alter world climateby pumping significant amounts of greenhouse gasesinto the atmosphere over the last 1,000 years. In turn,a warmer atmosphere heated the oceans making themmuch less efficient storehouses of carbon dioxide and re-inforcing global warming, possibly forestalling the onsetof a new glacial age.[46]

1.6.2 Position of the continents

The geological record appears to show that ice ages startwhen the continents are in positions which block or re-duce the flow of warmwater from the equator to the polesand thus allow ice sheets to form. The ice sheets increaseEarth’s reflectivity and thus reduce the absorption of solarradiation. With less radiation absorbed the atmospherecools; the cooling allows the ice sheets to grow, whichfurther increases reflectivity in a positive feedback loop.The ice age continues until the reduction in weatheringcauses an increase in the greenhouse effect.There are three known configurations of the continentswhich block or reduce the flow of warm water from theequator to the poles:

• A continent sits on top of a pole, as Antarctica doestoday.

• A polar sea is almost land-locked, as the ArcticOcean is today.

• A supercontinent covers most of the equator, asRodinia did during the Cryogenian period.

Since today’s Earth has a continent over the South Poleand an almost land-locked ocean over the North Pole,geologists believe that Earth will continue to experienceglacial periods in the geologically near future.Some scientists believe that the Himalayas are a majorfactor in the current ice age, because these mountainshave increased Earth’s total rainfall and therefore the rateat which carbon dioxide is washed out of the atmosphere,

1.6. CAUSES OF ICE AGES 7

decreasing the greenhouse effect.[42] The Himalayas’ for-mation started about 70 million years ago when the Indo-Australian Plate collided with the Eurasian Plate, and theHimalayas are still rising by about 5 mm per year becausethe Indo-Australian plate is still moving at 67 mm/year.The history of the Himalayas broadly fits the long-termdecrease in Earth’s average temperature since the mid-Eocene, 40 million years ago.

1.6.3 Fluctuations in ocean currents

Another important contribution to ancient climateregimes is the variation of ocean currents, which aremodified 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 abil-ity to warm (e.g. giving the British Isles a temperate asopposed to a boreal climate). The closing of the Isthmusof Panama about 3 million years ago may have ushered inthe present period of strong glaciation over North Amer-ica by ending the exchange of water between the tropicalAtlantic and Pacific Oceans.[47]

Analyses suggest that ocean current fluctuations can ade-quately account for recent glacial oscillations. During thelast glacial period the sea-level has fluctuated 20–30 m aswater was sequestered, primarily in the northern hemi-sphere ice sheets. When ice collected and the sea leveldropped sufficiently, flow through the Bering Strait (thenarrow strait between Siberia and Alaska is ~50 m deeptoday) was reduced, resulting in increased flow from theNorth Atlantic. This realigned the thermohaline circu-lation in the Atlantic, increasing heat transport into theArctic, which melted the polar ice accumulation and re-duced other continental ice sheets. The release of waterraised sea levels again, restoring the ingress of colder wa-ter from the Pacific with an accompanying shift to north-ern hemisphere ice accumulation.[48]

1.6.4 Uplift of the Tibetan plateau and sur-rounding mountain areas above thesnowline

Matthias Kuhle’s geological theory of Ice Age develop-ment was suggested by the existence of an ice sheet cover-ing the Tibetan plateau during the Ice Ages (Last GlacialMaximum?). According to Kuhle, the plate-tectonic up-lift of Tibet past the snow-line has led to a surface of c.2,400,000 square kilometres (930,000 sq mi) changingfrom bare land to ice with a 70% greater albedo. The re-flection of energy into space resulted in a global cooling,triggering the Pleistocene Ice Age. Because this highlandis at a subtropical latitude, with 4 to 5 times the insolationof high-latitude areas, what would be Earth’s strongestheating surface has turned into a cooling surface.Kuhle explains the interglacial periods by the 100,000-

year cycle of radiation changes due to variations in Earth’sorbit. This comparatively insignificant warming, whencombined with the lowering of the Nordic inland ice areasand Tibet due to the weight of the superimposed ice-load,has led to the repeated complete thawing of the inland iceareas.[49][50][51][52]

1.6.5 Variations in Earth’s orbit (Mi-lankovitch cycles)

The Milankovitch cycles are a set of cyclic variations incharacteristics of the Earth’s orbit around the Sun. Eachcycle has a different length, so at some times their effectsreinforce each other and at other times they (partially)cancel each other.

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

There is strong evidence that the Milankovitch cyclesaffect the occurrence of glacial and interglacial periodswithin an ice age. The present ice age is the most stud-ied and best understood, particularly the last 400,000years, since this is the period covered by ice cores thatrecord atmospheric composition and proxies for temper-ature and ice volume. Within this period, the match ofglacial/interglacial frequencies to the Milanković orbitalforcing periods is so close that orbital forcing is gener-ally accepted. The combined effects of the changing dis-tance to the Sun, the precession of the Earth’s axis, andthe changing tilt of the Earth’s axis redistribute the sun-light received by the Earth. Of particular importance arechanges in the tilt of the Earth’s axis, which affect the in-tensity of seasons. For example, the amount of solar in-flux in July at 65 degrees north latitude varies by as muchas 22% (from 450 W/m² to 550 W/m²). It is widely be-lieved that ice sheets advance when summers become toocool to melt all of the accumulated snowfall from the pre-vious winter. Some workers believe that the strength ofthe orbital forcing is too small to trigger glaciations, butfeedback mechanisms like CO2 may explain this mismatch.While Milankovitch forcing predicts that cyclic changesin the Earth’s orbital elements can be expressed in theglaciation record, additional explanations are necessary toexplain which cycles are observed to be most importantin the timing of glacial–interglacial periods. In partic-ular, during the last 800,000 years, the dominant periodof glacial–interglacial oscillation has been 100,000 years,which corresponds to changes in Earth’s orbital eccentric-ity and orbital inclination. Yet this is by far the weakestof the three frequencies predicted by Milankovitch. Dur-

8 CHAPTER 1. ICE AGE

ing the period 3.0–0.8 million years ago, the dominantpattern of glaciation corresponded to the 41,000-year pe-riod of changes in Earth’s obliquity (tilt of the axis). Thereasons for dominance of one frequency versus anotherare poorly understood and an active area of current re-search, but the answer probably relates to some form ofresonance in the Earth’s climate system.The “traditional” Milankovitch explanation struggles toexplain the dominance of the 100,000-year cycle over thelast 8 cycles. Richard A. Muller, Gordon J. F. MacDon-ald,[53][54][55] and others have pointed out that those cal-culations are for a two-dimensional orbit of Earth but thethree-dimensional orbit also has a 100,000-year cycle oforbital inclination. They proposed that these variationsin orbital inclination lead to variations in insolation, asthe Earth moves in and out of known dust bands in thesolar system. Although this is a different mechanism tothe traditional view, the “predicted” periods over the last400,000 years are nearly the same. The Muller and Mac-Donald theory, in turn, has been challenged by Jose An-tonio Rial.[56]

Another worker, William Ruddiman, has suggesteda model that explains the 100,000-year cycle by themodulating effect of eccentricity (weak 100,000-year cy-cle) on precession (26,000-year cycle) combined withgreenhouse gas feedbacks in the 41,000- and 26,000-year cycles. Yet another theory has been advanced byPeter Huybers who argued that the 41,000-year cycle hasalways been dominant, but that the Earth has entereda mode of climate behavior where only the second orthird cycle triggers an ice age. This would imply thatthe 100,000-year periodicity is really an illusion createdby averaging together cycles lasting 80,000 and 120,000years.[57] This theory is consistent with a simple empiricalmulti-state model proposed by Didier Paillard.[58] Pail-lard suggests that the late Pleistocene glacial cycles can beseen as jumps between three quasi-stable climate states.The jumps are induced by the orbital forcing, while inthe early Pleistocene the 41,000-year glacial cycles re-sulted from jumps between only two climate states. Adynamical model explaining this behavior was proposedby Peter Ditlevsen.[59] This is in support of the suggestionthat the late Pleistocene glacial cycles are not due to theweak 100,000-year eccentricity cycle, but a non-linear re-sponse to mainly the 41,000-year obliquity cycle.

1.6.6 Variations in the Sun’s energy output

There are at least two types of variation in the Sun’s en-ergy output

• In the very long term, astrophysicists believe that theSun’s output increases by about 7% every one billion(109) years.

• Shorter-term variations such as sunspot cycles, andlonger episodes such as the Maunder minimum,

which occurred during the coldest part of the LittleIce Age.

The long-term increase in the Sun’s output cannot be acause of ice ages.

1.6.7 Volcanism

Volcanic eruptions may have contributed to the incep-tion and/or the end of ice age periods. At times dur-ing the paleoclimate, carbon dioxide levels were two orthree times greater than today. Volcanoes and move-ments in continental plates contributed to high amountsof CO2 in the atmosphere. Carbon dioxide from volca-noes probably contributed to periods with highest over-all temperatures.[60] One suggested explanation of thePaleocene-Eocene Thermal Maximum is that underseavolcanoes released methane from clathrates and thuscaused a large and rapid increase in the greenhouse ef-fect. There appears to be no geological evidence for sucheruptions at the right time, but this does not prove theydid not happen.

1.7 Recent glacial and interglacialphases

Northern hemisphere glaciation during the last ice ages. The setup of 3 to 4 km thick ice sheets caused a sea level lowering ofabout 120 m.

Main article: Timeline of glaciation

1.7.1 Glacial stages in North America

The major glacial stages of the current ice age in NorthAmerica are the Illinoian, Sangamonian and Wisconsinstages. The use of the Nebraskan, Afton, Kansan,and Yarmouthian (Yarmouth) stages to subdivide the iceage in North America have been discontinued by Qua-ternary geologists and geomorphologists. These stages

1.8. EFFECTS OF GLACIATION 9

have all been merged into the Pre-Illinoian Stage in the1980s.[61][62][63]

During the most recent North American glaciation, dur-ing the latter part of the Wisconsin Stage (26,000 to13,300 years ago), ice sheets extended to about 45 de-grees north latitude. These sheets were 3 to 4 kmthick.[62]

This Wisconsin glaciation left widespread impacts onthe North American landscape. The Great Lakes andthe Finger Lakes were carved by ice deepening old val-leys. Most of the lakes in Minnesota and Wisconsin weregouged out by glaciers and later filled with glacial meltwa-ters. The old Teays River drainage system was radicallyaltered and largely reshaped into the Ohio River drainagesystem. Other rivers were dammed and diverted to newchannels, such as the Niagara, which formed a dramaticwaterfall and gorge, when the waterflow encountered alimestone escarpment. Another similar waterfall, at thepresent Clark Reservation State Park near Syracuse, NewYork, is now dry.The area fromLong Island to Nantucket was formed fromglacial till, and the plethora of lakes on the CanadianShield in northern Canada can be almost entirely at-tributed to the action of the ice. As the ice retreated andthe rock dust dried, winds carried the material hundredsof miles, forming beds of loess many dozens of feet thickin the Missouri Valley. Isostatic rebound continues to re-shape the Great Lakes and other areas formerly under theweight of the ice sheets.The Driftless Zone, a portion of western and southwest-ern Wisconsin along with parts of adjacent Minnesota,Iowa, and Illinois, was not covered by glaciers.See also: Glacial history of Minnesota

1.7.2 Last Glacial Period in the semi-arid Andes around Aconcagua andTupungato

A specially interesting climatic change during glacialtimes has taken place in the semi-arid Andes. Be-side the expected cooling down in comparison with thecurrent climate, a significant precipitation is concernedhere. So, researches in the presently semiarid subtropicAconcagua-massif (6,962 m) have shown an unexpect-edly extensive glacial glaciation of the type “ice streamnetwork”.[64][65][66][67][68] The connected valley glaciersexceeding 100 km in length, flowed down on the East-side of this section of the Andes at 32–34°S and 69–71°W as far as a height of 2,060 m and on the westernluff-side still clearly deeper.[68][69] Where current glaciersscarcely reach 10 km in length, the snowline (ELA) runsat a height of 4,600 m and at that time was lowered to3,200 m asl, i.e. about 1,400 m. From this follows that—beside of an annual depression of temperature about c.

8.4°C— here was an increase in precipitation. Accord-ingly, at glacial times the humid climatic belt that todayis situated several latitude degrees further to the S, wasshifted much further to the N.[67][68]

1.8 Effects of glaciation

Scandinavia exhibits some of the typical effects of ice age glacia-tion such as fjords and lakes.

See also: Glacial landforms

Although the last glacial period ended more than 8,000years ago, its effects can still be felt today. For exam-ple, the moving ice carved out the landscape in Canada(See Canadian Arctic Archipelago), Greenland, north-ern Eurasia and Antarctica. The erratic boulders, till,drumlins, eskers, fjords, kettle lakes, moraines, cirques,horns, etc., are typical features left behind by the glaciers.The weight of the ice sheets was so great that they de-formed the Earth’s crust and mantle. After the ice sheetsmelted, the ice-covered land rebounded. Due to the highviscosity of the Earth’s mantle, the flow of mantle rockswhich controls the rebound process is very slow—at a rateof about 1 cm/year near the center of rebound area today.During glaciation, water was taken from the oceans toform the ice at high latitudes, thus global sea leveldropped by about 110 meters, exposing the continentalshelves and forming land-bridges between land-massesfor animals to migrate. During deglaciation, the meltedice-water returned to the oceans, causing sea level to rise.This process can cause sudden shifts in coastlines and

10 CHAPTER 1. ICE AGE

hydration systems resulting in newly submerged lands,emerging lands, collapsed ice dams resulting in salinationof lakes, new ice dams creating vast areas of freshwater,and a general alteration in regional weather patterns ona large but temporary scale. It can even cause tempo-rary reglaciation. This type of chaotic pattern of rapidlychanging land, ice, saltwater and freshwater has been pro-posed as the likely model for the Baltic and Scandinavianregions, as well as much of central North America at theend of the last glacial maximum, with the present-daycoastlines only being achieved in the last few millenniaof prehistory. Also, the effect of elevation on Scandi-navia submerged a vast continental plain that had existedunder much of what is now the North Sea, connecting theBritish Isles to Continental Europe.[70]

The redistribution of ice-water on the surface of theEarth and the flow of mantle rocks causes changes in thegravitational field as well as changes to the distributionof the moment of inertia of the Earth. These changes tothe moment of inertia result in a change in the angularvelocity, axis, and wobble of the Earth’s rotation.The weight of the redistributed surface mass loaded thelithosphere, caused it to flex and also induced stress withinthe Earth. The presence of the glaciers generally sup-pressed the movement of faults below.[71][72][73] How-ever, during deglaciation, the faults experience acceler-ated slip triggering earthquakes. Earthquakes triggerednear the ice margin may in turn accelerate ice calving andmay account for the Heinrich events.[74] Asmore ice is re-moved near the ice margin, more intraplate earthquakesare induced and this positive feedback may explain thefast collapse of ice sheets.In Europe, glacial erosion and isostatic sinking fromweight of ice made the Baltic Sea, which before the IceAge 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 mys-

tery. Short Hills NJ: Enslow Publishers. ISBN 978-0-89490-015-0.

[2] Gribbin, J.R. (1982). Future Weather: Carbon Diox-ide, Climate and the Greenhouse Effect. Penguin. ISBN0140224599.

[3] Rémis, F.; Testus, L.; Testut (2006). “Mais com-ment s’écoule donc un glacier ? Aperçu his-torique” (PDF). C. R. Geoscience (in French) 338(5): 368–385. 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) Anaccount of the glacieres or ice alps in Savoy, in two letters,one from an English gentleman to his friend at Geneva ;the other from PierreMartel , engineer, to the said Englishgentleman”. In Mathews, C.E. The annals of Mont Blanc.London: Unwin. p. 327. See (Montgomery 2010) for afull bibliography

[6] Krüger, Tobias (2013). Discovering the Ice Ages. Interna-tional Reception and Consequences for a Historical Under-standing of Climate (German editon: Basel 2008). Leiden.p. 47. ISBN 978-90-04-24169-5.

[7] Krüger 2013, pp. 78-83

[8] Krüger 2013, p. 150

[9] Krüger 2013, pp. 83, 151

[10] Goethe, Johann Wolfgang von: Geologische Problemeund Versuch ihrer Auflösung, Mineralogie und Geologiein Goethes Werke, Weimar 1892, ISBN 3-423-05946-X,book 73 (WA II,9), p. 253, 254.

[11] Krüger 2013, p. 83

[12] Krüger 2013, p. 38

[13] Krüger 2013, pp. 61-2

[14] Krüger 2013, pp. 88–90

[15] Krüger 2013, pp. 91-6

[16] Andersen, Bjørn G. (1992). "Jens Esmark—a pioneer inglacial geology” 21. Boreas. pp. 97–102.

[17] Davies, Gordon L. (1969). The Earth in Decay. A His-tory of British Geomorphology 1578–1878. London. pp.267f.Cunningham, Frank F. (1990). James David Forbes. Pio-neer Scottish Glaciologist. Edinburgh: Scottish AcademicPress. p. 15. ISBN 0707303206.

[18] Krüger 2013, pp. 142-47

[19] Krüger 2013, pp. 104–05

[20] Krüger 2013, pp. 150–53

[21] Krüger 2013, pp. 155–59

[22] Krüger 2013, pp. 167-70

[23] Krüger 2013, p. 173

[24] Krüger 2008, pp. 177–78

1.10. REFERENCES 11

[25] Agassiz, Louis; Bettannier, Joseph (1840). Études sur lesglaciers. Ouvrage accompagné d'un atlas de 32 planches,Neuchâtel. H. Nicolet.

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[31] Lockwood, J.G.; van Zinderen-Bakker, E. M. (Novem-ber 1979). “The Antarctic Ice-Sheet: Regulator of GlobalClimates?: Review”. The Geographical Journal 145 (3):469–471. doi:10.2307/633219. JSTOR 633219.

[32] Warren, John K. (2006). Evaporites: sediments, resourcesand hydrocarbons. Birkhäuser. p. 289. ISBN 978-3-540-26011-0.

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[34] Chris Clowes. ""Snowball” Scenarios of the Cryogenian”.Paleos: Life through deep time. Archived from the originalon 20 Dec 2010. Retrieved April 2012.

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[36] Berger A, Loutre MF (August 2002). “Climate. Anexceptionally long interglacial ahead?". Science 297(5585): 1287–8. doi:10.1126/science.1076120. PMID12193773.

[37] “Next Ice Age Delayed By Rising Carbon Dioxide Lev-els”. ScienceDaily. 2007. Retrieved 2008-02-28.

[38] Ewing, M.; Donn, W.L.; Donn (June 1956).“A Theory of Ice Ages”. Science 123(3207): 1061–6. Bibcode:1956Sci...123.1061E.doi:10.1126/science.123.3207.1061. PMID 17748617.

[39] Black, Richard (9 January 2012). “Carbon emissions 'willdefer Ice Age'". BBC News. Retrieved 10 August 2012.

[40] Luthi, Dieter; et al. (2008-03-17). “High-resolution carbon dioxide concentration record650,000–800,000 years before present”. Nature453 (7193): 379–382. Bibcode:2008Natur.453..379L.doi:10.1038/nature06949. PMID 18480821.

[41] Ruddiman, W.F.; Kutzbach, J.E. (1991). “PlateauUplift and Climate Change”. Scientific American264 (3): 66–74. Bibcode:1991SciAm.264...66R.doi:10.1038/scientificamerican0391-66.

[42] Raymo, M.E.; Ruddiman, W.F.; Froelich, P.N.;Ruddiman; Froelich (July 1988). “Influence oflate Cenozoic mountain building on ocean geo-chemical cycles”. Geology 16 (7): 649–653.Bibcode:1988Geo....16..649R. doi:10.1130/0091-7613(1988)016<0649:IOLCMB>2.3.CO;2.

[43] Davies, Ella (2012-05-07). “BBCNature - Dinosaur gases'warmed the Earth'". Bbc.co.uk. Retrieved 2012-08-07.

[44] Clark, Peter U.; Dyke, Arthur S.; Shakun, Jeremy D.;Carlson, Anders E.; Clark, Jorie; Wohlfarth, Barbara;Mitrovica, Jerry X.; Hostetler, Steven W. & McCabe, A.Marshall (2009). “The Last Glacial Maximum”. Science325 (5941): 710–714. Bibcode:2009Sci...325..710C.doi:10.1126/science.1172873. PMID 19661421.

[45] Ruddiman, William F. (2003). “The Anthro-pogenic Greenhouse Era Began Thousands ofYears Ago”. Climatic Change 61 (3): 261–293.doi:10.1023/B:CLIM.0000004577.17928.fa.

[46] Did Early Climate Impact Divert a New Glacial Age?Newswise, Retrieved on December 17, 2008.

[47] Svitil, K.A. (April 1996). “We are all Panamanians”. Dis-cover. Retrieved April 2012.—formation of Isthmus ofPanama may have started a series of climatic changes thatled to evolution of hominids

[48] Hu, Aixue; Gerald Meehl, Bette L. Otto-Bliesner, ClaireWaelbroeck, Weiqing Han, Marie-France Loutre, KurtLambeck, Jerry X. Mitrovica & Nan Rosenbloom (2010).“Influence of Bering Strait flow and North Atlantic cir-culation on glacial sea-level changes”. Nature Geo-science 3 (2): 118. Bibcode:2010NatGe...3..118H.doi:10.1038/ngeo729.

[49] Kuhle, Matthias (December 1988). “Tibet and High-Asia: Results of the Sino-German Joint Expeditions (I)".GeoJournal 17 (4): 581–595. JSTOR 41144345. |chap-ter= ignored (help)

[50] 2c (Quaternary Glaciation — Extent and Chronology,Part III: South America, Asia, Africa, Australia, Antarcti-caKuhle, M. (2004). “The High Glacial (Last Ice Age andLGM) ice cover in High and Central Asia”. In Ehlers,J.; Gibbard, P.L. Quaternary Glaciations: South Amer-ica, Asia, Africa, Australasia, Antarctica. Development inQuaternary Science: Quaternary Glaciations: Extent andChronology Vol. 3. Amsterdam: Elsevier. pp. 175–199.ISBN 978-0-444-51593-3.

[51] Kuhle, M. (1999). “Reconstruction of an approximatelycomplete Quaternary Tibetan inland glaciation betweenthe Mt. Everest- and Cho Oyu Massifs and the Aksai

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Chin. A new glaciogeomorphological SE–NW diagonalprofile through Tibet and its consequences for the glacialisostasy and Ice Age cycle”. GeoJournal 47 (1–2): 3–276.doi:10.1023/A:1007039510460.

[52] Kuhle, M. (2011). “Ice Age Development Theory”. InSingh, V.P.; Singh, P.; Haritashya, U.K. Encyclopedia ofSnow, Ice and Glaciers. Springer. pp. 576–581.

[53] Muller, R.A.; MacDonald, G.J.; MacDonald (August1997). “Spectrum of 100-kyr glacial cycle: orbital incli-nation, not eccentricity”. Proc. Natl. Acad. Sci. U.S.A.94 (16): 8329–34. Bibcode:1997PNAS...94.8329M.doi:10.1073/pnas.94.16.8329. PMC 33747. PMID11607741.

[54] Richard A. Muller. “A New Theory of Glacial Cycles”.Muller.lbl.gov. Retrieved 2012-08-07.

[55] Muller, R.A.; MacDonald, G.J.; MacDonald (July 1997).“Glacial Cycles and Astronomical Forcing”. Science277 (5323): 215–8. Bibcode:1997Sci...277..215M.doi:10.1126/science.277.5323.215.

[56] Rial, J.A. (July 1999). “Pacemaking the ice agesby frequency modulation of Earth’s orbital ec-centricity” (PDF). Science 285 (5427): 564–8.doi:10.1126/science.285.5427.564. PMID 10417382.

[57] Huybers, P.; Wunsch, C.; Wunsch (March2005). “Obliquity pacing of the late Pleis-tocene glacial terminations”. Nature 434(7032): 491–4. Bibcode:2005Natur.434..491H.doi:10.1038/nature03401. PMID 15791252.

[58] Paillard, D. (22 January 1998). “The timing ofPleistocene glaciations from a simple multiple-stateclimate model”. Nature 391 (6665): 378–381.Bibcode:1998Natur.391..378P. doi:10.1038/34891.

[59] Ditlevsen, P.D. (2009). “Bifurcation structureand noise-assisted transitions in the Pleistoceneglacial cycles”. Paleoceanography 24 (3): PA3204.arXiv:0902.1641. Bibcode:2009PalOc..24.3204D.doi:10.1029/2008PA001673. as PDF

[60] Rieke, George. “Long TermClimate”. Retrieved 25April2013.

[61] Hallberg, G.R. (1986). “Pre-Wisconsin glacial stratig-raphy of the Central Plains region in Iowa, Ne-braska, Kansas, and Missouri”. Quaternary ScienceReviews 5: 11–15. Bibcode:1986QSRv....5...11H.doi:10.1016/0277-3791(86)90169-1.

[62] Richmond, G.M.; Fullerton, D.S. (1986). “Summa-tion of Quaternary glaciations in the United Statesof America”. Quaternary Science Reviews 5: 183–196. Bibcode:1986QSRv....5..183R. doi:10.1016/0277-3791(86)90184-8.

[63] Gibbard, P.L., S. Boreham, K.M. Cohen and A.Moscariello, 2007, Global chronostratigraphical correla-tion table for the last 2.7 million years v. 2007b., jpgversion 844 KB. Subcommission on Quaternary Stratigra-phy, Department of Geography, University of Cambridge,Cambridge, England

[64] Kuhle, M. (1984). “Spuren hocheiszeitlicher Gletscherbe-deckung in der Aconcagua-Gruppe (32–33° S)". Zen-tralblatt für Geologie und Paläontologie Teil I, Geologie.11/12: 1635–46. ISSN 0340-5109. Verhandlungsblattdes Südamerika-Symposiums 1984 in Bamberg.

[65] Kuhle, M. (1986). “Die Vergletscherung Tibets und dieEntstehung von Eiszeiten”. Spektrum der Wissenschaft(9/86): 42–54. ISSN 0170-2971.

[66] Kuhle, Matthias (June 1987). “SubtropicalMountain- andHighland-Glaciation as Ice Age Triggers and the Wan-ing of the Glacial Periods in the Pleistocene”. GeoJour-nal 14 (4): 393–421. doi:10.1007/BF02602717. JSTOR41144132.

[67] Kuhle, M. (2004). “The Last Glacial Maximum (LGM)glacier cover of the Aconcagua group and adjacent mas-sifs in the Mendoza Andes (South America)". In Ehlers,J.; Gibbard, P.L. Quaternary Glaciations: South Amer-ica, Asia, Africa, Australasia, Antarctica. Development inQuaternary Science. Amsterdam: Elsevier. pp. 75–81.ISBN 978-0-444-51593-3.

[68] Kuhle, M. (2011). “Ch 53: The High-Glacial (LastGlacial Maximum) Glacier Cover of the AconcaguaGroup and Adjacent Massifs in the Mendoza Andes(South America) with a Closer Look at Further Empiri-cal Evidence”. In Ehlers, J.; Gibbard, P.L.; Hughes, P.D.Quaternary Glaciations – Extent and Chronology: A CloserLook. Development in Quaternary Science. Amsterdam:Elsevier. pp. 735–8. ISBN 978-0-444-53447-7.

[69] Brüggen, J. (1929). “Zur Glazialgeologie der chilenis-chen Anden”. Geol. Rundsch. 20 (1): 1–35.doi:10.1007/BF01805072.

[70] Andersen, Bjørn G.; Borns, Harold W. Jr. (1997).The Ice Age World: an introduction to quaternary his-tory and research with emphasis on North America andNorthern Europe during the last 2.5 million years. Oslo:Universitetsforlaget. ISBN 97-88200376-83-5. Re-trieved 2013-10-14.

[71] Johnston, A. (1989). “The effect of large ice sheets onearthquake genesis”. In Gregersen, S.; Basham, P. Earth-quakes at North-Atlantic passive margins: Neotectonicsand postglacial rebound. Dordrecht: Kluwer. pp. 581–599. ISBN 0792301501.

[72] Wu, P.; Hasegawa, H.S.; Hasegawa (October 1996).“Induced stresses and fault potential in eastern Canadadue to a realistic load: a preliminary analysis”.Geophysical Journal International 127 (1): 215–229.Bibcode:1996GeoJI.127..215W. doi:10.1111/j.1365-246X.1996.tb01546.x.

[73] Turpeinen, H.; Hampel, A.; Karow, T.; Maniatis, G.(2008). “Effect of ice sheet growth and melting on the slipevolution of thrust faults”. Earth and Planetary ScienceLetters 269: 230–241. Bibcode:2008E&PSL.269..230T.doi:10.1016/j.epsl.2008.02.017.

[74] Hunt, A.G.; Malin, P.E.; Malin (14 May 1998).“Possible triggering of Heinrich events by ice-load-induced earthquakes”. Nature 393 (6681): 155–8.Bibcode:1998Natur.393..155H. doi:10.1038/30218.

1.11. EXTERNAL LINKS 13

1.11 External links• Cracking the Ice Age from PBS

• Montgomery, Keith (2010). “Development of theglacial theory, 1800–1870”. Historical Simulation

• Raymo, M. (July 2011). “Overview of the Uplift-Weathering Hypothesis”.

• Eduard Y. Osipov ., Oleg M. Khlystov. Glaciersand meltwater flux to Lake Baikal during the LastGlacial Maximum.

• Black, R. (9 January 2012). “Carbon emissions 'willdefer Ice Age'". BBC News: Science and Environ-ment.

Chapter 2

Megafauna

This article is about large land animals. For naked-eyevisible bottom-dwelling animals, see Macrobenthos. Forgiant animals in mythology, see Megafauna (mythology).In terrestrial zoology,megafauna (Ancient Greekmegas

The African bush elephant, Earth’s largest living land animal

“large” + New Latin fauna “animal”) are large or giantanimals. The most common thresholds used are 45 kilo-grams (100 lb)[1][2] or 100 kilograms (220 lb).[2][3] Thisthus includes many species not popularly thought of asoverly large, such as white-tailed deer, red kangaroo, andhumans.In practice, the most common usage encountered in aca-demic and popular writing describes land animals roughlylarger than a human that are not (solely) domesticated.The term is especially associated with the Pleistocenemegafauna – the land animals often larger than moderncounterparts considered archetypical of the last ice age,such as mammoths, the majority of which in northern

Eurasia, the Americas and Australia became extinct asrecently as 10,000–40,000 years ago.[4] It is also com-monly used for the largest extant wild land animals, espe-cially elephants, giraffes, hippopotamuses, rhinoceroses,and large bovines. Megafauna may be subcategorizedby their trophic position into megaherbivores (e.g., elk),megacarnivores (e.g., lions), and, more rarely, megaom-nivores (e.g., bears).Other common uses are for giant aquatic species, espe-cially whales, any larger wild or domesticated land ani-mals such as larger antelope and cattle, as well as numer-ous dinosaurs and other extinct giant reptilians.The term is also sometimes applied to animals (usuallyextinct) of great size relative to a more common or sur-viving type of the animal, for example the 1 m (3 ft)dragonflies of the Carboniferous period.

2.1 Ecological strategy

Megafauna – in the sense of the largest mammals andbirds – are generally K-strategists, with high longevity,slow population growth rates, low mortality rates, and (atleast for the largest) few or no natural predators capable ofkilling adults. These characteristics, although not exclu-sive to such megafauna, make them vulnerable to humanoverexploitation, in part because of their slow populationrecovery rates.

2.2 Evolution of large body size

One observation that has been made about the evolutionof larger body size is that rapid rates of increase that areoften seen over relatively short time intervals are not sus-tainable over much longer time periods. In an exam-ination of mammal body mass changes over time, themaximum increase possible in a given time interval wasfound to scale with the interval length raised to the 0.25power.[5] This is thought to reflect the emergence, dur-ing a trend of increasing maximum body size, of a se-ries of anatomical, physiological, environmental, geneticand other constraints that must be overcome by evolu-

14

2.2. EVOLUTION OF LARGE BODY SIZE 15

tionary innovations before further size increases are pos-sible. A strikingly faster rate of change was found forlarge decreases in body mass, such as may be associatedwith the phenomenon of insular dwarfism. When nor-malized to generation length, the maximum rate of bodymass decrease was found to be over 30 times greater thanthe maximum rate of body mass increase for a ten-foldchange.[5]

2.2.1 In terrestrial mammals

Subsequent to the Cretaceous–Paleogene extinction eventthat eliminated the non-avian dinosaurs about 66 Ma ago,terrestrial mammals underwent a nearly exponential in-crease in body size as they diversified to occupy the eco-logical niches left vacant.[6] Starting from just a few kgbefore the event, maximum size had reached ~50 kg afew million years later, and ~750 kg by the end of thePaleocene. This trend of increasing body mass appearsto level off about 40 Ma ago (in the late Eocene), suggest-ing that physiological or ecological constraints had beenreached, after an increase in body mass of over three or-ders of magnitude.[6] However, when considered from thestandpoint of rate of size increase per generation, the ex-ponential increase is found to have continued until theappearance of Indricotherium 30 Ma ago. (Since gener-ation time scales with body mass0.259, increasing genera-tion times with increasing size cause the log mass vs. timeplot to curve downward from a linear fit.)[5]

Megaherbivores eventually attained a body mass of over10 000 kg. The largest of these, indricotheres andproboscids, have been hindgut fermenters, which are be-lieved to have an advantage over foregut fermenters interms of being able to accelerate gastrointestinal tran-sit in order to accommodate very large food intakes.[7]A similar trend emerges when rates of increase of max-imum body mass per generation for different mam-malian clades are compared (using rates averaged overmacroevolutionary time scales). Among terrestrial mam-mals, 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 ofwhich are hindgut fermenters. The rate of increase forartiodactyls (0.74) was about a third that of perisso-dactyls. The rate for carnivorans (0.65) was slightly loweryet, while primates, perhaps constrained by their arborealhabits, had the lowest rate (0.39) among the mammaliangroups studied.[5]

Terrestrial mammalian carnivores from several eutheriangroups (the mesonychid Andrewsarchus, the creodontsMegistotherium and Sarkastodon, and the carnivoransAmphicyon and Arctodus) all reached a maximum sizeof about 1000 kg[6] (the carnivoran Arctotherium appar-ently actually got somewhat larger). The largest knownmetatherian carnivore, Proborhyaena gigantea, appar-ently reached 600 kg, also close to this limit.[8] A similartheoretical maximum size for mammalian carnivores has

been predicted based on the metabolic rate of mammals,the energetic cost of obtaining prey, and themaximum es-timated rate coefficient of prey intake.[9] It has also beensuggested that maximum size for mammalian carnivoresis constrained by the stress the humerus can withstand attop running speed.[8]

Analysis of the variation of maximum body size over thelast 40 Ma suggests that decreasing temperature and in-creasing continental land area are associated with increas-ing maximum body size. The former correlation wouldbe consistent with Bergmann’s rule,[10] and might be re-lated to the thermoregulatory advantage of large bodymass in cool climates,[6] better ability of larger organ-isms to cope with seasonality in food supply,[10] or otherfactors;[10] the latter correlation could be explainable interms of range and resource limitations.[6] However, thetwo parameters are interrelated (due to sea level drops ac-companying increased glaciation), making the driver ofthe trends in maximum size more difficult to identify.[6]

2.2.2 In marine mammals

The ancestors of cetaceans are believed to have been thesemiaquatic pakicetids, no larger than wolves, of about53 million years (Ma) ago.[11] By 40 Ma ago, cetaceanshad attained a length of 20 m or more in Basilosaurus,an elongated, serpentine whale that differed from mod-ern whales in many respects and was not ancestral tothem. Following this, the evolution of large body sizein cetaceans appears to have come to a temporary halt,and then to have backtracked, although the available fossilrecords are limited. However, in the period from 31 Maago (in the Oligocene) to the present, cetaceans under-went a significantly more rapid sustained increase in bodymass (a rate of increase in bodymass0.259 of a factor of 3.2per million years) than achieved by any group of terres-trial mammals.[5] This trend led to the largest animal ofall time, the modern blue whale. Several reasons for themore rapid evolution of large body size in cetaceans arepossible. Fewer biomechanical constraints on increasesin body size may be associated with suspension in wateras opposed to standing against the force of gravity, andwith swimming movements as opposed to terrestrial lo-comotion. Also, the greater heat capacity and thermalconductivity of water compared to air may increase thethermoregulatory advantage of large body size in marineendotherms, although diminishing returns apply.[5]

Cetaceans are not the only marine mammals to reachunprecedented size in the modern era. The largestcarnivoran of all time is the mostly aquatic modernsouthern elephant seal.

2.2.3 In flightless birds

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

16 CHAPTER 2. MEGAFAUNA

tebrates had a roughly ten million year long window ofopportunity (during the Paleocene) for evolution of gi-gantism without much competition.[12] During this inter-val, apex predator niches were often occupied by reptiles,such as terrestrial crocodilians (e.g. Pristichampsus),large snakes (e.g. Titanoboa) or varanid lizards, or byflightless birds[6] (e.g. Gastornis in Europe and NorthAmerica, Paleopsilopterus in South America). This isalso the period when flightless herbivorous paleognathbirds evolved to large size on a number of Gondwananland masses. These birds, termed ratites, have tradi-tionally been viewed as representing a lineage separatefrom that of their small flighted relatives, the Neotropictinamous. However, recent genetic studies have foundthat tinamous nest well within the ratite tree, and are thesister group of the extinct moa of New Zealand.[13][12][14]Similarly, the small kiwi of NewZealand have been foundto be the sister group of the extinct elephant birds ofMadagascar.[12] These findings indicate that flightlessnessand gigantism arose independently multiple times amongratites via parallel evolution.In the northern continents, large predatory birds were dis-placed when large eutherian carnivores evolved. In iso-lated South America, the phorusrhacids could not be out-competed by the local metatherian sparassodonts and re-mained dominant until advanced eutherian predators ar-rived from North America (as part of the Great Amer-ican Interchange) during the Pliocene. However, noneof the largest predatory (Brontornis), possibly omnivo-rous (Dromornis[15]) or herbivorous (Aepyornis) flightlessbirds of the Cenozoic ever grew to masses much above500 kg, and thus never attained the size of the largestmammalian carnivores, let alone that of the largest mam-malian herbivores. It has been suggested that the increas-ing thickness of avian eggshells in proportion to egg masswith increasing egg size places an upper limit on the sizeof birds.[16][note 1] The largest species of Dromornis, D.stirtoni, may have gone extinct after it attained the max-imum avian body mass and was then outcompeted bymarsupial diprotodonts that evolved to sizes several timeslarger.[19]

2.3 Megafaunal mass extinctions

2.3.1 Timing and possible causes

A well-known mass extinction of megafauna, theHolocene extinction (see also Quaternary extinctionevent), occurred at the end of the last ice age glacial pe-riod (a.k.a. the Würm glaciation) and wiped out manygiant ice age animals, such as woolly mammoths, inthe Americas and northern Eurasia. Various theorieshave attributed the wave of extinctions to human hunt-ing, climate change, disease, a putative extraterrestrialimpact, or other causes. However, this extinction pulsenear the end of the Pleistocene was just one of a series

of megafaunal extinction pulses that have occurred dur-ing the last 50,000 years over much of the Earth’s surface,with Africa and southern Asia being largely spared. Thelatter areas did suffer a gradual attrition of megafauna,particularly of the slower-moving species (a class of vul-nerable megafauna epitomized by giant tortoises), overthe last several million years.[20][21]

Outside the mainland of Afro-Eurasia, these megafau-nal extinctions followed a highly distinctive landmass-by-landmass pattern that closely parallels the spread of hu-mans into previously uninhabited regions of the world,and which shows no correlation with climatic history(which can be visualized with plots over recent geolog-ical time periods of climate markers such as marine oxy-gen isotopes or atmospheric carbon dioxide levels).[22][23]Australia was struck first around 45,000 years ago,[24]followed by Tasmania about 41,000 years ago (afterformation of a land bridge to Australia about 43,000years ago),[25][26][27] Japan apparently about 30,000 yearsago,[28] North America 13,000 years ago, South Amer-ica about 500 years later,[29][30] Cyprus 10,000 yearsago,[31][32] the Antilles 6000 years ago,[33] New Caledo-nia[34] and nearby islands[35] 3000 years ago, Madagascar2000 years ago,[36] New Zealand 700 years ago,[37] theMascarenes 400 years ago,[38] and the Commander Is-lands 250 years ago.[39] Nearly all of the world’s iso-lated islands could furnish similar examples of extinc-tions occurring shortly after the arrival of Homo sapi-ens, though most of these islands, such as the HawaiianIslands, never had terrestrial megafauna, so their extinctfauna were smaller.[22][23]

An analysis of Sporormiella fungal spores (which de-rive mainly from the dung of megaherbivores) in swampsediment cores spanning the last 130,000 years fromLynch’s Crater in Queensland, Australia showed thatthe megafauna of that region virtually disappeared about41,000 years ago, at a time when climate changes wereminimal; the change was accompanied by an increasein charcoal, and was followed by a transition from rain-forest to fire-tolerant sclerophyll vegetation. The high-resolution chronology of the changes supports the hypoth-esis that human hunting alone eliminated the megafauna,and that the subsequent change in flora was most likelya consequence of the elimination of browsers and an in-crease in fire.[40][41][42] The increase in fire lagged the dis-appearance of megafauna by about a century, and mostlikely resulted from accumulation of fuel once brows-ing stopped. Over the next several centuries grass in-creased; sclerophyll vegetation increased with a lag ofanother century, and a sclerophyll forest developed af-ter about another thousand years.[42] During two periodsof climate change about 120 and 75 thousand years ago,sclerophyll vegetation had also increased at the site in re-sponse to a shift to cooler, drier conditions; neither ofthese episodes had a significant impact on megafaunalabundance.[42] Similar conclusions regarding the culpa-bility of human hunters in the disappearance of Pleis-

2.4. EXAMPLES 17

tocene megafauna were obtained via an analysis of a largecollection of eggshell fragments of the flightless Aus-tralian bird Genyornis newtoni[43] and from analysis ofSporormiella fungal spores from a lake in eastern NorthAmerica.[44][45]

Continuing human hunting and environmental distur-bance has led to additional megafaunal extinctions in therecent past, and has created a serious danger of furtherextinctions in the near future (see examples below).A number of other mass extinctions occurred earlier inEarth’s geologic history, in which some or all of themegafauna of the time also died out. Famously, in theCretaceous–Paleogene extinction event the dinosaurs andmost other giant reptilians were eliminated. However, theearlier mass extinctions were more global and not so se-lective 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 havebeen caused by more generalized types of disturbances tothe biosphere.

2.3.2 Consequences of depletion ofmegafauna

Effect on nutrient transport

Megafauna play a significant role in the lateral trans-port of mineral nutrients in an ecosystem, tending totranslocate them from areas of high to those of lowerabundance. They do so by their movement between thetime they consume the nutrient and the time they re-lease it through elimination (or, to a much lesser extent,through decomposition after death).[47] In South Amer-ica’s Amazon Basin, it is estimated that such lateral dif-fusion was reduced over 98% following the megafaunalextinctions that occurred roughly 12,500 years ago.[48][49]Given that phosphorus availability is thought to limit pro-ductivity in much of the region, the decrease in its trans-port from the western part of the basin and from flood-plains (both of which derive their supply from the upliftof the Andes) to other areas is thought to have signifi-cantly impacted the region’s ecology, and the effects maynot yet have reached their limits.[49]

Effect on methane emissions

Large populations of megaherbivores have the potentialto contribute greatly to the atmospheric concentration ofmethane, which is an important greenhouse gas. Mod-ern ruminant herbivores produce methane as a byprod-uct of foregut fermentation in digestion, and release itthrough belching. Today, around 20% of annual methaneemissions come from livestock methane release. In theMesozoic, it has been estimated that sauropods couldhave emitted 520 million tons of methane to the atmo-sphere annually,[50] contributing to the warmer climate

of the time (up to 10 C warmer than at present).[50][51]This large emission follows from the enormous estimatedbiomass of sauropods, and because methane productionof individual herbivores is believed to be almost propor-tional to their mass.[50]

Recent studies have indicated that the extinction ofmegafaunal herbivores may have caused a reductionin atmospheric methane. This hypothesis is relativelynew.[52] One study examined the methane emissions fromthe bison that occupied the Great Plains of North Amer-ica before contact with European settlers. The study es-timated that the removal of the bison caused a decreaseof as much as 2.2 million tons per year.[53] Another studyexamined the change in the methane concentration in theatmosphere at the end of the Pleistocene epoch after theextinction of megafauna in the Americas. After early hu-mans migrated to the Americas about 13,000 BP, theirhunting and other associated ecological impacts led tothe extinction of many megafaunal species there. Cal-culations suggest that this extinction decreased methaneproduction by about 9.6 million tons per year. This sug-gests that the absence of megafaunal methane emissionsmay have contributed to the abrupt climatic cooling at theonset of the Younger Dryas.[52] The decrease in atmo-spheric methane that occurred at that time, as recordedin ice cores, was 2-4 times more rapid than any other de-crease in the last half million years, suggesting that anunusual mechanism was at work.[52]

2.4 Examples

The following are some notable examples of animals of-ten considered as megafauna (in the sense of the “largeanimal” definition). This list is not intended to be ex-haustive:

• Clade Synapsida• Class Mammalia (phylogenetically, a cladewithin Therapsida; see below)• Infraclass Metatheria

• Order Diprotodontia• The red kangaroo (Macropus ru-

fus) is the largest living Australianmammal and marsupial at a weightof up to 85 kg (187 lb). How-ever, its extinct relative, the giantshort-faced kangaroo Procoptodongoliah reached 230 kg (510 lb),while extinct diprotodonts attainedthe largest size of any marsupial inhistory, up to an estimated 2,750 kg(6,060 lb). The extinct marsupiallion (Thylacleo carnifex), at up to160 kg (350 lb) was much largerthan any extant carnivorous marsu-pial.

18 CHAPTER 2. MEGAFAUNA

• Infraclass Eutheria• Superorder Afrotheria• Order Proboscidea• Elephants are the largest livingland animals. They and their rel-atives arose in Africa, but untilrecently had a nearly worldwidedistribution. The African bushelephant (Loxodonta africana) hasa shoulder height of up to 4.3m (14 ft) and weighs up to 13tons. Among recently extinctproboscideans, mammoths (Mam-muthus) were close relatives of ele-phants, while mastodons (Mam-mut) were much more distantlyrelated. The steppe mammoth(M. trogontherii) is estimated tohave commonly weighed around10 tonnes, making it possibly thelargest proboscid, which wouldmake it the second largest landmammal after indricotherines.

• Order Sirenia• The largest sirenian at up to 1500kg is the West Indian manatee(Trichechus manatus). Steller’ssea cow (Hydrodamalis gigas) wasprobably around five times as mas-sive, but was exterminated by hu-mans within 27 years of its dis-covery off the remote CommanderIslands in 1741. In prehistorictimes this sea cow also lived alongthe coasts of northeastern Asiaand northwestern North America;it was apparently eliminated fromthese more accessible locations byaboriginal hunters.

• Superorder Xenarthra• Order Cingulata• The glyptodonts were a group oflarge, heavily armored ankylosaur-like xenarthrans related to liv-ing armadillos. They originatedin South America, invaded NorthAmerica during the Great Amer-ican Interchange, and went ex-tinct at the end of the Pleistoceneepoch.[54]

• Order Pilosa• Ground sloths were another groupof slow, terrestrial xenarthrans,related to modern tree sloths.They had a similar history, al-though they reached North Amer-ica earlier, and spread farther

north (e.g., Megalonyx). Thelargest genera, Megatherium andEremotherium, reached sizes com-parable to elephants.[54]

• Superorder Euarchontoglires• Order Primates• The largest living primate, at upto 266 kg (586 lb), is the gorilla(Gorilla beringei and Gorilla go-rilla, with three of four sub-species being critically endan-gered). The extinct Malagasysloth lemur Archaeoindris reacheda similar size, while the ex-tinct Gigantopithecus blacki ofSoutheast Asia is believed to havebeen several times larger. Somepopulations of archaic Homo weresignificantly larger than recentHomo sapiens;[55][56] for example,Homo heidelbergensis in south-ern Africa may have commonlyreached 7 feet (2.1m) in height,[57]while Neanderthals were about30% more massive.[58]

• Order Rodentia• The extant capybara (Hydro-

choerus hydrochaeris) of SouthAmerica, the largest living rodent,weighs up to 65 kg (143 lb).Several recently extinct NorthAmerican forms were larger: thecapybara Neochoerus pinckneyi(another neotropic migrant) wasabout 40% heavier; the giantbeaver (Castoroides ohioensis) wassimilar. The extinct blunt-toothedgiant hutia (Amblyrhiza inundata)of several Caribbean islands mayhave been larger still. However,several million years ago SouthAmerica harbored much moremassive rodents. Phoberomys pat-tersoni, known from a nearly fullskeleton, probably reached 700 kg(1,500 lb). Fragmentary remainssuggest that Josephoartigasiamonesi grew to upwards of 1,000kg (2,200 lb).

• Superorder Laurasiatheria• Order Carnivora• Big cats include the tiger (Pan-

thera tigris) and lion (Pantheraleo). The largest subspecies, at upto 306 kg (675 lb), is the Siberiantiger (P. tigris altaica), in accordwith Bergmann’s rule. Members

2.4. EXAMPLES 19

of Panthera are distinguished bymorphological features which en-able them to roar. Larger ex-tinct felids include the Americanlion (Panthera leo atrox) and theSouth American saber-toothed catSmilodon populator.

• Bears are large carnivorans of thecaniform suborder. The largestliving forms are the polar bear (Ur-sus maritimus), with a body weightof up to 680 kg (1,500 lb), andthe similarly sized Kodiak bear(Ursus arctos middendorffi), againconsistent with Bergmann’s rule.Arctotherium augustans, an ex-tinct short-faced bear from SouthAmerica, was the largest predatoryland mammal ever with an esti-mated average weight of 1,600 kg(3,500 lb).[59]

• Seals, sea lions, and walrusesare amphibious marine carnivo-rans that evolved from bearlikeancestors. The southern ele-phant seal (Mirounga leonina) ofAntarctic and subantarctic watersis the largest carnivoran of all time,with bull males reaching a maxi-mum length of 6–7 m (20–23 ft)and maximum weight of 5,000 kg(11,000 lb).

• Order Perissodactyla• Tapirs are browsing animals, witha short prehensile snout and pig-like form that appears to havechanged little in 20 million years.They inhabit tropical forests ofSoutheast Asia and South andCentral America, and include thelargest surviving land animals ofthe latter two regions. There arefour species.

•Rhinoceros, from Dürer’s woodcut

Rhinoceroses are odd-toed ungu-lates with horns made of keratin,the same type of protein compos-ing hair. They are among thelargest living land mammals af-ter elephants (hippos attain a sim-ilar size). Three of five extantspecies are critically endangered.Their extinct central Asian rela-tives the indricotherines were thelargest terrestrial mammals of alltime.

• Order Artiodactyla (or cladistically,Cetartiodactyla)• Giraffes (Giraffa camelopardalis)are the tallest living land animals,reaching heights of up to nearly 6m (20 ft).

• Bovine ungulates include thelargest surviving land animals ofEurope and North America. Thewater buffalo (Bubalis arnee),bison (Bison bison and B. bona-sus), and gaur (Bos gaurus) can allgrow to weights of over 900 kg(2,000 lb).

• The semiaquatic hippopotamus(Hippopotamus amphibius) is theheaviest living even-toed ungulate;it and the critically endangeredpygmy hippo (Choeropsis liberien-sis) are believed to be the closestextant relatives of cetaceans.Hippos are among the megafau-nal species most dangerous tohumans.[60]

• Order Cetacea (or cladistically,Cetartiodactyla)• Whales, dolphins, and porpoisesare marine mammals. The bluewhale (Balaenoptera musculus) isthe largest baleen whale and thelargest animal that has ever lived,at 30 metres (98 ft)[61] in lengthand 170 tonnes (190 short tons)[62]or more in weight. The spermwhale (Physeter macrocephalus) isthe largest toothed whale, as wellas the planet’s loudest and brainiestanimal (with a brain about fivetimes as massive as a human’s).The killer whale (Orcinus orca) isthe largest dolphin.

• Order Pelycosauria (traditional; paraphyletic)• Cotylorhynchus was a large, big-clawed,herbivorous caseid of Early Permian

20 CHAPTER 2. MEGAFAUNA

North America, reaching 6 m (20 ft) and2 tonnes.

• Order Therapsida• Anteosaurus was a headbutting, semi-aquatic, carnivorous dinocephalian ofMiddle Permian South Africa. It reached5–6m (16–20 ft) long, and weighed about500–600 kg (1,100–1,300 lb).[63]

• Clade Sauropsida• Class Aves (phylogenetically, a clade withinCoelurosauria, a taxon within the orderSaurischia; see below)• Order Struthioniformes

• The ratites are an ancient anddiverse group of flightless birdsthat are found on fragments of theformer supercontinent Gondwana.The largest living bird, the ostrich(Struthio camelus) was surpassed bythe extinct Aepyornis of Madagascar,the heaviest of the group (400 kg(880 lb)), and the extinct giant moa(Dinornis) of New Zealand, thetallest, growing to heights of 3.4 m(11 ft). The latter two are examplesof island gigantism.

• Order Anseriformes• Extinct dromornithids of Australiasuch as Dromornis may have ex-ceeded the largest ratites in size.(Due to its small size for a continentand its isolation, Australia is some-times viewed as the world’s largest is-land; thus, these species could also beconsidered insular giants.)

• Class Reptilia (traditional; paraphyletic)• Order Crocodilia

• Alligators and crocodiles are largesemiaquatic reptiles, the largestof which, the saltwater crocodile(Crocodylus porosus), can grow toa weight of 1,360 kg (3,000 lb).Crocodilians’ distant ancestors andtheir kin, the crurotarsans, domi-nated the world in the late Triassic,until the Triassic–Jurassic extinctionevent allowed dinosaurs to over-take them. They remained diverseduring the later Mesozoic, whencrocodyliforms such as Deinosuchusand Sarcosuchus reached lengthsof 12 m. Similarly large crocodil-ians, such as Mourasuchus andPurussaurus, were present as re-cently as the Miocene in SouthAmerica.

• Order Saurischia• Saurischian dinosaurs of the Jurassicand Cretaceous include sauropods,the longest (at up to 40 m or 130 ft)and most massive terrestrial animalsknown (Argentinosaurus reached 80–100 metric tonnes, or 90–110 tons),as well as theropods, the largest ter-restrial carnivores (Spinosaurus grewto 7–9 tonnes; the more famousTyrannosaurus, to 6.8 tonnes).

• Order Squamata• While the largest extant lizard, theKomodo dragon (Varanus komod-oensis), another island giant, canreach 3 m (10 ft) in length, its extinctAustralian relative Megalania mayhave reached more than twice thatsize. These monitor lizards' marinerelatives, the mosasaurs, were apexpredators in late Cretaceous seas.

• The heaviest extant snake is consid-ered to be the green anaconda (Eu-nectes murinus), while the reticulatedpython (Python reticulatus), at upto 8.7 m or more, is consideredthe longest. An extinct AustralianPliocene species of Liasis, the BluffDowns giant python, reached 10 m,while the Paleocene Titanoboa ofSouth America reached lengths of12–15 m and an estimated weight ofabout 1135 kilograms (2500 lb).

• Order Testudines• The largest turtle is the critically en-dangered marine leatherback turtle(Dermochelys coriacea), weighingup to 900 kg (2,000 lb). It is dis-tinguished from other sea turtles byits lack of a bony shell. The mostmassive terrestrial chelonians arethe giant tortoises of the GalápagosIslands (Chelonoidis nigra) andAldabra Atoll (Aldabrachelys gigan-tea), at up to 300 kg (660 lb). Thesetortoises are the biggest survivorsof an assortment of giant tortoisespecies that were widely presenton continental landmasses[64][65]and additional islands[64] during thePleistocene.

• Class Amphibia (in the wide, probably paraphyletic,sense)

• Order Temnospondyli (relationship to extantamphibians is unclear)

2.5. GALLERY 21

• The Permian temnospondylPrionosuchus, the largest amphibianknown, reached 9 m in length andwas an aquatic predator resembling acrocodilian. After the appearance ofreal crocodilians, temnospondyls such asKoolasuchus (5 m long) had retreated tothe Antarctic region by the Cretaceous,before going extinct.

• Class Actinopterygii

• Order Tetraodontiformes• The largest extant bony fish is the oceansunfish (Mola mola), whose average adultweight is 1,000 kg (2,200 lb). While phy-logenetically a “bony fish”, its skeleton isprimarily cartilage (which is lighter thanbone). It has a disk-shaped body, and pro-pels itself with its long, thin dorsal andanal fins; it feeds primarily on jellyfish.In these three respects (as well as in itssize and diving habits), it resembles aleatherback turtle.

• Order Acipenseriformes• The critically endangered beluga (Eu-ropean sturgeon, Huso huso) at up to1,476 kg (3,254 lb) is the largest sturgeon(which are also mostly cartilaginous) andis considered the largest anadromous fish.

• Order Siluriformes• The critically endangered Mekong giantcatfish (Pangasianodon gigas), at up to293 kg (646 lb), is often viewed as thelargest freshwater fish.

• Class Chondrichthyes

• Order Lamniformes• The largest living predatory fish, the greatwhite shark (Carcharodon carcharias),reaches weights up to 2,240 kg (4,940lb). Its extinct relative C. megalodon (thedisputed genus being either Carcharodonor Carcharocles) was more than an orderof magnitude larger, and is the largestpredatory shark or fish of all time (andpossibly the largest predator in vertebratehistory); it preyed on whales and othermarine mammals.

• Order Orectolobiformes• The largest extant shark, cartilaginousfish, and fish overall is the whale shark(Rhincodon typus), which reaches weightsin excess of 21.5 tonnes (47,000 pounds).Like baleen whales, it is a filter feeder andprimarily consumes plankton.

• Order Rajiformes• Themanta ray (Manta birostris) is anotherfilter feeder and the largest ray, growingto up to 2300 kg.

• Class Placodermi

• Order Arthrodira• The largest armored fish, Dunkleosteus,arose during the late Devonian. At upto 10 metres (33 ft) in length[66] and 3.6tonnes (4.0 short tons) in mass,[67] it was ahypercarnivorous apex predator that em-ployed suction feeding.[68][69] Its contem-porary, Titanichthys, apparently an earlyfilter feeder, rivaled it in size. The an-throdires were eliminated by the environ-mental upheavals of the Late Devonianextinction, after existing for only about 50million years.

• Class Cephalopoda

• Order Teuthida• A number of deep ocean creatures ex-hibit abyssal gigantism. These includethe giant squid (Architeuthis) and colossalsquid (Mesonychoteuthis hamiltoni); both(although rarely seen) are believed to at-tain lengths of 12 m (39 ft) or more. Thelatter is the world’s 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 di-verse group of aquatic and possibly am-phibious predators that included the mostmassive arthropods to have existed. Theysurvived over 200 million years, but fi-nally died out in the Permian–Triassic ex-tinction event along with trilobites andmost other forms of life present at thetime, including most of the dominant ter-restrial therapsids. The Early DevonianJaekelopterus reached an estimated lengthof 2.5 m (8.2 ft), not including itsraptorial chelicerae, and is thought tohave been a freshwater species.

2.5 Gallery

2.5.1 Extinct

• Some Paleozoic sea scorpions (Eurypterus shown)were larger than a man.

22 CHAPTER 2. MEGAFAUNA

• Dunkleosteus was a 10 m (33 ft) long toothlessarmored predatory Devonian placoderm fish.

• Sail-backed pelycosaur Dimetrodon andtemnospondyl Eryops from North America’sPermian.

• Pliosaur Pliosaurus (right) harassing the filter feederfish Leedsichthys during the Jurassic.

• Macronarian sauropods; from left, Camarasaurus,Brachiosaurus, Giraffatitan, Euhelopus.

• Tyrannosaurus was a 12.3 m (40 ft) long theropoddinosaur, an apex predator of west North America.

• Indricotheres, the land mammals closest tosauropods in size and lifestyle, were Asian rhinos.

• The Late Miocene teratorn Argentavis of SouthAmerica had a 7 m (23 ft) wingspan.

• C. megalodon (two possible sizes) with a whaleshark, great white shark and human for scale.

• Deinotherium had downward-curving tusks andranged widely over Afro-Eurasia.

• Titanis walleri, the only terror bird known to haveinvaded North America, was 2.5 m (8 ft 2 in) tall.

• Hippo-sized Diprotodon of Australia, the largestmarsupial of all time, went extinct 40,000 years ago.

• Megalania, a giant carnivorous goanna of Australia,might have grown to 7 metres long.

• Elephant-sizedMegatherium, from South America’sPleistocene, was the largest sloth.[1]

• Toxodon, one of South America’s largest and lastnotoungulates.[1] It had a relative in Mexico.

• American lions exceeded extant lions in size andranged over two continents until 10,000 BP.

• Woolly mammoths vanished after humans invadedtheir habitat in Eurasia and N. America.[2]

• Haast’s eagle, the largest eagle known, attackingmoa (which included the tallest bird known).

1. ^ Cite error: The named referenceFari.C3.B1a2013 was invoked but never defined(see the help page).

2. ^ Cite error: The named reference Stuart was in-voked but never defined (see the help page).

2.5.2 Living

• The gorilla is the largest and one of the mostendangered primates on the planet.

• Siberian tigers are the biggest living cats, exempli-fying Bergmann’s rule.

• Polar bears, the largest bears and semiaquatic carni-vores, are vulnerable to global warming.

• The critically endangered black rhinoceros, up to 14feet (4.3 m) long, is threatened by poaching.

• Wild Bactrian camels are critically endangered.Their ancestors originated in North America.

• Unlike woolly rhinos and mammoths, muskoxennarrowly survived the Quaternary extinctions.[1]

• Hippos, the heaviest and most aquatic even-toed un-gulates, are whales' closest living relatives.

• A filter feeder up to 33 m (108 ft) long, the bluewhale is the largest animal of all time.

• The orca, the largest dolphin and pack predator, ishighly intelligent and lives in complex societies.

• The ostrich is the largest ratite, the heaviest liv-ing bird and, at 70 km/h,[2] the fastest runningbiped.[3][note 1]

• The saltwater crocodile is the largest living reptileand a dangerous predator of humans.

• The Komodo dragon, an insular giant, is the largestlizard and has infectious and venomous saliva.

• The green anaconda, an aquatic constrictor, is theheaviest snake, weighing up to 97.5 kg (215 lb).

• The deep-diving ocean sunfish is the largest bonyfish, but its skeleton is mostly cartilaginous.

• The Nile perch, one of the largest freshwater fish, isalso a damaging invasive species.[note 2]

• The whale shark is the largest extant shark or fishspecies, growing up to 12.6 m (41 ft) in length.

• The manta, a filter feeder, is the largest ray at up to7.6 m across, yet can breach clear of the water.

• Examination of a 9 m giant squid, an abyssal giantand the second largest cephalopod.

1. ^ Cite error: The named reference Stuart was in-voked but never defined (see the help page).

2. ^ Davies, S.J.J.F. (2003). “Birds I Tinamous andRatites to Hoatzins”. In Hutchins, Michael. Grz-imek’s Animal Life Encyclopedia 8 (2 ed.). Farm-ington Hills, MI: Gale Group. pp. 99–101. ISBN0-7876-5784-0.

2.8. REFERENCES 23

3. ^ Cite error: The named reference Stewart2006 wasinvoked but never defined (see the help page).

4. ^ Penny, M. (2002). The Secret World of Kanga-roos. Austin TX: Raintree Steck-Vaughn. ISBN 0-7398-4986-7.

Cite error: There are <ref group=note> tags on thispage, but the references will not show without a {{re-flist|group=note}} template (see the help page).

2.6 See also

• Australian megafauna

• Bergmann’s rule

• Charismatic megafauna

• Cope’s rule

• Deep-sea gigantism

• Fauna

• Giant animals in fiction and mythology

• Island dwarfism

• 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 be-

cause they had a different relationship between body massand egg size than birds. The 400 kg Aepyornis had largereggs than nearly all dinosaurs.[17][18]

2.8 References[1] Stuart, A. J. (November 1991). “Mammalian extinctions

in the Late Pleistocene of northern Eurasia and NorthAmerica”. Biological Reviews (Wiley) 66 (4): 453–562.doi:10.1111/j.1469-185X.1991.tb01149.x.

[2] Johnson, C. N. (2002-09-23). “Determinants of Loss ofMammal Species during the Late Quaternary 'Megafauna'Extinctions: Life History and Ecology, but Not BodySize”. Proceedings of the Royal Society of London B (TheRoyal Society) 269 (1506): 2221–2227 (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 VertebratePaleontology 2. New York: Kluwer/Plenum. pp. 17–56.ISBN 978-0-306-46092-0. OCLC 41368299. Retrieved2011-08-23.

[4] Ice Age Animals. Illinois State Museum

[5] Evans, A. R.; et al. (2012-01-30). “The max-imum rate of mammal evolution”. PNAS 109.doi:10.1073/pnas.1120774109. Retrieved 2011-02-11.

[6] Smith, F. A.; Boyer, A. G.; Brown, J. H.; Costa, D. P.;Dayan, T.; Ernest, S. K. M.; Evans, A. R.; Fortelius,M.; Gittleman, J. L.; Hamilton, M. J.; Harding, L.E.; Lintulaakso, K.; Lyons, S. K.; McCain, C.; Okie,J. G.; Saarinen, J. J.; Sibly, R. M.; Stephens, P. R.;Theodor, J.; Uhen, M. D. (2010-11-26). “The EvolutionofMaximumBody Size of Terrestrial Mammals”. Science330 (6008): 1216–1219. Bibcode:2010Sci...330.1216S.doi:10.1126/science.1194830. Retrieved 2012-01-07.

[7] Clauss, M.; Frey, R.; Kiefer, B.; Lechner-Doll, M.;Loehlein, W.; Polster, C.; Roessner, G. E.; Streich, W.J. (2003-04-24). “The maximum attainable body size ofherbivorous mammals: morphophysiological constraintson foregut, and adaptations of hindgut fermenters”.Oecologia 136 (1): 14–27. doi:10.1007/s00442-003-1254-z. PMID 12712314. Retrieved 2012-01-08.

[8] Sorkin, B. (2008-04-10). “A biomechanical con-straint on body mass in terrestrial mammalian preda-tors”. Lethaia 41 (4): 333–347. doi:10.1111/j.1502-3931.2007.00091.x. Retrieved 2011-08-02.

[9] Carbone, C.; Teacher, A; Rowcliffe, J. M. (2007-01-16).“The Costs of Carnivory”. PLoS Biology 5 (2, e22): 363–368. doi:10.1371/journal.pbio.0050022. PMC 1769424.PMID 17227145. Retrieved 2012-01-08.

[10] Ashton, K. G.; Tracy, M. C.; de Queiroz, A. (Oc-tober 2000). “Is Bergmann’s Rule Valid for Mam-mals?". The American Naturalist 156 (4): 390–415.doi:10.1086/303400. Retrieved 2012-01-07.

[11] Thewissen, J. G. M.; Bajpai, S. (1 January 2001).“Whale Origins as a Poster Child for Macroevolution”.BioScience 51 (12): 1037–1049. doi:10.1641/0006-3568(2001)051[1037:WOAAPC]2.0.CO;2. ISSN 0006-3568.

24 CHAPTER 2. MEGAFAUNA

[12] Mitchell, K. J.; Llamas, B.; Soubrier, J.; Rawlence,N. J.; Worthy, T. H.; Wood, J.; Lee, M. S. Y.;Cooper, A. (2014-05-23). “Ancient DNA revealselephant birds and kiwi are sister taxa and clarifiesratite bird evolution”. Science 344 (6186): 898–900.doi:10.1126/science.1251981. PMID 24855267.

[13] Phillips MJ, Gibb GC, Crimp EA, Penny D (January2010). “Tinamous and moa flock together: mitochondrialgenome sequence analysis reveals independent losses offlight among ratites”. Systematic Biology 59 (1): 90–107.doi:10.1093/sysbio/syp079. PMID 20525622.

[14] Baker, A. J.; Haddrath, O.; McPherson, J. D.; Cloutier,A. (2014). “Genomic Support for a Moa-TinamouClade and Adaptive Morphological Convergence inFlightless Ratites”. Molecular Biology and Evolution.doi:10.1093/molbev/msu153.

[15] Murray, Peter F.; Vickers-Rich, Patricia (2004).Magnificent Mihirungs: The Colossal Flightless Birds ofthe Australian Dreamtime. Indiana University Press. pp.51, 314. ISBN 978-0-253-34282-9. Retrieved 7 January2012.

[16] Ibid. p. 212.

[17] Kenneth Carpenter (1999). Eggs, Nests, and Baby Di-nosuars: A Look at Dinosaur Reproduction. IndianaUniversity Press. ISBN 978-0-253-33497-8. OCLC42009424. Retrieved 6 May 2013.

[18] Jackson, F. D.; Varricchio, D. J.; Jackson, R. A.; Vila, B.;Chiappe, L. M. (2008). “Comparison of water vapor con-ductance in a titanosaur egg from the Upper Cretaceous ofArgentina and a Megaloolithus siruguei egg from Spain”.Paleobiology 34 (2): 229–246. doi:10.1666/0094-8373(2008)034[0229:COWVCI]2.0.CO;2. ISSN 0094-8373.

[19] Ibid. p. 277.

[20] Corlett, R. T. (2006). “Megafaunal extinctions in tropicalAsia”. Tropinet 17 (3): 1–3. Retrieved 2010-10-04.

[21] Edmeades, Baz. “Megafauna — First Victims of theHuman-Caused Extinction”. (internet-published bookwith Foreword by Paul S. Martin). Retrieved 2010-10-04.

[22] Martin, P. S. (2005). “Chapter 6. Deadly Syncopation”.Twilight of the Mammoths: Ice Age Extinctions and theRewilding of America. University of California Press. pp.118–128. ISBN 0-520-23141-4. OCLC 58055404. Re-trieved 2014-11-11.

[23] Burney, D. A.; Flannery, T. F. (July 2005). “Fifty mil-lennia of catastrophic extinctions after human contact”.Trends in Ecology & Evolution (Elsevier) 20 (7): 395–401. doi:10.1016/j.tree.2005.04.022. PMID 16701402.Retrieved 2014-11-11.

[24] Roberts, R. G.; Flannery, T. F.; Ayliffe, L. K.; Yoshida,H.; Olley, J. M.; Prideaux, G. J.; Laslett, G. M.; Baynes,A.; Smith, M. A.; Jones, R.; Smith, B. L. (2001-06-08).“NewAges for the Last AustralianMegafauna: Continent-Wide Extinction About 46,000 Years Ago”. Science

292 (5523): 1888–1892. Bibcode:2001Sci...292.1888R.doi:10.1126/science.1060264. PMID 11397939. Re-trieved 2011-08-26.

[25] Diamond, Jared (2008-08-13). “Palaeontology: Thelast giant kangaroo”. Nature 454 (7206): 835–836.Bibcode:2008Natur.454..835D. doi:10.1038/454835a.PMID 18704074. Retrieved 2011-05-08.

[26] Turney, C. S. M.; Flannery, T. F.; Roberts, R. G.;et al. (2008-08-21). “Late-surviving megafaunain Tasmania, Australia, implicate human involve-ment in their extinction”. PNAS (NAS) 105 (34):12150–12153. Bibcode:2008PNAS..10512150T.doi:10.1073/pnas.0801360105. PMC 2527880. PMID18719103. Retrieved 2011-05-08.

[27] Roberts, R.; Jacobs, Z. (October 2008). “The Lost Gi-ants of Tasmania”. Australasian Science 29 (9): 14–17.Retrieved 2011-08-26.

[28] Norton, C. J.; Kondo, Y.; Ono, A.; Zhang, Y.;Diab, M. C. (2009-05-23). “The nature ofmegafaunal extinctions during the MIS 3–2 tran-sition in Japan”. Quaternary International 211(1–2): 113–122. Bibcode:2010QuInt.211..113N.doi:10.1016/j.quaint.2009.05.002. Retrieved 2011-08-30.

[29] Haynes, Gary (2009). “Introduction to the Volume”.In Haynes, Gary. American Megafaunal Extinctions atthe End of the Pleistocene. Springer. pp. 1–20.doi:10.1007/978-1-4020-8793-6_1. ISBN 978-1-4020-8792-9.

[30] Fiedel, Stuart (2009). “Sudden Deaths: The Chronol-ogy of Terminal Pleistocene Megafaunal Extinction”.In Haynes, Gary. American Megafaunal Extinctions atthe End of the Pleistocene. Springer. pp. 21–37.doi:10.1007/978-1-4020-8793-6_2. ISBN 978-1-4020-8792-9.

[31] Simmons, A. H. (1999). Faunal extinction in anisland society: pygmy hippopotamus hunters ofCyprus. Interdisciplinary Contributions to Archae-ology. Kluwer Academic/Plenum Publishers. p. 382.doi:10.1007/b109876. ISBN 978-0-306-46088-3.OCLC 41712246.

[32] Simmons, A. H.; Mandel, R. D. (December 2007).“Not Such a New Light: A Response to Ammermanand Noller”. World Archaeology 39 (4): 475–482.doi:10.1080/00438240701676169. JSTOR 40026143.

[33] Steadman, D. W.; Martin, P. S.; MacPhee, R. D.E.; Jull, A. J. T.; McDonald, H. G.; Woods, C.A.; Iturralde-Vinent, M.; Hodgins, G. W. L. (2005-08-16). “Asynchronous extinction of late Quater-nary sloths on continents and islands”. Proc. Natl.Acad. Sci. USA (National Academy of Sciences) 102(33): 11763–11768. Bibcode:2005PNAS..10211763S.doi:10.1073/pnas.0502777102. PMC 1187974. PMID16085711.

[34] Anderson, A.; Sand, C.; Petchey, F.; Worthy, T. H.(2010). “Faunal extinction and human habitation in New

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Caledonia: Initial results and implications of new researchat the Pindai Caves”. Journal of Pacific Archaeology 1 (1):89–109. hdl:10289/5404.

[35] White, A. W.; Worthy, T. H.; Hawkins, S.; Bed-ford, S.; Spriggs, M. (2010-08-16). “Megafaunalmeiolaniid horned turtles survived until earlyhuman settlement in Vanuatu, Southwest Pa-cific”. Proc. Natl. Acad. Sci. USA 107 (35):15512–15516. Bibcode:2010PNAS..10715512W.doi:10.1073/pnas.1005780107. PMC 2932593. PMID20713711.

[36] Burney, D. A.; Burney, L. P.; Godfrey, L. R.; Jungers,W. L.; Goodman, S. M.; Wright, H. T.; Jull. A. J. T.(July 2004). “A chronology for late prehistoric Mada-gascar”. Journal of Human Evolution 47 (1–2): 25–63. doi:10.1016/j.jhevol.2004.05.005. PMID 15288523.Retrieved 2011-08-30.

[37] Holdaway, R. N.; Jacomb, C. (2000-03-24). “RapidExtinction of the Moas (Aves: Dinornithiformes):Model, Test, and Implications”. Science 287(5461): 2250–2254. Bibcode:2000Sci...287.2250H.doi:10.1126/science.287.5461.2250. PMID 10731144.Retrieved 2011-08-30.

[38] Janoo, A. (April 2005). “Discovery of isolated dodobones (Raphus cucullatus (L.), Aves, Columbiformes)from Mauritius cave shelters highlights human predation,with a comment on the status of the family RaphidaeWet-more, 1930”. Annales de Paléontologie 91 (2): 167–180.doi:10.1016/j.annpal.2004.12.002. Retrieved 2011-08-30.

[39] Anderson, P. K. (July 1995). “Competition, Pre-dation, and the Evolution and Extinction of Steller’sSea Cow, Hydrodamalis gigas". Marine Mammal Sci-ence (Society for Marine Mammalogy) 11 (3): 391–394. doi:10.1111/j.1748-7692.1995.tb00294.x. Re-trieved 2011-08-30.

[40] Biello, D. (2012-03-22). “Big Kill, Not Big Chill, Fin-ished Off Giant Kangaroos”. Scientific American news.Retrieved 2012-03-25.

[41] McGlone, M. (2012-03-23). “The HuntersDid It”. Science 335 (6075): 1452–1453. Bibcode:2012Sci...335.1452M.doi:10.1126/science.1220176. Retrieved 2012-03-25.

[42] Rule, S.; Brook, B. W.; Haberle, S. G.; Turney,C. S. M.; Kershaw, A. P. (2012-03-23). “The Af-termath of Megafaunal Extinction: Ecosystem Trans-formation in Pleistocene Australia”. Science 335(6075): 1483–1486. Bibcode:2012Sci...335.1483R.doi:10.1126/science.1214261. Retrieved 2012-03-25.

[43] Miller, G. H.; Magee, J. W.; Johnson, B. J.; Fogel, M. L.;Spooner, N. A.; McCulloch, M. T.; Ayliffe, L. K. (1999-01-08). “Pleistocene Extinction of Genyornis newtoni:Human Impact on Australian Megafauna”. Science 283(5399): 205–208. doi:10.1126/science.283.5399.205.PMID 9880249.

[44] Johnson, C. (2009-11-20). “Megafaunal De-cline and Fall”. Science 326 (5956): 1072–1073.doi:10.1126/science.1182770. PMID 19965418.

[45] Gill, J. L.; Williams, J.W.; Jackson, S. T.; Lininger, K. B.;Robinson, G. S. (2009-11-20). “Pleistocene MegafaunalCollapse, Novel Plant Communities, and Enhanced FireRegimes in North America”. Science 326 (5956): 1100–1103. doi:10.1126/science.1179504. PMID 19965426.

[46] Alroy, J. (2008-08-12). “Dynamics of orig-ination and extinction in the marine fossilrecord”. PNAS. 105 Suppl 1 (Supplement_1):11536–11542. Bibcode:2008PNAS..10511536A.doi:10.1073/pnas.0802597105. PMC 2556405. PMID18695240.

[47] Wolf, A.; Doughty, C. E.; Malhi, Y. (2013-08-09). “Lat-eral Diffusion of Nutrients by Mammalian Herbivoresin Terrestrial Ecosystems”. PLoS ONE 8 (8): e71352.doi:10.1371/journal.pone.0071352.

[48] Marshall, M. (2013-08-11). “Ecosystems still feel thepain of ancient extinctions”. New Scientist. Retrieved2013-08-12.

[49] Doughty, C. E.; Wolf, A.; Malhi, Y. (2013-08-11).“The legacy of the Pleistocene megafauna extinctions onnutrient availability in Amazonia”. Nature Geoscience.doi:10.1038/ngeo1895.

[50] Wilkinson, D. M.; Nisbet, E. G.; Ruxton, G. D.(2012-05-08). “Could methane produced by sauro-pod dinosaurs have helped drive Mesozoic climatewarmth?". Current Biology 22 (9): R292–R293.doi:10.1016/j.cub.2012.03.042. Retrieved 2012-05-08.

[51] “Dinosaur gases 'warmed the Earth'". BBC Nature News.2012-05-07. Retrieved 2012-05-08.

[52] Smith, F. A.; Elliot, S. M.; Lyons, S. K. (2010-05-23).“Methane emissions from extinct megafauna”. NatureGeoscience (Nature Publishing Group) 3 (6): 374–375.Bibcode:2010NatGe...3..374S. doi:10.1038/ngeo877.Retrieved 2011-02-26.

[53] Kelliher, F. M.; Clark, H. (2010-03-15). “Methaneemissions from bison—An historic herd esti-mate for the North American Great Plains”.Agricultural and Forest Meteorology 150 (3): 473–577. doi:10.1016/j.agrformet.2009.11.019.

[54] Fariña, Richard A.; Vizcaíno, Sergio F.; De Iuliis, Gerry(22 May 2013). Megafauna: Giant Beasts of PleistoceneSouth America. Indiana University Press. ISBN 0-253-00719-4. OCLC 779244424.

[55] Ruff, C. B.; Trinkaus, E.; Holliday, T. W. (1997-05-08). “Body mass and encephalization in Pleis-tocene Homo”. Nature 387 (6629): 173–176.Bibcode:1997Natur.387..173R. doi:10.1038/387173a0.PMID 9144286. Retrieved 2012-05-25.

[56] Grine, F. E.; Jumgers, W. L.; Tobias, P. V.; Pearson, O.M. (June 1995). “Fossil Homo femur from Berg Aukas,northern Namibia”. American Journal of Physical Anthro-pology 97 (2): 151–185. doi:10.1002/ajpa.1330970207.PMID 7653506.

26 CHAPTER 2. MEGAFAUNA

[57] Smith, Chris; Burger, Lee (November 2007). “Our Story:Human Ancestor Fossils”. The Naked Scientists. Re-trieved 2011-02-19.

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[59] Soibelzon, L. H.; Schubert, B. W. (January 2011). “TheLargest Known Bear, Arctotherium angustidens, from theEarly Pleistocene Pampean Region of Argentina: With aDiscussion of Size and Diet Trends in Bears”. Journalof Paleontology (Paleontological Society) 85 (1): 69–75.doi:10.1666/10-037.1. Retrieved 2011-06-01.

[60] Swift, E. M. (1997-11-17). “What Big Mouths TheyHave: Travelers in Africa who run afoul of hippos maynot live to tell the tale”. Sports Illustrated Vault. Time Inc.Retrieved 2011-11-16.

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[64] Hansen, D. M.; Donlan, C. J.; Griffiths, C. J.; Campbell,K. J. (April 2010). “Ecological history and latent con-servation potential: large and giant tortoises as a modelfor taxon substitutions”. Ecography (Wiley) 33 (2): 272–284. doi:10.1111/j.1600-0587.2010.06305.x. Retrieved2011-02-26.

[65] Cione, A. L.; Tonni, E. P.; Soibelzon, L. (2003). “TheBroken Zig-Zag: Late Cenozoic large mammal and tor-toise extinction in South America”. Rev. Mus. ArgentinoCienc. Nat., n.s. 5 (1): 1–19. ISSN 1514-5158. Retrieved2011-02-06.

[66] Palmer, D. (1 July 2002). The Marshall Illustrated Ency-clopedia of Dinosaurs and Prehistoric Animals. New LineBooks. ISBN 978-1-57717-293-2. OCLC 183092423.Retrieved 2013-06-10.

[67] Monster fish crushed opposition with strongest bite ever.The Sydney Morning Herald. November 30, 2006.

[68] Anderson, P. S.L; Westneat, M. W (2006-11-28). “Feed-ing mechanics and bite force modelling of the skull ofDunkleosteus terrelli, an ancient apex predator”. BiologyLetters 3 (1): 77–80. doi:10.1098/rsbl.2006.0569. ISSN1744-9561.

[69] Anderson, P.S.L. (2010-05-04). “Using linkage modelsto explore skull kinematic diversity and functional conver-gence in arthrodire placoderms”. Journal of Morphology:990–1005. doi:10.1002/jmor.10850. ISSN 0362-2525.

[70] Stewart, D. (2006-08-01). “A Bird Like No Other”.National Wildlife. National Wildlife Federation.Archived from the original on 2012-02-09. Retrieved2014-05-30.

2.9 External links• Megafauna – “First Victims of the Human-CausedExtinction”

Chapter 3

Pleistocene

Earth during the Pleistocene epoch.

The Pleistocene /ˈplaɪstɵsiːn/ (symbol PS[1]) is the ge-ological epoch which lasted from about 2,588,000 to11,700 years ago, spanning the world’s recent period ofrepeated glaciations.Charles Lyell introduced this term in 1839 to describestrata in Sicily that had at least 70% of their mol-luscan fauna still living today. This distinguished itfrom the older Pliocene Epoch, which Lyell had origi-nally thought to be the youngest fossil rock layer. Heconstructed the name “Pleistocene” (“Most New” or“Newest”) from the Greek πλεῖστος, pleīstos, “most”,and καινός, kainós (latinized as cænus), “new";[2] thiscontrasting with the immediately preceding Pleiocene(“More New” or “Newer”, from πλείων, pleíōn, “more”,and kainós; usual spelling: Pliocene), and the imme-diately subsequent Holocene (“wholly new” or “entirelynew”, from ὅλος, hólos, “whole”, and kainós) epoch,which extends to the present time.The Pleistocene is the first epoch of the QuaternaryPeriod or sixth epoch of the Cenozoic Era.[3] The end ofthe Pleistocene corresponds with the end of the last glacialperiod. It also corresponds with the end of the Paleolithicage used in archaeology. In the ICS timescale, the Pleis-tocene is divided into four stages or ages, the Gelasian,Calabrian, Ionian and Tarantian. All of these stages weredefined in southern Europe. In addition to this interna-tional subdivision, various regional subdivisions are oftenused.Before a change finally confirmed in 2009 by theInternational Union of Geological Sciences, the timeboundary between the Pleistocene and the precedingPliocene was regarded as being at 1.806 million years

before the present, as opposed to the currently accepted2.588 million years BP: publications from the precedingyears may use either definition of the period.

3.1 Dating

The Pleistocene has been dated from 2.588 million(±.005) to 11,700 years before present (BP), with the enddate expressed in radiocarbon years as 10,000 carbon-14years BP.[4] It covers most of the latest period of repeatedglaciation, up to and including the Younger Dryas coldspell. The end of the Younger Dryas has been dated toabout 9640 BC (11,654 calendar years BP). It was notuntil after the development of radiocarbon dating, how-ever, that Pleistocene archaeological excavations shiftedto stratified caves and rock-shelters as opposed to open-air river-terrace sites.[5]

In 2009 the International Union of Geological Sciences(IUGS) confirmed a change in time period for the Pleis-tocene, changing the start date from 1.806 to 2.588 mil-lion years BP, and accepted the base of the Gelasian asthe base of the Pleistocene, namely the base of the MonteSan Nicola GSSP.[6] The IUGS has yet to approve a typesection, Global Boundary Stratotype Section and Point(GSSP), for the upper Pleistocene/Holocene boundary(i.e. the upper boundary). The proposed section is theNorth Greenland Ice Core Project ice core 75° 06' N 42°18' W.[7] The lower boundary of the Pleistocene Seriesis formally defined magnetostratigraphically as the baseof the Matuyama (C2r) chronozone, isotopic stage 103.Above this point there are notable extinctions of the cal-careous nanofossils: Discoaster pentaradiatus and Dis-coaster surculus.[8][9]

The Pleistocene covers the recent period of repeatedglaciations. The name Plio-Pleistocene has in the pastbeen used to mean the last ice age. The revised defini-tion of the Quaternary, by pushing back the start date ofthe Pleistocene to 2.58 Ma, results in the inclusion of allthe recent repeated glaciations within the Pleistocene.

27

28 CHAPTER 3. PLEISTOCENE

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

3.2 Paleogeography and climate

The modern continents were essentially at their presentpositions during the Pleistocene, the plates upon whichthey sit probably having moved no more than 100 km rel-ative to each other since the beginning of the period.According to Mark Lynas (through collected data), thePleistocene’s overall climate could be characterized asa continuous El Niño with trade winds in the southPacific weakening or heading east, warm air rising nearPeru, warm water spreading from the west Pacific andthe Indian Ocean to the east Pacific, and other El Niñomarkers.[10]

3.2.1 Glacial features

Pleistocene climate wasmarked by repeated glacial cyclesin which continental glaciers pushed to the 40th parallelin some places. It is estimated that, at maximum glacialextent, 30% of the Earth’s surface was covered by ice. Inaddition, a zone of permafrost stretched southward fromthe edge of the glacial sheet, a few hundred kilometresin North America, and several hundred in Eurasia. The

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 incontinental ice sheets 1,500 to 3,000 metres (4,900–9,800 ft) thick, resulting in temporary sea-level drops of100 metres (300 ft) or more over the entire surface ofthe Earth. During interglacial times, such as at present,drowned coastlines were common, mitigated by isostaticor other emergent motion of some regions.The effects of glaciation were global. Antarctica was ice-bound throughout the Pleistocene as well as the precedingPliocene. The Andes were covered in the south by thePatagonian ice cap. There were glaciers in New Zealandand Tasmania. The current decaying glaciers of MountKenya, Mount Kilimanjaro, and the Ruwenzori Range ineast and central Africa were larger. Glaciers existed inthe mountains of Ethiopia and to the west in the Atlasmountains.In the northern hemisphere, many glaciers fused into one.The Cordilleran ice sheet covered the North Americannorthwest; the east was covered by the Laurentide. TheFenno-Scandian ice sheet rested on northern Europe, in-cluding Great Britain; the Alpine ice sheet on the Alps.Scattered domes stretched across Siberia and the Arcticshelf. The northern seas were ice-covered.South of the ice sheets large lakes accumulated becauseoutlets were blocked and the cooler air slowed evapora-tion. When the Laurentide ice sheet retreated, north cen-tral North America was totally covered by Lake Agassiz.Over a hundred basins, now dry or nearly so, were over-flowing in the North American west. Lake Bonneville,for example, stood where Great Salt Lake now does. InEurasia, large lakes developed as a result of the runofffrom the glaciers. Rivers were larger, had a more copiousflow, and were braided. African lakes were fuller, appar-ently from decreased evaporation. Deserts on the otherhand were drier and more extensive. Rainfall was lowerbecause of the decrease in oceanic and other evaporation.

3.2.2 Major events

Further information: Timeline of glaciationOver 11 major glacial events have been identified, aswell as many minor glacial events.[11] A major glacialevent is a general glacial excursion, termed a “glacial.”Glacials are separated by “interglacials”. During a glacial,the glacier experiences minor advances and retreats. Theminor excursion is a “stadial"; times between stadials are“interstadials”.These events are defined differently in different regionsof the glacial range, which have their own glacial historydepending on latitude, terrain and climate. There is a gen-eral correspondence between glacials in different regions.Investigators often interchange the names if the glacialgeology of a region is in the process of being defined.

3.2. PALEOGEOGRAPHY AND CLIMATE 29

Ice ages as reflected in atmospheric CO2, stored in bubbles fromglacial ice of Antarctica.

However, it is generally incorrect to apply the name of aglacial in one region to another.For most of the 20th century only a few regions had beenstudied and the names were relatively few. Today the ge-ologists of different nations are taking more of an interestin Pleistocene glaciology. As a consequence, the num-ber of names is expanding rapidly and will continue toexpand. Many of the advances and stadials remain un-named. Also, the terrestrial evidence for some of themhas been erased or obscured by larger ones, but evidenceremains from the study of cyclical climate changes.The glacials in the following tables show historical us-ages, are a simplification of a much more complex cycleof variation in climate and terrain, and are generally nolonger used. These names have been abandoned in favorof numeric data because many of the correlations werefound to be either inexact or incorrect and more than fourmajor glacials have been recognized since the historicalterminology was established.[11][12][13]

Corresponding to the terms glacial and interglacial, theterms pluvial and interpluvial are in use (Latin: pluvia,rain). A pluvial is a warmer period of increased rainfall;an interpluvial, of decreased rainfall. Formerly a pluvialwas thought to correspond to a glacial in regions not iced,and in some cases it does. Rainfall is cyclical also. Plu-vials and interpluvials are widespread.There is no systematic correspondence of pluvials toglacials, however. Moreover, regional pluvials do not cor-respond to each other globally. For example, some haveused the term “Riss pluvial” in Egyptian contexts. Anycoincidence is an accident of regional factors. Only a fewof the names for pluvials in restricted regions have beenstrategraphically defined.

3.2.3 Palaeocycles

The sum of transient factors acting at the Earth’s surfaceis cyclical: climate, ocean currents and other movements,wind currents, temperature, etc. The waveform response

comes from the underlying cyclical motions of the planet,which eventually drag all the transients into harmony withthem. The repeated glaciations of the Pleistocene werecaused by the same factors.

Milankovitch cycles

Main article: Milankovitch cycles

Glaciation in the Pleistocene was a series of glacials andinterglacials, stadials and interstadials, mirroring periodicchanges in climate. The main factor at work in climatecycling is now believed to be Milankovitch cycles. Theseare periodic variations in regional and planetary solar ra-diation reaching the Earth caused by several repeatingchanges in the Earth’s motion.Milankovitch cycles cannot be the sole factor responsi-ble for the variations in climate since they explain neitherthe long term cooling trend over the Plio-Pleistocene, northe millennial variations in the Greenland Ice Cores. Mi-lankovitch pacing seems to best explain glaciation eventswith periodicity of 100,000, 40,000, and 20,000 years.Such a pattern seems to fit the information on climatechange found in oxygen isotope cores. The timing ofour present interglacial interval (known as the Holocene,Postglacial, or the Present Interglacial) to that of the pre-vious interglacial, beginning about 130,000 years ago(The Eemian Interglacial), suggests that the next glacialwould likely begin in about 3,000 years.

Oxygen isotope ratio cycles

Main article: Oxygen isotope ratio cycle

In oxygen isotope ratio analysis, variations in the ratio of18O to 16O (two isotopes of oxygen) by mass (measured by a massspectrometer) present in the calcite of oceanic core sam-ples is used as a diagnostic of ancient ocean temperaturechange and therefore of climate change. Cold oceans arericher in 18O, which is included in the tests of the microorganisms(foraminifera) contributing the calcite.A more recent version of the sampling process makes useof modern glacial ice cores. Although less rich in 18O than sea water, the snow that fell on the glacier year byyear nevertheless contained 18O and 16O in a ratio that depended on the mean annual tempera-ture.Temperature and climate change are cyclical when plot-ted on a graph of temperature versus time. Temperaturecoordinates are given in the form of a deviation from to-day’s 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 difference from theratio found in standard mean ocean water (SMOW).The graph in either form appears as a waveform withovertones. 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 MISstages beginning at about 2.588Ma BP in the Early Pleis-tocene Gelasian. Early Pleistocene stages were shallowand frequent. The latest were the most intense and mostwidely spaced.By convention, stages are numbered from the Holocene,which is MIS1. Glacials receive an even number; inter-glacials, odd. The first major glacial was MIS2-4 at about85–11 ka BP. The largest glacials were 2, 6, 12, and 16;the warmest interglacials, 1, 5, 9 and 11. For matchingof MIS numbers to named stages, see under the articlesfor those names.

3.3 Fauna

See also: Quaternary extinction event

Both marine and continental faunas were essentially mod-ern and many animals, specifically, mammals were muchlarger in body form than their modern relatives .

Pleistocene of Northern Spain showing woolly mammoth, cavelions eating a reindeer, tarpans, and woolly rhinoceros.

The severe climatic changes during the ice age had ma-jor impacts on the fauna and flora. With each advance ofthe ice, large areas of the continents became totally de-populated, and plants and animals retreating southwardin front of the advancing glacier faced tremendous stress.The most severe stress resulted from drastic climaticchanges, reduced living space, and curtailed food supply.A major extinction event of large mammals (megafauna),which included mammoths, mastodons, saber-toothedcats, glyptodons, ground sloths, Irish elk, cave bears, andshort-faced bears, began late in the Pleistocene and con-tinued into the Holocene. Neanderthals also became ex-tinct 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

Pleistocene of South America showing Megatherium and twoGlyptodon.

had replaced the megafauna and migrated north.The extinctions were especially severe in North Americawhere native horses and camels were eliminated.

• Asian land mammal ages (ALMA) includeZhoukoudianian, Nihewanian, and Yushean.

• European land mammal ages (ELMA) includeGelasian (2.5—1.8 Ma).

• North American land mammal ages (NALMA) in-clude Blancan (4.75–1.8), Irvingtonian (1.8–0.24)and Rancholabrean (0.24–0.01) in millions of years.The Blancan extends significantly back into thePliocene.

• South American land mammal ages (SALMA) in-clude Uquian (2.5–1.5), Ensenadan (1.5–0.3) andLujanian (0.3–0.01) in millions of years. TheUquian previously extended significantly back intothe Pliocene, although the new definition places itentirely within the Pleistocene.

3.4 Humans

Main articles: Human evolution, Paleolithic and Modelsof migration to the New World

Scientific evidence[14] indicates that humans evolved intotheir present form during the Pleistocene.[15] In the be-ginning of the Pleistocene Paranthropus species are stillpresent, as well as early human ancestors, but during thelower Palaeolithic they disappeared, and the only hominidspecies found in fossilic records isHomo erectus for muchof the Pleistocene. The Middle and late Palaeolithic sawthe appearance of new types of humans, as well as thedevelopment of more elaborate tools than found in previ-ous eras. According to mitochondrial timing techniques,modern humans migrated from Africa after the Rissglaciation in the middle Palaeolithic during the Eemian

3.7. REFERENCES 31

Stage, spreading all over the ice-free world during the latePleistocene.[16][17][18] A 2005 study posits that humansin this migration interbred with archaic human forms al-ready outside of Africa by the late Pleistocene, incorpo-rating archaic human genetic material into the modernhuman gene pool.[19]

3.5 Deposits

Pleistocene non-marine sediments are found primarilyin fluvial deposits, lakebeds, slope and loess deposits aswell as in the large amounts of material moved about byglaciers. Less common are cave deposits, travertines andvolcanic deposits (lavas, ashes). Pleistocene marine de-posits are found primarily in shallowmarine basinsmostly(but with important exceptions) in areas within a few tensof kilometers of the modern shoreline. In a few geolog-ically active areas such as the Southern California coast,Pleistocene marine deposits may be found at elevationsof 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 Pleis-tocene and Holocene Epochs” Chapter 22 PDF (3.1 MB)InGradstein, F. M., Ogg, James G., and Smith, A. Gilbert(eds.), A Geologic Time Scale 2004 Cambridge UniversityPress, 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) “TheNeogene Period”. In: Gradstein, F., Ogg, J., Smith, A.G.(Eds.), A Geologic Time Scale 2004. Cambridge: Cam-bridge University Press.

[5] Moore, Mark; Brumm (2007). “Stone artifacts and ho-minins in island Southeast Asia: New insights from Flo-res, eastern Indonesia”. Journal of Human Evolution 52:88. Retrieved 10 April 2014.

[6] Riccardi, Alberto C. (30 June 2009) “IUGS ratified ICSRecommendation on redefinition of Pleistocene and for-mal definition of base of Quaternary” International Unionof Geological Sciences

[7] Svensson, A., S. W. Nielsen, S. Kipfstuhl, S. J. Johnsen,J. P. Steffensen, M. Bigler, U. Ruth, and R. Röthlisberger(2005) “Visual stratigraphy of the North Greenland IceCore Project (NorthGRIP) ice core during the last glacialperiod” Journal of Geophysical Research 110: (D02108)

[8] Gradstein, Felix M.; Ogg, James G. and Smith, A. Gilbert(eds.) (2005) A Geologic Time Scale 2004 CambridgeUniversity Press, Cambridge, UK, p. 28, ISBN 0-521-78142-6

[9] Rio, D.; Sprovieri, R.; Castradori, D. and Di Stefano, E.(1998) “The Gelasian Stage (Upper Pliocene): a new unitof the global standard chronostratigraphic scale” Episodes21: pp. 82-87

[10] National Geographic Channel, Six Degrees Could ChangeThe World, Mark Lynas interview. Retrieved February14, 2008.

[11] Richmond, G.M. and D.S. Fullerton, 1986, Summationof Quaternary glaciations in the United States of America.Quaternary Science Reviews. vol. 5, pp. 183-196.

[12] Roy, M., P.U. Clark, R.W. Barendregt, J.R., Glasmann,and R.J. Enkin, 2004, Glacial stratigraphy and paleomag-netism of late Cenozoic deposits of the north-central UnitedStates, PDF version, 1.2MB. Geological Society of Amer-ica 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 theNebraskan, Aftonian, Kansan, and Yarmouthian stageswere abandoned by modern stratigraphers).

[14] Rogers, A.R. and Jorde, L.B. (1995) “Genetic evidenceon modern human origins” Human Biology 67: pp. 1–36

[15] Wall, J.D. and Przeworski, M. (2000) “When did thehuman population start increasing?" Genetics 155: pp.1865–1874

[16] Cann, R.L.; Stoneking, M. and Wilson, A.C.(1987) “Mi-tochondrial DNA and human evolution” Nature 325: pp.31–36

[17] Stringer, C.B. (1992) “Evolution of early modern hu-mans” In: Jones, Steve; Martin, R. and Pilbeam, DavidR. (eds.) (1992) The Cambridge encyclopedia of humanevolution Cambridge University Press, Cambridge, ISBN0-521-32370-3, pp. 241–251.

[18] Templeton, A. (2002) “Out of Africa again and again”Na-ture 416: p. 45

[19] Eswarana, Vinayak; Harpendingb, Henry and Rogers,Alan R. (2005) “Genomics refutes an exclusively Africanorigin of humans” Journal of Human Evolution 49(1): pp.1–18 Abstract

• Ogg, Jim; June, 2004, Overview of GlobalBoundary Stratotype Sections and Points (GSSP’s,Stratigraphy.org, Accessed April 30, 2006.

32 CHAPTER 3. PLEISTOCENE

3.8 External links• The SMU-in-Taos Research Publications digitalcollection of anthropological and archaeologicalmonographs contains Late Pleistocene environ-ments of the southern high plains.

• Pleistocene Microfossils: 50+ images ofForaminifera

Chapter 4

Prehistoric mammal

An early drawing depicting prehistoric mammals

Prehistoric mammals are groups of mammals that be-came extinct before humans developed writing. 164million years ago, in the Jurassic period, Castorocaudalutrasimilis, a mammaliaform (mammal-shaped) animalweighing about 500 grams (1.1 lb), had a full mammalianpelt, with guard hairs and underfur, webbed feet, andscales on the tail like a modern beaver, as well as teethspecialized for catching fish.Later, about 130 million years ago in the Cretaceous,there existed larger mammals; a fossil of Repenomamusgiganticus indicates that the animal was about 1 me-ter (3¼ ft) long. In the stomach of a smaller cousin,Repenomamus robustus at 52 cm (20½ in), the remainsof a juvenile dinosaur have been preserved.The lineages of many varieties continued through theCenozoic era where some reached very large sizes. Mostof the very large mammals became extinct in the last iceage, but have smaller descendants.

4.1 List of prehistoric mammals

Main article: List of prehistoric mammals

Prehistoric mammals include:

4.2 See also• Cynodont

• List of extinct mammals

• Mammaliaformes

• Megamammals

• Pleistocene extinctions

• Pleistocene megafauna

• Synapsid

• Therapsid

33

Chapter 5

Stone Age

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

Modern Awash River, Ethiopia, descendant of the Palaeo-Awash, source of the sediments in which the oldest Stone Agetools have been found

which stone was widely used to make implements witha sharp edge, a point, or a percussion surface. The periodlasted roughly 3.4 million years, and ended between 6000BCE and 2000 BCE with the advent of metalworking.[1]Stone Age artifacts include tools used by humans and bytheir predecessor species in the genusHomo, as well as theearlier partly contemporaneous genera Australopithecusand Paranthropus. Bone tools were used during this pe-riod as well but are rarely preserved in the archaeologicalrecord. The Stone Age is further subdivided by the typesof stone tools in use.The Stone Age is the first of the three-age systemof archaeology, which divides human technologicalprehistory into three periods:

• The Stone Age

• The Bronze Age

• The Iron Age

5.1 Historical significance

The Stone Age is contemporaneous with the evolution ofthe genus Homo, the only exception possibly being at thevery beginning, when species prior to Homo may havemanufactured tools. According to the age and locationof the current evidence, the cradle of the genus is theEast African Rift System, especially toward the north inEthiopia, where it is bordered by grasslands. The closestrelative 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 conduitfor movement into southern Africa and also north downthe Nile into North Africa and through the continuationof the rift in the Levant to the vast grasslands of Asia.Starting from about 3 million years ago (mya) a singlebiome established itself from South Africa through therift, North Africa, and across Asia to modern China,which has been called “transcontinental 'savannahstan'"recently.[2] Starting in the grasslands of the rift, Homoerectus, the predecessor of modern humans, found anecological niche as a tool-maker and developed a de-pendence on it, becoming a “tool equipped savannadweller.”[3]

5.2 The Stone Age in archaeology

5.2.1 Beginning of the Stone Age

During 2010, fossilised animal bones bearing marks fromstone tools were found in the Lower Awash Valley inEthiopia. Discovered by an international team led byShannon McPherron, at 3.4 million years old they are theoldest evidence of stone tool use ever found anywhere inthe world.[1]

The oldest known stone tools have been excavated fromseveral sites at Gona, Ethiopia, on the sediments of thepaleo-Awash River, which serve to date them. All thetools come from the Busidama Formation, which liesabove a disconformity, or missing layer, which wouldhave been from 2.9 to 2.7 mya. The oldest sites contain-ing tools are dated to 2.6–2.55 mya.[4] One of the most

34

5.2. THE STONE AGE IN ARCHAEOLOGY 35

Obsidian projectile point

striking circumstances about these sites is that they arefrom the Late Pliocene, where previous to their discoverytools were thought to have evolved only in the Pleistocene.Rogers and Semaw, excavators at the locality, point outthat:[5]

"...the earliest stone tool makers were skilledflintknappers .... The possible reasons behindthis seeming abrupt transition from the absenceof stone tools to the presence thereof include ...gaps in the geological record.”

The excavators are confident that more tools will befound elsewhere from 2.9 mya. The species whomade the Pliocene tools remains unknown. Fragmentsof Australopithecus garhi, Australopithecus aethiopicus[6]and Homo, possibly Homo habilis, have been found insites near the age of the oldest tools.[7]

5.2.2 End of the Stone Age

Innovation of the technique of smelting ore ended theStone Age and began the Bronze Age. The first mostsignificant metal manufactured was bronze, an alloy ofcopper and tin, each of which was smelted separately.The transition from the Stone Age to the Bronze Age wasa period during which modern people could smelt cop-per, but did not yet manufacture bronze, a time knownas the Copper Age, or more technically the Chalcolithic,“copper-stone” age. The Chalcolithic by convention is

the initial period of the Bronze Age and is unquestionablypart of the Age of Metals. The Bronze Age was followedby the Iron Age. During this entire time stone remainedin use in parallel with the metals for some objects, includ-ing those also used in the Neolithic, such as stone pottery.The transition out of the Stone Age occurred between6000 BCE and 2500 BCE for much of humanity living inNorth Africa and Eurasia. The first evidence of humanmetallurgy dates to between the 5th and 6th millenniumBCE in the archaeological sites of Majdanpek, Yarmovacand Pločnik (a copper axe from 5500 BCE belonging tothe Vinca culture), though not conventionally consideredpart of the Chalcolithic or “Copper Age”, this providesthe earliest known example of copper metallurgy.[8] andthe Rudna Glava mine in Serbia. Ötzi the Iceman, amummy from about 3300 BCE carried with him a copperaxe and a flint knife.In regions such as Subsaharan Africa, the Stone Age wasfollowed directly by the Iron Age. The Middle East andsoutheastern Asian regions progressed past Stone Agetechnology around 6000 BCE. Europe, and the rest ofAsia became post–Stone Age societies by about 4000BCE. The proto-Inca cultures of South America contin-ued at a Stone Age level until around 2000 BCE, whengold, copper and silver made their entrance, the rest fol-lowing later. Australia remained in the Stone Age untilthe 17th century. Stone tool manufacture continued. InEurope and North America, millstones were in use untilwell into the 20th century, and still are in many parts ofthe world.

5.2.3 The concept of Stone Age

The terms was never meant to suggest that advancementand time periods in prehistory are only measured by thetype of tool material, rather than, for example, social or-ganization, food sources exploited, adaptation to climate,adoption of agriculture, cooking, settlement and religion.Like pottery, the typology of the stone tools combinedwith the relative sequence of the types in various regionsprovide a chronological framework for the evolution ofman and society. They serve as diagnostics of date, ratherthan characterizing the people or the society.Lithic analysis is a major and specialised form of ar-chaeological investigation. It involves the measurementof the stone tools to determine their typology, functionand the technology involved. It includes scientific studyof the lithic reduction of the raw materials, examininghow the artifacts were made. Much of this study takesplace in the laboratory in the presence of various special-ists. In experimental archaeology, researchers attempt tocreate replica tools, to understand how they were made.Flintknappers are craftsmen who use sharp tools to re-duce flintstone to flint tool.In addition to lithic analysis, the field prehistorian uti-lizes a wide range of techniques derived from multiple

36 CHAPTER 5. STONE AGE

A variety of stone tools

fields. The work of the archaeologist in determining thepaleocontext and relative sequence of the layers is supple-mented by the efforts of the geologic specialist in identi-fying layers of rock over geologic time, of the paleonto-logical specialist in identifying bones and animals, of thepalynologist in discovering and identifying plant species,of the physicist and chemist in laboratories determiningdates by the carbon-14, potassium-argon and other meth-ods. Study of the Stone Age has never been mainly aboutstone tools and archaeology, which are only one form ofevidence. The chief focus has always been on the societyand the physical people who belonged to it.Useful as it has been, the concept of the Stone Age has itslimitations. The date range of this period is ambiguous,disputed, and variable according to the region in ques-tion. While it is possible to speak of a general 'stoneage' period for the whole of humanity, some groups neverdeveloped metal-smelting technology, so remained in a'stone age' until they encountered technologically devel-oped cultures. The term was innovated to describe thearchaeological cultures of Europe. It may not always bethe best in relation to regions such as some parts of theIndies and Oceania, where farmers or hunter-gatherersused stone for tools until European colonisation began.The archaeologists of the late 19th and early 20th cen-turies CE, who adapted the three-age system to theirideas, hoped to combine cultural anthropology and ar-chaeology in such a way that a specific contemporaneoustribe can be used to illustrate the way of life and beliefsof the people exercising a specific Stone-Age technology.As a description of people living today, the term stone ageis controversial. The Association of Social Anthropolo-gists discourages this use, asserting:[9]

“To describe any living group as 'primitive'or 'Stone Age' inevitably implies that they areliving representatives of some earlier stage ofhuman development that the majority of hu-mankind has left behind. For some, this couldbe a positive description, implying, for exam-ple, that such groups live in greater harmonywith nature .... For others, ... 'primitive' is

a negative characterisation. For them, 'primi-tive' denotes irrational use of resources and ab-sence of the intellectual and moral standards of'civilised' human societies.... From the stand-point of anthropological knowledge, both theseviews are equally one-sided and simplistic.”

5.2.4 The three-stage system

In the 1920s, South African archaeologists organizing thestone tool collections of that country observed that theydid not fit the newly detailed Three-Age System. In thewords of J. Desmond Clark,[10]

“It was early realized that the threefold di-vision of culture into Stone, Bronze and IronAges adopted in the nineteenth century for Eu-rope had no validity in Africa outside the Nilevalley.”

Consequently they proposed a new system for Africa, theThree-stage System. Clark regarded the Three-age Sys-tem as valid for North Africa; in sub-Saharan Africa, theThree-stage System was best.[11] In practice, the failureof African archaeologists either to keep this distinctionin mind, or to explain which one they mean, contributesto the considerable equivocation already present in the lit-erature. There are in effect two Stone Ages, one part ofthe Three-age and the other constituting the Three-stage.They refer to one and the same artifacts and the sametechnologies, but vary by locality and time.The Three-stage System was proposed in 1929 by AstleyJohn Hilary Goodwin, a professional archaeologist, andClarence van Riet Lowe, a civil engineer and amateur ar-chaeologist, in an article titled “Stone Age Cultures ofSouth Africa” in the journal Annals of the South AfricanMuseum. By then, the dates of the Early Stone Age, orPaleolithic, and Late StoneAge, or Neolithic (neo = new),were fairly solid and were regarded by Goodwin as abso-lute. He therefore proposed a relative chronology of peri-ods with floating dates, to be called the Earlier and LaterStone Age. The Middle Stone Age would not change itsname, but it would not mean Mesolithic.[12]

The duo thus reinvented the Stone Age. In Sub-SaharanAfrica, however, it was ended by the intrusion of the IronAge from the north. The Neolithic and the Bronze Agenever occurred. Moreover, the technologies included inthose 'stages’, as Goodwin called them, were not exactlythe same. Since then, the original relative terms have be-come identified with the technologies of the Paleolithicand Mesolithic, so that they are no longer relative. More-over, there has been a tendency to drop the comparativedegree in favor of the positive: resulting in two sets ofEarly, Middle and Late Stone Ages of quite different con-tent and chronologies.

5.3. CHRONOLOGY 37

By voluntary agreement, archaeologists respect the deci-sions of the Pan-African Congress of Prehistory, whichmeets every four years to resolve archaeological businessbrought before it. Delegates are actually international; theorganization takes its name from the topic. Louis Leakeyhosted the first one in Nairobi in 1947. It adopted Good-win and Lowe’s 3-stage system at that time, the stages tobe called Early, Middle and Later.

5.2.5 The problem of the transitions

The problem of the transitions in archaeology is a branchof the general philosophic continuity problem, whichexamines how discrete objects of any sort that arecontiguous in any way can be presumed to have a rela-tionship of any sort. In archaeology, the relationship isone of causality. If Period B can be presumed to descendfrom Period A, there must be a boundary between A andB, the A–B boundary. The problem is in the nature ofthis boundary. If there is no distinct boundary, then thepopulation of A suddenly stopped using the customs char-acteristic of A and suddenly started using those of B, anunlikely scenario in the process of evolution. More real-istically, a distinct border period, the A/B transition, ex-isted, in which the customs of A were gradually droppedand those of B acquired. If transitions do not exist, thenthere is no proof of any continuity between A and B.The Stone Age of Europe is characteristically in deficitof known transitions. The 19th and early 20th-centuryinnovators of the modern three-age system recognizedthe problem of the initial transition, the “gap” betweenthe Paleolithic and the Neolithic. Louis Leakey providedsomething of an answer by proving that man evolved inAfrica. The Stone Age must have begun there to be car-ried repeatedly to Europe by migrant populations. Thedifferent phases of the Stone Age thus could appear therewithout transitions. The burden on African archaeolo-gists became all the greater, because now they must findthe missing transitions in Africa. The problem is difficultand ongoing.After its adoption by the First Pan African Congress in1947, the Three-Stage Chronology was amended by theThird Congress in 1955 to include a First Intermedi-ate Period between Early and Middle, to encompass theFauresmith and Sangoan technologies, and the Second In-termediate Period between Middle and Later, to encom-pass the Magosian technology and others. The chrono-logic basis for definition was entirely relative. With thearrival of scientific means of finding an absolute chronol-ogy, the two intermediates turned out to be will-of-the-wisps. 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 artificial mix of two different periods.”[14]

Once seriously questioned, the intermediates did not waitfor the next Pan African Congress two years hence,

but were officially rejected in 1965 (again on an advi-sory basis) by Burg Wartenstein Conference #29, Sys-tematic Investigation of the African Later Tertiary andQuaternary,[15] a conference in anthropology held by theWenner-Gren Foundation, at Burg Wartenstein Castle,which it then owned in Austria, attended by the samescholars that attended the Pan African Congress, includ-ing Louis Leakey and Mary Leakey, who was deliveringa pilot presentation of her typological analysis of EarlyStone Age tools, to be included in her 1971 contributionto Olduvai Gorge, “Excavations in Beds I and II, 1960–1963.”[16]

However, although the intermediate periods were gone,the search for the transitions continued.

5.3 Chronology

2

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-8

2

2.5

3

3.5

4

4.5

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18Equ

ival

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(°C

)100 kyr cycle41 kyr cycle

Five Million Years ofClimate Change

From Sediment Cores

Millions of Years Ago

Time series plot of temperature over the previous 5 million years

In 1859 Jens Jacob Worsaae first proposed a division ofthe Stone Age into older and younger parts based on hiswork with Danish kitchenmiddens that began in 1851.[17]In the subsequent decades this simple distinction devel-oped into the archaeological periods of today. The majorsubdivisions of the Three-age Stone Age cross two epochboundaries on the geologic time scale:

• The geologic Pliocene–Pleistocene boundary(highly glaciated climate)

• The Paleolithic period of archaeology

• The geologic Pleistocene–Holocene boundary(modern climate)

• Mesolithic or Epipaleolithic period of archae-ology

• Neolithic period of archaeology

The succession of these phases varies enormously fromone region (and culture) to another.

5.3.1 Three-age chronology

Main articles: Paleolithic, Human evolution and Three-age system

38 CHAPTER 5. STONE AGE

The Paleolithic or Palaeolithic (from Greek: παλαιός,palaios, “old"; and λίθος, lithos, “stone” lit. “old stone,”coined by archaeologist John Lubbock and published in1865) is the earliest division of the Stone Age. It cov-ers the greatest portion of humanity’s 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 thefirst documented use of stone tools by hominans such asHomo habilis, to the end of the Pleistocene around 10,000BCE.[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 (about2,500,000 to 200,000 years ago), simple pebble toolshave been found in association with the remains of whatmay have been the earliest human ancestors. A somewhatmore sophisticated Lower Paleolithic tradition, known asthe Chopper chopping-tool industry, is widely distributedin the Eastern Hemisphere. This tradition is thoughtto have been the work of the hominin species namedHomo erectus. Although no such fossil tools have yetbeen found, it is believed that H. erectus probably madetools of wood and bone as well as stone. About 700,000years ago, a new Lower Paleolithic tool, the hand ax, ap-peared. The earliest European hand axes are assignedto the Abbevillian industry, which developed in northernFrance in the valley of the Somme River; a later, more re-fined hand-ax tradition is seen in the Acheulian industry,evidence of which has been found in Europe, Africa, theMiddle East, and Asia. Some of the earliest known handaxes were found at Olduvai Gorge (Tanzania) in associ-ation with remains of H. erectus. Alongside the hand-ax tradition there developed a distinct and very differ-ent stone-tool industry, based on flakes of stone: specialtools were made from worked (carefully shaped) flakes offlint. In Europe, the Clactonian industry is one exampleof a flake tradition. The early flake industries probablycontributed to the development of the Middle Paleolithicflake tools of theMousterian industry, which is associatedwith the remains of Neanderthal man.[19]

Oldowan in Africa Main article: Oldowan

The earliest documented stone tools were found in EastAfrica, manufacturers unknown. They belonged to anindustry now known as Oldowan, after the type site ofOlduvai Gorge in Tanzania; however, sites in Ethiopialater proved to be older.The tools were formed by knocking pieces off a river peb-ble, or stones like it, with a hammerstone to obtain largeand small pieces with one or more sharp edges. The origi-

This is a Mode 1, or Oldowan, stone tool from the western Sa-hara.

nal stone is called a core; the resultant pieces, flakes. Typ-ically, but not necessarily, small pieces are detached froma larger piece, in which case the larger piecemay be calledthe core and the smaller pieces the flakes. The prevalentusage, however, is to call all the results flakes, which canbe confusing. A split in half is called bipolar flaking.Consequently the method is often called “core-and-flake”. More recently, the tradition has been called “smallflake” since the flakes were small compared to subsequentAcheulean tools.[20]

“The essence of the Oldowan is the makingand often immediate use of small flakes.”

Another naming scheme is “Pebble Core Technology(PBC)":[21]

“Pebble cores are ... artifacts that havebeen shaped by varying amounts of hard-hammer percussion.”

Various refinements in the shape have been called chop-pers, discoids, polyhedrons, subspheroid, etc. To date noreasons for the variants have been ascertained:[22]

“From a functional standpoint, pebblecores seem designed for no specific purpose.”

However, they would not have been manufactured for nopurpose:[22]

“Pebble cores can be useful in many cut-ting, scraping or chopping tasks, but ... theyare not particularly more efficient in such tasksthan a sharp-edged rock ....”

The whole point of their utility is that each is a “sharp-edged rock” in locations where nature has not providedany. There is additional evidence that Oldowan, or Mode

5.3. CHRONOLOGY 39

1, tools were utilized in “percussion technology"; that is,they were designed to be gripped at the blunt end andstrike something with the edge, from which use they weregiven the name of choppers. Modern science has beenable to detect mammalian blood cells on Mode 1 toolsat Sterkfontein, Member 5 East, in South Africa. As theblood must have come from a fresh kill, the tool usersare likely to have done the killing and used the tools forbutchering. Plant residues bonded to the silicon of sometools confirm the use to chop plants.[23]

Although the exact species authoring the tools remainsunknown, Mode 1 tools in Africa were manufactured andused predominantly byHomo habilis. They cannot be saidto have developed these tools or to have contributed thetradition to technology. They continued a tradition of yetunknown origin. As chimpanzees sometimes naturallyuse percussion to extract or prepare food in the wild, andmay use either unmodified stones or stones that they havesplit, creating an Oldowan tool, the tradition may well befar older than its current record.Towards the end of Oldowan in Africa a new species ap-peared over the range of Homo habilis: Homo erectus.The earliest “unambiguous” evidence is a whole cranium,KNM-ER 3733 (a find identifier) from Koobi Fora inKenya, dated to 1.78 mya.[24] An early skull fragment,KNM-ER 2598, dated to 1.9 mya, is considered a goodcandidate also.[25] Transitions in paleoanthropology arealways hard to find, if not impossible, but based on the“long-legged” limb morphology shared by H. habilis andH. rudolfensis in East Africa, an evolution from one ofthose two has been suggested.[26]

The most immediate cause of the new adjustments ap-pears to have been an increasing aridity in the region andconsequent contraction of parkland savanna, interspersedwith trees and groves, in favor of open grassland, dated1.8–1.7 mya. During that transitional period the percent-age of grazers among the fossil species increased from15–25% to 45%, dispersing the food supply and requir-ing a facility among the hunters to travel longer distancescomfortably, which H. erectus obviously had.[27] The ul-timate proof is the “dispersal” of H. erectus “across muchof Africa and Asia, substantially before the developmentof theMode 2 technology and use of fire ....”[26] H. erectuscarried Mode 1 tools over Eurasia.According to the current evidence (which may change atany time) Mode 1 tools are documented from about 2.6mya to about 1.5 mya in Africa,[28] and to 0.5 mya outsideof it.[29] The genus Homo is known from H. habilis andH. rudolfensis from 2.3 to 2.0 mya, with the latest habilisbeing an upper jaw from Koobi Fora, Kenya, from 1.4mya. H. erectus is dated 1.8–0.6 mya.[30]

According to this chronology Mode 1 was inher-ited by Homo from unknown Hominans, probablyAustralopithecus and Paranthropus, who must have con-tinued on with Mode 1 and then with Mode 2 until theirextinction no later than 1.1 mya. Meanwhile living con-

temporaneously in the same regions H. habilis inheritedthe tools around 2.3 mya. At about 1.9 mya H. erectuscame on stage and lived contemporaneously with the oth-ers. Mode 1 was now being shared by a number of Hom-inans over the same ranges, presumably subsisting in dif-ferent niches, but the archaeology is not precise enoughto say which.

Oldowan out of Africa Tools of the Oldowan tra-dition first came to archaeological attention in Europe,where, being intrusive and not well defined, comparedto the Acheulean, they were puzzling to archaeologists.The mystery would be elucidated by African archaeol-ogy 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 theUnited States, used the term to describe a “chalky boulderclay” underlying a layer of gravel at Hoxne, central Eng-land, where Acheulean tools had been found.[31] Whetherany tools would be found in it and what type was notknown. Hugo Obermaier, a contemporary German ar-chaeologist working in Spain, quipped:

“Unfortunately, the stage of human indus-try which corresponds to these deposits cannotbe positively identified. All we can say is thatit is pre-Acheulean....”

This uncertainty was clarified by the subsequent excava-tions at Olduvai; nevertheless, the term is still in use forpre-Acheulean contexts, mainly across Eurasia, that areyet unspecified or uncertain but with the understandingthat they are or will turn out to be pebble-tool.[32]

There are ample associations ofMode 2 withH. erectus inEurasia. H. erectus –Mode 1 associations are scantier butthey do exist, especially in the Far East. One strong pieceof evidence prevents the conclusion that only H. erectusreached Eurasia: at Yiron, Israel, Mode 1 tools have beenfound dating to 2.4 mya,[33] about 0.5 my earlier than theknown H. erectus finds. If the date is correct, either an-other Hominan preceded H. erectus out of Africa or theearliest H. erectus has yet to be found.After the initial appearance at Gona in Ethiopia at 2.7mya, 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 alreadyleft pebble tools at Yiron, Israel, at 2.4 mya, Riwat, Pak-istan, at 2.0 mya, and Renzidong, South China, at over 2mya.[34] The identification of a fossil skull at Mojokerta,Pernung Peninsula on Java, dated to 1.8 mya, as H. erec-tus, suggests that the African finds are not the earliest tobe found in Africa, or that, in fact, erectus did not origi-nate in Africa after all but on the plains of Asia.[26] Theoutcome of the issue waits for more substantial evidence.Erectus was found also at Dmanisi, Georgia, from 1.75mya in association with pebble tools.

40 CHAPTER 5. STONE AGE

Pebble tools are found the latest first in southern Europeand then in northern. They begin in the open areas of Italyand Spain, the earliest dated to 1.6 mya at Pirro Nord,Italy. The mountains of Italy are rising at a rapid ratein the framework of geologic time; at 1.6 mya they werelower and covered with grassland (as much of the high-lands still are). Europe was otherwise mountainous andcovered over with dense forest, a formidable terrain forwarm-weather savanna dwellers. Similarly there is no ev-idence that the Mediterranean was passable at Gibraltaror anywhere else to H. erectus or earlier hominans. Theymight have reached Italy and Spain along the coasts.In northern Europe pebble tools are found earliest atHappisburgh, United Kingdom, from 0.8 mya. Thelast traces are from Kent’s Cavern, dated 0.5 mya. Bythat time H. erectus is regarded as having been ex-tinct; however, a more modern version apparently hadevolved, Homo heidelbergensis, who must have inheritedthe tools.[35] He also explains the last of the Acheulean inGermany at 0.4 mya.In the late 19th and early 20th centuries archaeologistsworked on the assumptions that a succession of Homi-nans and cultures prevailed, that one replaced another.Today the presence of multiple hominans living contem-poraneously near each other for long periods is acceptedas proved true; moreover, by the time the previously as-sumed “earliest” culture arrived in northern Europe, therest of Africa and Eurasia had progressed to the Mid-dle and Upper Palaeolithic, so that across the earth allthree were for a time contemporaneous. In any given re-gion there was a progression fromOldowan to Acheulean,Lower to Upper, no doubt.

Acheulean in Africa Main article: AcheuleanThe end of Oldowan in Africa was brought on by the

An Acheulean tool, not worked over the entire surface

appearance of Acheulean, or Mode 2, stone tools. Theearliest known instances are in the 1.7–1.6 mya layer

at Kokiselei, West Turkana, Kenya.[25] At Sterkfontein,South Africa, they are in Member 5 West, 1.7–1.4mya.[23] The 1.7 is a fairly certain, fairly standard date.Mode 2 is often found in association with H. erectus. Itmakes sense that the most advanced tools should havebeen innovated by the most advanced Hominan; conse-quently, they are typically given credit for the innovation.A Mode 2 tool is a biface consisting of two concavesurfaces intersecting to form a cutting edge all the wayaround, except in the case of tools intended to feature apoint. More work and planning go into the manufactureof aMode 2 tool. Themanufacturer hits a slab off a largerrock to use as a blank. Then large flakes are struck off theblank and worked them into bifaces by hard-hammer per-cussion on an anvil stone. Finally the edge is retouched:small flakes are hit off with a bone or wood soft ham-mer to sharpen or resharpen it. The core can be eitherthe blank or another flake. Blanks are ported for manu-facturing supply in places where nature has provided nosuitable stone.AlthoughmostMode 2 tools are easily distinguished fromMode 1, there is a close similarity of some Oldowan andsome Acheulean, which can lead to confusion. SomeOldowan tools are more carefully prepared to form amore regular edge. One distinguishing criterion is thesize of the flakes. In contrast to the Oldowan “small flake”tradition, Acheulean is “large flake:" “The primary tech-nological distinction remaining between Oldowan andthe Acheulean is the preference for large flakes (>10cm) as blanks for making large cutting tools (handaxesand cleavers) in the Acheulean.”[36] “Large Cutting Tool(LCT)" has become part of the standard terminology aswell.[22]

In North Africa, the presence of Mode 2 remains a mys-tery, as the oldest finds are from Thomas Quarry inMorocco at 0.9 mya.[34] Archaeological attention, how-ever, shifts to the Jordan Rift Valley, an extension ofthe East African Rift Valley (the east bank of the Jor-dan is slowly sliding northward as East Africa is thrustaway from Africa). Evidence of use of the Nile Valleyis in deficit, but Hominans could easily have reached thepalaeo-Jordan river from Ethiopia along the shores of theRed Sea, one side or the other. A crossing would not havebeen necessary, but it is more likely there than over a the-oretical but unproven land bridge through either Gibraltaror Sicily.Meanwhile Acheulean went on in Africa past the 1.0 myamark and also past the extinction of H. erectus there. Thelast 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, classified asH. heidelbergensis, a more advanced, but not yet mod-ern, descendant most likely of H. erectus. The ThomanQuarry Hominans in Morocco similarly are most likelyHomo rhodesiensis,[37] in the same evolutionary status as

5.3. CHRONOLOGY 41

H. heidelbergensis.

Acheulean out of Africa Mode 2 is first known outof Africa at 'Ubeidiya, Israel, a site now on the JordanRiver, then frequented over the long term (hundreds ofthousands of years) by Homo on the shore of a variable-level palaeo-lake, long since vanished. The geology wascreated by successive “transgression and regression” ofthe lake[38] resulting in four cycles of layers. The toolsare located in the first two, Cycles Li (Limnic Inferior)and Fi (Fluviatile Inferior), but mostly in Fi. The cy-cles represent different ecologies and therefore differentcross-sections of fauna, which makes it possible to datethem. They appear to be the same faunal assemblages asthe Ferenta Faunal Unit in Italy, known from excavationsat Selvella and Pieterfitta, dated to 1.6–1.2 mya.[39]

At 'Ubeidiya the marks on the bones of the animal speciesfound there indicate that the manufacturers of the toolsbutchered the kills of large predators, an activity that hasbeen termed “scavenging.”[40] There are no living floors,nor did they process bones to obtain the marrow. Theseactivities cannot be understood therefore as the only oreven the typical economic activity of Hominans. Theirinterests were selective: they were primarily harvestingthe meat of Cervids,[41] which is estimated to have beenavailable without spoiling for up to four days after the kill.The majority of the animals at the site were of“Palaearctic biogeographic origin.”[42] However, theseoverlapped in range on 30–60% of “African biogeo-graphic origin.”[43] The biome was Mediterranean, notsavanna. The animals were not passing through; therewas simply an overlap of normal ranges. Of the Homi-nans, H. erectus left several cranial fragments. Teeth ofundetermined species may have been H. ergaster.[44] Thetools are classified as “Lower Acheulean” and “DevelopedOldowan.” The latter is a disputed classification createdby Mary Leakey to describe an Acheulean-like traditionin Bed II at Olduvai. It is dated 1.53–1.27 mya. The dateof the tools therefore probably does not exceed 1.5 mya;1.4 is often given as a date. This chronology, which isdefinitely later than in Kenya, supports the “out of Africa”hypothesis for Acheulean, if not for the Hominans.From Southwest Asia, as the Levant is now called, theAcheulean extended itself more slowly eastward, arrivingat Isampur, India, about 1.2 mya. It does not appear inChina and Korea until after 1mya and not at all in Indone-sia. There is a discernible boundary marking the furthestextent of the Acheulean eastward before 1 mya, called theMovius Line, after its proposer, Hallam L. Movius. Onthe east side of the line the small flake tradition contin-ues, but the tools are additionally worked Mode 1, withflaking down the sides. In Athirampakkam at Chennai inTamil Nadu the Acheulean age started at 1.51 mya and itis also prior than North India and Europe.[45]

The cause of the Movius Line remains speculative,whether it represents a real change in technology or a lim-

itation of archeology, but after 1 mya evidence not avail-able to Movius indicates the prevalence of Acheulean.For example, the Acheulean site at Bose, China, is dated0.803±3K mya.[46] The authors of this chronologicallylater East Asian Acheulean remain unknown, as doeswhether it evolved in the region or was brought in.There is no named boundary line between Mode 1 andMode 2 on the west; nevertheless, Mode 2 is equally latein Europe as it is in the Far East. The earliest comesfrom a rock shelter at Estrecho de Quípar in Spain, datedto greater than 0.9 mya. Teeth from an undeterminedHominan were found there also.[47] The last Mode 2 inSouthern Europe is from a deposit at Fontana Ranuccionear Anagni in Italy dated to 0.45 mya, which is generallylinked toHomo cepranensis, a “late variant ofH. erectus,”a fragment of whose skull was found at Ceprano nearby,dated 0.46 mya.[48]

Middle Paleolithic

Main article: Middle Paleolithic

This period is best known as the era during which theNeanderthals lived in Europe and the Near East (c.300,000–28,000 years ago). Their technology is mainlythe Mousterian, but Neanderthal physical characteris-tics have been found also in ambiguous association withthe more recent Châtelperronian archeological culturein Western Europe and several local industries like theSzeletian in Eastern Europe/Eurasia. There is no evi-dence for Neanderthals in Africa, Australia or the Amer-icas.Neanderthals nursed their elderly and practised ritualburial indicating an organised society. The earliest evi-dence (Mungo Man) of settlement in Australia dates toaround 40,000 years ago when modern humans likelycrossed from Asia by island-hopping. Evidence for sym-bolic behavior such as body ornamentation and burial isambiguous for the Middle Paleolithic and still subject todebate. The Bhimbetka rock shelters exhibit the earliesttraces of human life in India, some of which are approx-imately 30,000 years old.

Upper Paleolithic

Main article: Upper Paleolithic

From 50,000 to 10,000 years ago in Europe, the Up-per Paleolithic ends with the end of the Pleistocene andonset of the Holocene era (the end of the last ice age).Modern humans spread out further across the Earth dur-ing the period known as the Upper Paleolithic. The Up-per Paleolithic is marked by a relatively rapid successionof often complex stone artifact technologies and a largeincrease in the creation of art and personal ornaments.

42 CHAPTER 5. STONE AGE

During period between 35 and 10 kya evolved: from 38to 30 kya Châtelperronian, 40–28 Aurignacian, 28–22Gravettian, 22–17 Solutrean, and 18–10 Magdalenian.All of these industries except the Châtelperronian are as-sociated with anatomically modern humans. Authorshipof the Châtelperronian is still the subject of much debate.The Americas were colonised via the Bering land bridgewhich was exposed during this period by lower sea levels.These people are called the Paleo-Indians, and the earli-est accepted dates are those of the Clovis culture sites,some 13,500 years ago. Globally, societies were hunter-gatherers but evidence of regional identities begins to ap-pear in the wide variety of stone tool types being devel-oped to suit very different environments.

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 charac-terized by rising sea levels and a need to adapt to a chang-ing environment and find new food sources. The devel-opment of Mode 5 (microlith) tools began in response tothese changes. They were derived from the previous Pale-olithic tools, hence the term Epipaleolithic, or were inter-mediate between the Paleolithic and the Neolithic, hencethe term Mesolithic (Middle Stone Age). The choice ofa word depends on exact circumstances and the inclina-tion of the archaeologists excavating the site. Microlithswere used in the manufacture of more efficient compositetools, resulting in an intensification of hunting and fish-ing and with increasing social activity the developmentof more complex settlements, such as Lepenski Vir. Do-mestication of the dog as a hunting companion probablydates to this period.The earliest known battle occurred during the Mesolithicperiod at a site in Egypt known as Cemetery 117.

Neolithic

Main article: NeolithicThe Neolithic, New Stone Age, was approximately char-acterized by the adoption of agriculture, the shift fromfood gathering to food producing in itself is one of themost revolutionary changes in human history so-calledNeolithic Revolution, the development of pottery, pol-ished stone tools and more complex, larger settlementssuch as Çatal Hüyük and Jericho. Some of these featuresbegan in certain localities even earlier, in the transitionalMesolithic. The first Neolithic cultures started around7000 BCE in the fertile crescent and spread concentri-cally to other areas of the world; however, the Near Eastwas probably not the only nucleus of agriculture, the cul-tivation of maize in Meso-America and of rice in the FarEast being others.

Ġgantija temples, Gozo. Some of the world’s oldest free-standingstructures.

Skara Brae, Scotland. Europe’s most complete Neolithic village

Due to the increased need to harvest and process plants,ground stone and polished stone artifacts became muchmore widespread, including tools for grinding, cutting,and chopping. Skara Brae located on Orkney islandoff Scotland is one of Europe’s best examples of a Ne-olithic village. The community contains stone beds,shelves and even an indoor toilet linked to a stream. Thefirst large-scale constructions were built, including set-tlement towers and walls, e.g., Jericho and ceremonialsites, e.g.: Stonehenge. The Ġgantija temples of Gozoin the Maltese archipelago are the oldest surviving freestanding structures in the world, erected c. 3600–2500BCE. The earliest evidence for established trade exists inthe Neolithic with newly settled people importing exoticgoods over distances of many hundreds of miles.These facts show that there were sufficient resources andco-operation to enable large groups to work on theseprojects. To what extent this was a basis for the devel-opment of elites and social hierarchies is a matter of on-going debate.[49] Although some late Neolithic societiesformed complex stratified chiefdoms similar to Polyne-sian societies such as the Ancient Hawaiians, based on thesocieties of modern tribesmen at an equivalent technolog-ical level, most Neolithic societies were relatively simpleand egalitarian.[50] A comparison of art in the two agesleads some theorists to conclude that Neolithic cultures

5.4. MATERIAL CULTURE 43

were noticeably more hierarchical than the Paleolithiccultures that preceded them.[51]

5.3.2 Three-stage chronology

The Earlier or Early Stone Age (ESA)

Main articles: Paleolithic and Lower PaleolithicThis period is not to be identified with “Old Stone Age”,

Acheulean biface from Lake Langano area, Ethiopia.

a translation of Paleolithic, or with Paleolithic, or withthe “Earlier Stone Age” that originally meant what be-came the Paleolithic andMesolithic. In the initial decadesof its definition by the Pan-African Congress of Prehis-tory, it was parallel in Africa to the Upper and MiddlePaleolithic. However, since then Radiocarbon dating hasshown that the Middle Stone Age is in fact contempo-raneous with the Middle Paleolithic.[52] The Early StoneAge therefore is contemporaneous with the Lower Pale-olithic and happens to include the same main technolo-gies, Oldowan and Acheulean, which produced Mode 1and Mode 2 stone tools respectively. A distinct regionalterm is warranted, however, by the location and chronol-ogy of the sites and the exact typology.

The Middle Stone Age (MSA)

Main article: Middle Stone Age

The Middle Stone Age was a period of African prehis-tory between Early Stone Age and Late Stone Age. It be-gan around 300,000 years ago and ended around 50,000

years ago.[53] It is considered as an equivalent of Euro-pean Middle Paleolithic.[54] It is associated with anatom-ically modern or almost modern Homo sapiens. Earlyphysical evidence comes from Omo [55] and Herto,[56]both in Ethiopia and dated respectively at c. 195 ka andat c. 160 ka.

The Later Stone Age (LSA)

Main article: Later Stone Age

The Later Stone Age (LSA, sometimes also called theLate Stone Age) refers to a period in African prehistory.Its beginnings are roughly contemporaneous with the Eu-ropean Upper Paleolithic. It lasts until historical timesand this includes cultures corresponding to Mesolithicand Neolithic in other regions.

5.4 Material culture

5.4.1 Tools

Stone tools were made from a variety of stone. For ex-ample, flint and chert were shaped (or chipped) for useas cutting tools and weapons, while basalt and sandstonewere used for ground stone tools, such as quern-stones.Wood, bone, shell, antler (deer) and other materials werewidely used, as well. During the most recent part ofthe period, sediments (such as clay) were used to makepottery. Agriculture was developed and certain animalswere domesticated.Some species of non-Primates are able to use stone tools,such as the Sea Otter, which breaks Abalone shells withthem. Primates can both use andmanufacture stone tools.This combination of abilities is more marked in apes andmen, but only men, or more generally Hominans, dependon tool use for survival.[57] The key anatomical and be-havioral features required for tool manufacture, which arepossessed only byHominans, are the larger thumb and theability to hold by means of an assortment of grips.[58]

5.4.2 Food and drink

Main articles: Paleolithic diet and Paleolithic diet andnutrition

Food sources of the Palaeolithic hunter-gatherers werewild plants and animals harvested from the environment.They liked animal organ meats, including the livers,kidneys and brains. Large seeded legumes were partof the human diet long before the agricultural revolu-tion, as is evident from archaeobotanical finds from theMousterian layers of Kebara Cave, in Israel.[59] More-over, recent evidence indicates that humans processed

44 CHAPTER 5. STONE AGE

and consumed wild cereal grains as far back as 23,000years ago in the Upper Paleolithic.[60]

Near the end of theWisconsin glaciation, 15,000 to 9,000years ago, mass extinction of Megafauna such as theWoolymammoth occurred in Asia, Europe, North Amer-ica and Australia. This was the first Holocene extinc-tion event. It possibly forced modification in the dietaryhabits of the humans of that age and with the emergenceof agricultural practices, plant-based foods also becamea regular part of the diet. A number of factors have beensuggested for the extinction: certainly over-hunting, butalso deforestation and climate change.[61] The net effectwas to fragment the vast ranges required by the large an-imals and extinguish them piecemeal in each fragment.

5.4.3 Shelter and habitat

Around 2 million years ago, Homo habilis is believedto have constructed the first man-made structure in EastAfrica, consisting of simple arrangements of stones tohold branches of trees in position. A similar stone cir-cular arrangement believed to be around 380,000 yearsold was discovered at Terra Amata, near Nice, France.(Concerns about the dating have been raised, see TerraAmata). Several human habitats dating back to the StoneAge have been discovered around the globe, including:

• A tent-like structure inside a cave near the Grotte duLazaret, Nice, France.

• A structure with a roof supported with timber, dis-covered in Dolni Vestonice, The Czech Republic,dates to around 23,000 BCE. The walls were madeof packed clay blocks and stones.

• Many huts made of mammoth bones were found inEastern Europe and Siberia. The people who madethese huts were expert mammoth hunters. Exam-ples have been found along the Dniepr river valleyof Ukraine, including near Chernihiv, in Moravia,Czech Republic and in southern Poland.

• An animal hide tent dated to around 15000 to 10000BCE, in the Magdalenian, was discovered at PlateauParain, France.

5.4.4 Art

Prehistoric art is visible in the artifacts. Prehistoric musicis inferred from found instruments, while parietal art canbe found on rocks of any kind. The latter are petroglyphsand rock paintings. The art may or may not have had areligious function.

Petroglyphs

Main article: Petroglyph

Petroglyphs appeared in the Neolithic. A Petroglyph is anintaglio abstract or symbolic image engraved on naturalstone by various methods, usually by prehistoric peoples.They were a dominant form of pre-writing symbols. Pet-roglyphs have been discovered in different parts of theworld, including Asia (Bhimbetka, India), North Amer-ica (DeathValleyNational Park), SouthAmerica (CumbeMayo, Peru), and Europe (Finnmark, Norway).

Rock paintings

Rock painting at Bhimbetka, India, a World heritage site

Main article: Cave painting

In paleolithic times, mostly animals were painted, in the-ory ones that were used as food or represented strength,such as the rhinoceros or large cats (as in the ChauvetCave). Signs such as dots were sometimes drawn.Rare human representations include handprints and half-human/half-animal figures. The Cave of Chauvet in theArdèche département, France, contains the most impor-tant cave paintings of the paleolithic era, dating fromabout 31,000 BCE. The Altamira cave paintings in Spainwere done 14,000 to 12,000 BCE and show, among oth-ers, bisons. The hall of bulls in Lascaux, Dordogne,France, dates from about 15,000 to 10,000 BCE.The meaning of many of these paintings remains un-known. They may have been used for seasonal ritu-als. The animals are accompanied by signs that suggesta possible magic use. Arrow-like symbols in Lascaux aresometimes interpreted as calendar or almanac use, but theevidence remains interpretative.[62]

Some scenes of the Mesolithic, however, can be typedand therefore, judging from their various modifications,are fairly clear. One of these is the battle scene betweenorganized bands of archers. For example, “the marchingWarriors,” a rock painting at Cingle de la Mola, Castellónin Spain, dated to about 7,000–4,000 BCE, depicts about

5.5. MODERN POPULAR CULTURE AND THE STONE AGE 45

50 bowmen in two groups marching or running in steptoward each other, each man carrying a bow in one handand a fistful of arrows in the other. A file of fivemen leadsone band, one of whom is a figure with a “high crownedhat.” In other scenes elsewhere, the men wear head-dresses and knee ornaments but otherwise fight nude.Some scenes depict the dead and wounded, bristling witharrows.[63] One is reminded of Ötzi the Iceman, a CopperAge mummy revealed by an Alpine melting glacier, whocollapsed from loss of blood due to an arrow wound inthe back.

5.4.5 Stone Age rituals and beliefs

Main articles: Paleolithic religion, Prehistoric religionand Mother goddessModern studies and the in-depth analysis of finds dating

Poulnabrone dolmen in County Clare, Ireland

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

from the Stone Age indicate certain rituals and beliefs ofthe people in those prehistoric times. It is now believedthat activities of the Stone Age humans went beyond theimmediate requirements of procuring food, body cover-ings, and shelters. Specific rites relating to death andburial were practiced, though certainly differing in styleand execution between cultures.

• Megalithic tombs, multichambered, and dolmens,single-chambered, were graves with a huge stoneslab stacked over other similarly large stone slabs;they have been discovered all across Europe andAsia and were built in the Neolithic and the BronzeAge.

5.5 Modern popular culture andthe Stone Age

Imaginative depiction of the Stone Age, by Viktor Vasnetsov

The image of the caveman is commonly associated withthe Stone Age. For example, the 2003 documentary se-ries showing the evolution of humans through the StoneAge was calledWalking with Cavemen, although only thelast programme showed humans living in caves. Whilethe idea that human beings and dinosaurs coexisted issometimes portrayed in popular culture in cartoons, filmsand computer games, such as The Flintstones, OneMillionYears B.C. and Chuck Rock, the notion of hominids andnon-avian dinosaurs co-existing is not supported by anyscientific evidence.Other depictions of the Stone Age include the best-sellingEarth’s Children series of books by Jean M. Auel, whichare set in the Paleolithic and are loosely based on ar-chaeological and anthropological findings. The 1981 filmQuest for Fire by Jean-Jacques Annaud tells the story ofa group of neanderthals searching for their lost fire. Atwenty first century series, Chronicles of Ancient Dark-ness by Michelle Paver tells of two New Stone Age chil-dren fighting to fulfil a prophecy and save their clan.

5.6 See also

• Megalith

• Prehistoric warfare

• Ice Age

• Pleistocene

46 CHAPTER 5. STONE AGE

• Homo

• Timeline of the Stone Age

5.7 Notes[1] http://www.nhm.ac.uk/about-us/news/2010/august/

oldest-tool-use-and-meat-eating-revealed75831.html

[2] Barham & Mitchell 2008, p. 106

[3] Barham & Mitchell 2008, p. 147

[4] Rogers & Semaw 2009, pp. 162–163

[5] Rogers & Semaw 2009, p. 155

[6] As to whether aethiopicus is the genus Australopithecus orthe genus Paranthropus, broken out to include the morerobust forms, anthropological opinion is divided and bothusages occur in the professional sources.

[7] Rogers & Semaw 2009, p. 164

[8] “Neolithic Vinca was a metallurgical culture”. ArchaeoNews. Reuters. 17 November 2007. Retrieved 25 Jan-uary 2011.

[9] “ASA Statement on the use of 'primitive' as a descriptor ofcontemporary human groups”. ASA News (Association ofSocial Anthropologists of the UK and Commonwealth).27 August 2007.

[10] Clark 1970, p. 22

[11] Clark 1970, pp. 18–19

[12] Deacon & Deacon 1999, pp. 5–6

[13] Isaac, Glynn (1982). “The Earliest ArchaeologicalTraces”. In Clark, J. Desmond. The Cambridge Historyof Africa. Volume. I: From the Earliest Times to C. 500BC. Cambridge: Cambridge University Press. p. 246.

[14] Willoughby, Pamela R. (2007). The evolution of modernhumans in Africa: a comprehensive guide. Lanham, MD:AltaMira Press. p. 54.

[15] Barham & Mitchell 2008, p. 477

[16] “History: Systematic Investigation of the African LaterTertiary and Quaternary”. TheWenner-Gren Foundation.Retrieved 3 March 2011.

[17] “Worsaae, Jens Jacob Asmussen”. Encyclopædia Britan-nica.

[18] Toth, Nicholas; Schick, Kathy (2007). “21 Overviewof Paleolithic Archaeology”. In Henke, H.C. Winfried;Hardt, Thorolf; Tattersall, Ian. Handbook of Paleoan-thropology. 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.

[21] Shea 2010, p. 49

[22] Shea 2010, p. 50

[23] Barham & Mitchell 2008, p. 132

[24] Barham & Mitchell 2008, pp. 126–127.

[25] Barham & Mitchell 2008, p. 128

[26] Barham & Mitchell 2008, p. 145

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

[28] Barham & Mitchell 2008, p. 112

[29] Shea 2010, p. 57

[30] Barham & Mitchell 2008, p. 73

[31] Brooks, Charles E.P. (1919), “The Correlation of theQuaternary Deposits of the British Isles with Those ofthe Continent of Europe”, Annual Report of the Board ofRegents of the Smithsonian Institution 1917, Washington:Government Pronting Office, p. 277

[32] Hugo Obermaier; Christine Matthew; Henry Osborne(1924). Fossil Man in Spain. New Haven: Yale UniversityPress for the Hispanic Society of America. p. 272.

[33] Barham & Mitchell 2008, pp. 106–107

[34] Shea 2010, pp. 55–57

[35] Barham & Mitchell 2008, p. 24

[36] Barham & Mitchell 2008, p. 130

[37] Jean-Paul Raynal; et al. (2010). “Hominid Cave atThomas Quarry I (Casablanca, Morocco): Recent find-ings and their context”. Quaternary International (223–224): 369–382.

[38] Belmaker 2006, p. 9

[39] Belmaker 2006, pp. 119–120

[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”. TheHindu.

[46] “Bose, China”. What Does It Mean to be Human?. Smith-sonian National Museum of Natural History.

[47] Dalton, Rex (2 September 2009). “Europe’soldest axes discovered”. Nature News (Nature).doi:10.1038/news.2009.878.

[48] Giovanni Muttoni; et al. (2009). “Pleistocene mag-netochronology of early hominid sites at Ceprano andFontana Ranuccio, Italy”. Earth and Planetary ScienceLetters 286: 255–268. doi:10.1016/j.epsl.2009.06.032.

5.8. REFERENCES 47

[49] Kuijt, Ian (2000). “Chapter 13: Near Eastern NeolithicResearch: Directions and Trends”. In Kuijt, Ian. Lifein Neolithic Farming Communities: Social Organization,Identity, and differentiation. Fundamental Issues in Ar-chaeology. New York: Kluwer Academic/Plenum Pub-lishers. p. 317

[50] Boehm, Christopher (2000). “The Origin of Morality asSocial Control”. In Katz, Leonard D. Evolutionary Ori-gins of Morality: Cross-disciplinary Perspectives. Journalof Consciousness Studies Volume 7. Thorverton: ImprintAcademic. p. 158. ISBN 0-7190-5612-8

[51] Guthrie, Russell Dale (2005). The Nature of PaleolithicArt. Chicago: University of Chicago Press. pp. 419–420.ISBN 978-0-226-31126-5.

[52] Clark, J. Desmond (1982). “The Culture of the MiddlePaleolithic/MIddle Stone Age”. In Clark, J. Desmond.The Cambridge History of Africa. Volume. I: From theEarliest Times to C. 500 BC. Cambridge: CambridgeUniversity Press. p. 248.

[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 (March2005). “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 inthe Upper Palaeolithic revealed by starch grain analysis”.Nature 430 (7000): 670–3. doi:10.1038/nature02734.PMID 15295598.

[61] Turvey, Samuel T. (2009). “Chapter 2: In the shadow ofthe megafauna: prehistoric mammal and bird extinctionsacross the Holocene”. In Turvey, Samuel T. Holocene Ex-tinctions. Oxford Biology. Oxford: Oxford UniversityPress. pp. 16–17

[62] Aczel, Amir D. (2000). The Cave and the Cathedral: Howa Real-Life Indiana Jones and a Research Scholar De-coded the Ancient Art of Man. Hoboken: John Wiley &Sons Inc. pp. 157–158.

[63] Martínez, Antonio Beltrán (1982) [1979]. Rock art of theSpanish Levant. The Imprint of Man. Cambridge: Cam-bridge University Press. pp. 48–51.

[64] Salvatore Piccolo, Ancient Stones: The Prehistoric Dol-mens of Sicily. Abingdon, 2013.

5.8 References• Barham, Lawrence; Mitchell, Peter (2008). The

First Africans: African Archaeology from the Earli-est Toolmakers to Most Recent Foragers. CambridgeWorld Archaeology. Oxford: Oxford UniversityPress.

• Belmaker, Miriam (March 2006). CommunityStructure through Time: 'Ubeidiya, a Lower Pleis-tocene Site as a Case Study (Thesis). Paleoanthro-pology Society.

• Clark, J. Desmond (1970). The Prehistory of Africa.Ancient People and Places, Volume 72. New York;Washington: Praeger Publishers.

• Deacon, Hilary John; Deacon, Janette (1999). Hu-man beginnings in South Africa: uncovering the se-crets of the Stone Age. Walnut Creek, Calif. [u.a.]:Altamira Press.

• Piccolo, Salvatore (2013). Ancient Stones: The Pre-historic Dolmens of Sicily. Abingdon (UK): BrazenHead Publishing.

• Rogers, Michael J.; Semaw, Sileshi (2009). “FromNothing to Something: The Appearance and Con-text of the Earliest Archaeological Record”. InCamps i Calbet, Marta; Chauhan, Parth R. Source-book of paleolithic transitions: methods, theories,and interpretations. New York: Springer.

• Schick, Kathy D.; Nicholas Toth (1993). Mak-ing Silent Stones Speak: Human Evolution and theDawn of Technology. New York: Simon & Schus-ter. ISBN 0-671-69371-9.

• Shea, John J. (2010). “Stone Age Visiting Cards Re-visited: a Strategic Perspective on the Lithic Tech-nology of Early Hominin Dispersal”. In Fleagle,John G.; Shea, John J.; Grine, Frederick E.; Boden,Andrea L.; Leakey, Richard E,. Out of Africa I: theFirst 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”. His-tory World International. Retrieved 22 February2011.

48 CHAPTER 5. STONE AGE

• Kowalski, D.R. “Stone Age Hand-axes”. Aerobi-ologicalEngineering.com. Retrieved 22 February2011.

• Kowalski, D.R. “Stone Age Habitats”. Aerobiolog-icalEngineering.com. Retrieved 22 February 2011.

• “PanAfrican Archaeological Association”. Re-trieved 28 February 2011.

• “Society of Africanist Archaeologists”. Retrieved 3March 2011.

• “The ASA”. Association of Social Anthropologistsof the UK and Commonwealth.

Chapter 6

Woolly mammoth

The woolly mammoth (Mammuthus primigenius) was aspecies of mammoth, the common name for the extinctelephant genus Mammuthus. The woolly mammoth wasone of the last in a line of mammoth species, beginningwithMammuthus subplanifrons in the early Pliocene. M.primigenius diverged from the steppe mammoth, M. tro-gontherii, about 200,000 years ago in eastern Asia. Itsclosest extant relative is the Asian elephant.The appearance and behaviour of this species are amongthe best studied of any prehistoric animal because of thediscovery of frozen carcasses in Siberia and Alaska, aswell as skeletons, teeth, stomach contents, dung, and de-piction from life in prehistoric cave paintings. Mammothremains had long been known in Asia before they be-came known to Europeans in the 17th century. The originof these remains was long a matter of debate, and oftenexplained as being remains of legendary creatures. Themammoth was identified as an extinct species of elephantby Georges Cuvier in 1796.The woolly mammoth was roughly the same size as mod-ern African elephants. Males reached shoulder heightsbetween 2.7 and 3.4 m (9 and 11 ft) and weighed up to6 tonnes (6.6 short tons). Females averaged 2.6–2.9 me-tres (8.5–9.5 ft) in height and weighed up to 4 tonnes (4.4short tons). A newborn calf weighed about 90 kilograms(200 lb). The woolly mammoth was well adapted to thecold environment during the last ice age. It was coveredin fur, with an outer covering of long guard hairs and ashorter undercoat. The colour of the coat varied fromdark to light. The ears and tail were short to minimisefrostbite and heat loss. It had long, curved tusks and fourmolars, which were replaced six times during the life-time of an individual. Its behaviour was similar to thatof modern elephants, and it used its tusks and trunk formanipulating objects, fighting, and foraging. The diet ofthe woolly mammoth was mainly grass and sedges. In-dividuals could probably reach the age of 60. Its habitatwas the mammoth steppe, which stretched across north-ern Eurasia and North America.The woolly mammoth coexisted with early humans, whoused its bones and tusks for making art, tools, anddwellings, and the species was also hunted for food.[1]It disappeared from its mainland range at the end of thePleistocene 10,000 years ago, most likely through climate

change and consequent shrinkage of its habitat, huntingby humans, or a combination of the two. Isolated popu-lations survived on St. Paul Island until 6,400 years agoand Wrangel Island until 4,000 years ago. After its ex-tinction, humans continued using its ivory as a raw mate-rial, a tradition that continues today. It has been proposedthe species could be recreated through cloning, but thismethod is as yet infeasible because of the degraded stateof the remaining genetic material.

6.1 Taxonomy

A mammoth tusk with Inuit carvings of scenes on the YukonRiver, 19th century.

Remains of various extinct elephants were known byEuropeans for centuries, but were generally interpreted,based on biblical accounts, as the remains of legendarycreatures such as behemoths or giants. It was also the-orised that they were remains of modern elephants thathad been brought to Europe during the Roman Repub-lic, for example the war elephants of Hannibal the Greatand Pyrrhus of Epirus, or animals that had wanderednorth.[2] The first woolly mammoth remains studied byEuropean scientists were examined by Hans Sloane in1728 and consisted of fossilised teeth and tusks fromSiberia. Sloane was the first to recognise that the remainsbelonged to elephants.[3] Sloane turned to another bibli-cal explanation for the presence of elephants in the Arc-tic, asserting that they had been buried during the GreatFlood, and that Siberia had previously been tropical priorto a drastic climate change.[4] Others interpreted Sloane’sconclusion slightly differently, arguing the flood had car-ried elephants from the Tropics to the Arctic. Sloane’s

49

50 CHAPTER 6. WOOLLY MAMMOTH

paper was based on travellers’ descriptions and a few scat-tered bones collected in Siberia and Britain. He discussedthe question of whether or not the remains were from ele-phants, but drew no conclusions.[5]

In 1738, Johann Philipp Breyne argued that mammothfossils represented some kind of elephant. He could notexplain why a tropical animal would be found in such acold area as Siberia, and suggested that they might havebeen transported there by the Great Flood.[6] In 1796,French anatomist Georges Cuvier was the first to identifythe woolly mammoth remains not as modern elephantstransported to the Arctic, but as an entirely new species.He argued this species had gone extinct and no longerexisted, a concept that was not widely accepted at thetime.[2][7]

6.1.1 Etymology

Cuvier’s 1796 comparison between the mandible of a woollymammoth and an Indian elephant.

Following Cuvier’s identification, Johann Friedrich Blu-menbach gave the woolly mammoth its scientific name,Elephas primigenius, in 1799, placing it in the same genusas the Asian elephant. This name means “the firstbornelephant”. Henry Fairfield Osborn chose a molar fromBlumenbach’s collection as the lectotype specimen for

the species, since holotype designation was not practisedin Blumenbach’s time.[8] In 1828 Joshua Brookes recog-nised the species was distinct enough to warrant a newgenus, and reclassified it as Mammuthus primigenius.[9]It is unclear where and how the word “mammoth” orig-inated. According to the Oxford English Dictionary, itcomes from an old Vogul word mēmoŋt 'earth-horn'. [10]It may be a version of mehemot, the Arabic version ofthe biblical word “behemoth”. Another possible origin isEstonian, where maameans earth, and mutt means mole.The word was first used in Europe during the early 17thcentury, when referring to maimanto tusks discovered inSiberia.[11] Thomas Jefferson, who had a keen interestin palaeontology, is partially responsible for transform-ing the word mammoth from a noun describing the pre-historic elephant to an adjective describing anything ofsurprisingly large size. The first recorded use of the wordas an adjective was in a description of a wheel of cheese(the "Cheshire Mammoth Cheese") given to Jefferson in1802.[12]

6.1.2 Evolution

The earliest known proboscideans, the clade which con-tains elephants, existed about 55 million years ago aroundthe Tethys Sea. The closest known relatives of the Pro-boscidea are the sirenians and the hyraxes. The fam-ily Elephantidae existed six million years ago in Africaand includes the modern elephants and the mammoths.Among many now extinct clades, the mastodon is only adistant relative of the mammoths, and part of the separateMammutidae family, which diverged 25million years be-fore the mammoths evolved.[13] The following cladogramshows the placement of the genus Mammuthus amongother proboscideans, based on hyoid characteristics:[14]

Comparison between a woolly mammoth (L) and an Americanmastodon (R).

In 2005, researchers assembled a complete mitochondrialgenome profile of the woolly mammoth, which allowedthem to trace the close evolutionary relationship betweenmammoths and Asian elephants, Elephas maximus.[15]African elephants, Loxodonta africana, branched awayfrom this clade around 6 million years ago, close tothe time of the similar split between chimpanzees andhumans. Before the publication of the Neanderthalgenome, many researchers expected the first fully se-quenced nuclear genome of an extinct species would bethat of the mammoth.[16] A 2010 study confirmed theserelationships, and suggested the mammoth and Asian ele-

6.2. DESCRIPTION 51

phant lineages diverged 5.8–7.8 million years ago, whileAfrican elephants diverged from an earlier common an-cestor 6.6–8.8 million years ago.[17] In 2008, much ofthe woolly mammoth’s chromosomal DNA was mapped.The analysis showed that the woolly mammoth and theAfrican elephant are 98.55% to 99.40% identical.[18] Theteam mapped the woolly mammoth’s nuclear genome se-quence by extracting DNA from the hair follicles of botha 20,000-year-old mammoth retrieved from permafrost,and another that died 60,000 years ago.[19] In 2012,proteins were confidently identified for the first time, col-lected from a 43,000-year-old woolly mammoth.[20]

M. jeffersonii, a possible hybrid between Columbian and woollymammoths.

Since many remains of each species of mammoth areknown from several localities, it is possible to recon-struct the evolutionary history of the genus throughmorphological studies. Mammoth species can be identi-fied from the number of enamel ridges on their molars;primitive species had few ridges, and the number in-creased gradually as new species evolved and replacedthe preceding ones. The crowns of the teeth lengthenedand the skulls became taller to accommodate this. At thesame time, the skulls became shorter from front to backto minimise the weight.[21] These adaptations were ac-quired gradually as mammoths turned to more abrasivefood items.[22]

The first known members of the genus Mammuthus arethe African species M. subplanifrons from the Pliocene,and M. africanavus from the Pleistocene. The former isthought to be the ancestor of later forms. Mammothsentered Europe around 3 million years ago. The earli-est type known from there has been named M. rumanus,which spread across Europe and China. Only its molarsare known, which show that it had 8–10 enamel ridges.A population evolved 12–14 ridges, splitting off from andreplacing the earlier type, becoming M. meridionalis. Inturn, this species was replaced by the steppe mammoth,M. trogontherii, with 18–20 ridges, which evolved in east-ern Asia c. 1 million years ago. Mammoths derived fromM. trogontherii evolved molars with 26 ridges 200,000years 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

Cast of an intermediate form between M. trogontherii and M.primigenius,M. p. fraasi

entered North America. A 2011 genetic study showedthat two examined specimens of the Columbian mam-moth were grouped within a subclade of woolly mam-moths. This suggests that the two populations interbredand produced fertile offspring. A North American formknown asM. jeffersonii may be a hybrid between the twospecies.[23]

Individuals and populations showing transitional mor-phologies between each of the mammoth species areknown, and primitive and derived species coexisted aswell until the former disappeared. The different speciesand their intermediate forms therefore can be termed"chronospecies". Many intermediate subspecies havealso been proposed, but their validity is uncertain; de-pending on author, they are either considered primi-tive forms of an advanced species or advanced forms ofa primitive species.[21] Regional and intermediate sub-species 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 populationswere described as dwarf varieties, much smaller than themainland woolly mammoth; the Wrangel Island popu-lation was also proposed to be a new subspecies, M. p.vrangeliensis.[25][26] The Wrangel mammoths were iso-lated for 5000 years, but experienced only a slight lossof genetic variation.[27]

6.2 Description

The appearance of the woolly mammoth is probably thebest known of any prehistoric animal due to the manyfrozen specimens with preserved soft tissue and depic-tions by contemporary humans in their art. Fully grownmales reached shoulder heights between 2.7 and 3.4 m (9and 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

Cave art from Les Combarelles, France

which commonly reach 3–3.4 m (9.8–11.2 ft), and isless than the size of the earlier mammoth species M.meridionalis and M. trogontherii, and the contemporaryM. columbi. The reason for the smaller size is unknown.Female woolly mammoths averaged 2.6–2.9 m (8.5–9.5ft) in height andwere built more lightly thanmales, weigh-ing up to 4 tonnes (4.4 short tons). A newborn calf wouldhave weighed about 90 kg (200 lb). These sizes are de-duced from comparison with modern elephants of sim-ilar size.[28] Though the mammoths on Wrangel Islandwere smaller than those of the mainland, their size var-ied, and they were not small enough to be considered“dwarves”.[29] It has been claimed that the last woollymammoth populations decreased in size and increasedtheir sexual dimorphism, but this was dismissed in a 2012study.[30]

Woolly mammoths had several adaptations to the cold,most noticeably the layer of fur covering all parts of thebody. Other adaptations to cold weather include ears thatare far smaller than those of modern elephants; they wereabout 38 cm (15 in) long and 18–28 cm (7.1–11.0 in)across, and the ear of the 6–12 month old frozen calf“Dima” was under 13 cm (5.1 in) long. The small ears re-duced heat loss and frostbite, and the tail was short for thesame reason, only 36 cm (14 in) long in the “Berezovkamammoth”. The tail contained 21 vertebrae, whereas thetails of modern elephants contain 28–33. Their skin wasno thicker than that of present-day elephants, between1.25 and 2.5 cm (0.49 and 0.98 in). They had a layer offat up to 10 cm (3.9 in) thick under the skin, which helpedto keep them warm. Woolly mammoths had broad flapsof skin under their tails which covered the anus; this isalso seen in modern elephants.[31]

Other characteristic features depicted in cave paintingsinclude a large, high, single-domed head and a slopingback with a high shoulder hump, resulting from longspinous processes on the neck vertebrae. These fea-tures were not present in juveniles, which had concavebacks like African elephants.[32] Another feature shownin cave paintings was confirmed by the discovery of afrozen specimen in 1924, an adult nicknamed the “Mid-

Model at the Royal BC Museum

dle Kolymamammoth”, which was preservedwith a com-plete trunk tip. Unlike the trunk lobes of modern ele-phants, the upper “finger” at the tip of the trunk had a longpointed lobe and was 10 cm (3.9 in) long, while the lower“thumb” was 5 cm (2.0 in) and was broader. The trunkof “Dima” was 76 cm (2.49 ft) long, whereas the trunkof 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 examinationof the skeleton. Males were generally larger and hadmorerobust skeletons and tusks. The best indication of sex isthe size of the pelvic girdle, as the birth canal is alwayswider in females than in males.[33]

6.2.1 Coat

Fur in Naturhistorisches Museum, Vienna

The coat consisted of an outer layer of long, coarse “guardhair”, which was 30 cm (12 in) on the upper part of thebody, up to 90 cm (35 in) in length on the flanks and un-derside, and 0.5 mm (0.020 in) in diameter, and a denserinner layer of shorter, slightly curly under-wool, up to 8cm (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, reachingthe toes. The hairs on the head were relatively short, butlonger on the underside and the sides of the trunk. Thetail 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 thatthe woolly mammoth moulted seasonally, and that theheaviest fur was shed during spring. Since mammoth car-casses were more likely to be preserved during autumn,it is possible that only the winter coat has been preservedin frozen specimens. Modern elephants have much lesshair, though juveniles have a more extensive covering ofhair than adults.[34] Comparison between the over-hairsof woolly mammoths and extant elephants show that theydid not differ much in overall morphology.[35] Woollymammoths had numerous sebaceous glands in their skin,which secreted oils into their hair; this would have im-proved the wool’s insulation, repelled water, and given thefur a glossy sheen.[36]

Preserved woolly mammoth fur is orange-brown, but thisis believed to be an artefact from the bleaching of pig-ment during burial. The amount of pigmentation var-ied from hair to hair and also within each hair.[31] A2006 study sequenced the Mc1r gene (which influenceshair colour in mammals) from woolly mammoth bones.Two alleles were found: a dominant (fully active) anda recessive (partially active) one. In mammals, reces-sive Mc1r alleles results in light hair. Mammoths bornwith at least one copy of the dominant allele would havehad dark coats, while those with two copies of the reces-sive allele would have had light coats.[37] A 2011 studyshowed that light individuals would have been rare.[38] A2014 study instead indicated that the colouration of anindividual varied from non-pigmented on the overhairs,bi coloured, non-pigmented and mixed red-brown guardhairs, and non-pigmented underhairs, which would give alight overall appearance.[39]

6.2.2 Dentition

Woolly mammoths had very long tusks, which were morecurved than those of modern elephants. The largestknown male tusk is 4.2 m (14 ft) long and weighs 91 kg(201 lb), but 2.4–2.7 m (7.9–8.9 ft) and 45 kg (99 lb) wasa more typical size. Female tusks averaged at 1.5–1.8 m(4.9–5.9 ft) and weighed 9 kg (20 lb). About a quarterof the length was inside the sockets. The tusks grew spi-rally in opposite directions from the base and continuedin a curve until the tips pointed towards each other. Inthis way, most of the weight would have been close to theskull, and there would be less torque than with straighttusks. The tusks were usually asymmetrical and showedconsiderable variation, with some tusks curving down in-stead of outwards and some being shorter due to break-age. Calves developed small milk tusks a few centimetreslong at six months old, which were replaced by perma-nent tusks a year later. Tusk growth continued throughoutlife but became slower as the animal reached adulthood.The tusks grew by 2.5–15 cm (0.98–5.91 in) each year.Some cave paintings show woolly mammoths with smallor no tusks, but it is unknown whether this reflected real-ity or was artistic license. Female Asian elephants have no

Specimen with asymmetrical tusks, the Smithsonian Museum

tusks, but there is no fossil evidence that any adult woollymammoths lacked them.[40]

Molar from font de Champdamoy, France

Woolly mammoths had four functional molar teeth at atime, two in the upper jaw and two in the lower. 23 cm(9.1 in) of the crownwas within the jaw, and 2.5 cm (1 in)was above. The crown was continually pushed forwardsand up as it wore down, comparable to a conveyor belt.The teeth had up to 26 separated ridges of enamel, whichwere themselves covered in “prisms” that were directedtowards the chewing surface. These were quite wear re-sistant and kept together by cementum and dentine. Amammoth had six sets of molars throughout a lifetime,which were replaced five times, though a few specimenswith a seventh set are known. The latter condition couldextend the lifespan of the individual, unless the tooth onlyconsisted of a few plates. The first molars were aboutthe size of those of a human, 1.3 cm (0.51 in), the thirdwere 15 cm (6 in) 15 cm (5.9 in) long, and the sixth wereabout 30 cm (1 ft) long and weighed 1.8 kg (4 lb). Themolars grew larger and contained more ridges with each

54 CHAPTER 6. WOOLLY MAMMOTH

replacement.[41]

Distortion in the molars is the most common health prob-lem found in woolly mammoth fossils. Sometimes thereplacement was disrupted, and the molars were pushedinto abnormal positions, but some animals are knownto have survived this. Teeth from Britain showed that2% of specimens had periodontal disease, with half ofthese containing caries. The teeth also sometimes hadcancerous growths.[42]

6.3 Palaeobiology

Restoration of a herd walking near the Somme River by CharlesR. Knight

Adult woolly mammoths could effectively defend them-selves from predators with their tusks, trunks and size, butjuveniles and weakened adults were vulnerable to packhunters such as wolves, cave hyenas and large felines. Thetusks may also have been used in intra-species fighting,such as territorial fights or fights over mates. Becauseof their curvature, the tusks were not suitable for stab-bing, but may have been used for hitting, as indicatedby injuries to some fossil shoulder blades. As in mod-ern elephants, the sensitive and muscular trunk workedas a limb-like organ with many functions. It was usedfor manipulating objects, and in social interactions. Thevery long hairs on the tail probably compensated for theshortness of the tail, enabling its use as a flyswatter, simi-lar to the tail on modern elephants.[43] As in reindeer andmusk oxen, the haemoglobin of the woolly mammoth wasadapted to the cold, with three mutations to improve oxy-gen delivery around the body and prevent freezing. Thisfeature may have helped the mammoths to live in highlatitudes.[44]

Mounted “family group”

Like modern elephants, woolly mammoths were likelyvery social and lived in matriarchal family groups. This is

supported by fossil assemblages and cave paintings show-ing groups. It is therefore probable that most of theirother social behaviour was similar to those of modernelephants. Accumulations of modern elephant remainshave been termed "elephants’ graveyards", as these siteswere erroneously thought to be where old elephants wentto die. Similar accumulations of woolly mammoth boneshave been found; it is thought these are the result of in-dividuals dying near or in the rivers over thousands ofyears, and their bones eventually being brought togetherby the streams, or due to animals being mired in mud.Some accumulations are also thought to be the remainsof herds that died together at the same time, perhaps dueto flooding.[45]

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 Reser-voir in Canada, showing that there were in this case al-most equal numbers of adults, sub-adults and juveniles.The adults had a stride of 2 m (6.6 ft), and the juve-niles ran to keep up.[46] The well-preserved foot of theadult male "Yukagir mammoth" shows that the soles ofthe feet contained many cracks that would have helped ingripping surfaces during locomotion. Like modern ele-phants, woolly mammoths walked on their toes and hadlarge, fleshy pads behind the toes.[31]

Evidence of several different bone diseases has beenfound in woolly mammoths. The most common ofthese diseases was osteoarthritis, found in 2% of speci-mens. One specimen from Switzerland had several fusedvertebrae as a result of this condition. The “Yukagirmammoth” had suffered from spondylitis in two verte-brae, and osteomyelitis is also known from some spec-imens. Several specimens have healed bone fractures,showing that the animals had survived these injuries.[47]Parasitic flies and protozoa were identified in the gut ofthe calf “Dima”.[48]

6.3.1 Diet

Food at various stages of digestion has been found in theintestines of several woollymammoths, giving a good pic-ture of their diet. Woolly mammoths sustained them-selves on plant food, mainly grass and sedges, which weresupplemented with herbaceous plants, flowering plants,

6.3. PALAEOBIOLOGY 55

The frozen calf “Lyuba” which still had food in its stomach

shrubs, mosses, and tree matter. The composition andexact varieties differed from location to location. Woollymammoths needed a varied diet to support their growth,like modern elephants. An adult of six tonnes would needto eat 180 kg (397 lb) daily, and may have foraged aslong as twenty hours every day. The two-fingered tip ofthe trunk was probably adapted for picking up the shortgrasses of the last ice age (Quaternary glaciation, 2.58million years ago to present) by wrapping around them,whereas modern elephants curl their trunks around thelonger grass of their tropical environments. The trunkcould also be used for pulling off large grass tufts, del-icately picking buds and flowers, and tearing off leavesand branches where trees and shrubs were present. The“Yukagir mammoth” had ingested plant matter that con-tained spores of dung fungus.[49] Isotope analysis showsthat woolly mammoths fed mainly on C3 plants, unlikehorses and rhinos.[50]

Scientists identified milk in the stomach and faecal mat-ter in the intestines of the mammoth calf "Lyuba".[51]The faecal matter may have been eaten by “Lyuba” topromote development of the intestinal microbes neces-sary for digestion of vegetation, as is the case in modernelephants.[52] An isotope analysis of woolly mammothsfromYukon, Canada, showed that the young nursed for atleast three years, and were weaned and gradually changedto a diet of plants when they were two to three years old.This is later than in modern elephants and may be due toa higher risk of predator attack or difficulty in obtainingfood during the long periods of winter darkness in highlatitudes.[53]

The molars were adapted to their diet of coarse tundragrasses, with more enamel plates and a higher crown thantheir earlier, southern relatives. The woolly mammothchewed its food by using its powerful jawmuscles tomovethe mandible forwards and close the mouth, then back-

Mandibles at various growth stages, Naturalis, Leiden

wards while opening, whichmade the sharp enamel ridgescut across each other and grind the food. The ridges werewear-resistant to enable the animal to chew large quan-tities of food, which often contained grit. Woolly mam-moths may have used their tusks as shovels to clear snowfrom the ground and reach the vegetation buried below,and to break ice to drink. This is indicated on many pre-served tusks by flat, polished sections up to 30 centime-tres (12 in) long on the part of the surface that would havereached the ground. The tusks were also used for ob-taining food in other ways, such as digging up plants andstripping off bark.[54]

6.3.2 Growth and reproduction

Cross sectioned tusk with growth rings

The age of a woolly mammoth can be determined bycounting the growth rings of its tusks when viewed incross section. Each major line represents a year, andweekly and daily ones can be found in between. Darkbands correspond to summers, and it is therefore possi-ble to determine the season in which a mammoth died.The growth of the tusks slowed when it became harder toforage, for example during disease, when a male was ban-ished from the herd, and during periods of severe glacia-tion. Woolly mammoths continued growing past adult-hood, like other elephants. Unfused limb bones show thatmales grew until they reached the age of 40, and femalesgrew until they were 25. The frozen calf “Dima” was 90cm (35 in) tall when it died at the age of 6–12 months.

56 CHAPTER 6. WOOLLY MAMMOTH

At this age, the second set of molars would be in the pro-cess of erupting, and the first set would be worn out at18 months of age. The third set of molars lasted for tenyears, and this process was repeated until the final, sixthset emerged when the animal was 30 years old. A woollymammoth could probably reach the age of 60, like mod-ern elephants of the same size. By then the last set ofmolars would be worn out, the animal would be unable tochew and feed, and it would die of starvation.[55]

Head of the “Yukagir mammoth”

The best preserved head of a frozen adult specimen, thatof a male nicknamed the “Yukagir mammoth”, showsthat woolly mammoths had temporal glands between theear and the eye.[56] This feature indicates that, like bullelephants, male woolly mammoths also entered "musth",a period of heightened aggressiveness. The glands areused especially bymales to produce an oily substance witha strong smell called temporin. Their fur may have helpedin spreading the scent further.[57]

Examination of preserved calves show that they wereall born during spring and summer, and since modernelephants have gestation periods of 21–22 months, itis probable that the mating season was from summerto autumn.[58] δ15N isotopic analysis of the teeth of“Lyuba” has demonstrated their prenatal development,and indicates its gestation period was similar to that ofa modern elephant, and that it was born in spring.[59]

6.4 Distribution and habitat

Possible distribution during the last glacial period, based on lo-cations of fossil finds

The habitat of the woolly mammoth is known as"mammoth steppe" or “tundra steppe”. This environment

stretched across northern Asia, many parts of Europe,and the northern part of North America during the last iceage. It was similar to the grassy steppes of modern Rus-sia, but the flora was more diverse, abundant, and grewfaster. Grasses, sedges, shrubs, and herbaceous plantswere present, and scattered trees were mainly found insouthern regions. This habitat was not dominated by iceand snow, as is popularly believed, since these regionsare thought to have been high-pressure areas at the time.The habitat of the woolly mammoth also supported othergrazing herbivores such as the woolly rhinoceros, wildhorses and bison.[60] A 2014 study concluded that forbs (agroup of herbaceous plants) were more important in thesteppe-tundra than previously acknowledged, and that itwas a primary food source for the ice-age megafauna.[61]

Restoration of a group in late Pleistocene northern Spain, byMauricio Antón

The southernmost woolly mammoth specimen known isfrom the Shandong province of China, and is 33,000years old.[62] The southernmost European remains arefrom the Depression of Granada in Spain and are ofroughly the same age.[63] DNA studies have helped de-termine the phylogeography of the woolly mammoth. A2008 DNA study showed there were two distinct groupsof woolly mammoths: one that went extinct 45,000 yearsago and another one that went extinct 12,000 years ago.The two groups are speculated to be divergent enoughto be characterised as subspecies. The group that wentextinct earlier stayed in the middle of the high Arctic,while the group with the later extinction had a muchwider range.[64] Recent stable isotope studies of Siberianand New World mammoths have shown there were alsodifferences in climatic conditions on either side of theBering land bridge, with Siberia being more uniformlycold and dry throughout the Late Pleistocene.[65] Dur-ing the Younger Dryas age, woolly mammoths brieflyexpanded into north-east Europe, whereafter the main-land populations became extinct.[66] A2008 genetic studyshowed that some of the woolly mammoths that enteredNorth America through the Bering land bridge from Asiamigrated back about 300,000 years ago and had replacedthe previous Asian population by about 40,000 years ago,not long before the entire species went extinct.[67] Woollymammoths have been found in the same locations as thoseof the Columbian mammoth in North America, but it isunknown whether the two species were sympatric. Thewoolly mammoth may have entered these southern ar-

6.5. RELATIONSHIP WITH HUMANS 57

eas during times when Columbian mammoth populationswere absent.[68]

6.5 Relationship with humans

Woolly mammoth carved in ivory, discovered by Édouard Lartetin 1864

Modern humans coexisted with woolly mammoths dur-ing the Upper Palaeolithic period when they entered Eu-rope from Africa between 30,000 and 40,000 years ago.Prior to this, Neanderthals had coexisted with mammothsduring the Middle Palaeolithic. Woolly mammoths werevery important to ice age humans, and human survivalmay have depended on the mammoth in some areas. Ev-idence for such coexistence was not recognised until the19th century. William Buckland published his discov-ery of the Red Lady of Paviland skeleton in 1823, whichwas found in a cave alongside woolly mammoth bones,but he mistakenly denied that these were contemporaries.In 1864, Édouard Lartet found an engraving of a woollymammoth on a piece of mammoth ivory in the Abri dela Madeleine cave in Dordogne, France. This was thefirst widely accepted evidence for the coexistence of hu-mans with prehistoric extinct animals and is the first con-temporary depiction of such a creature known to modernscience.[69]

Cro-Magnon artists painting mammoths in Font-de-Gaume, byCharles R. Knight

The woolly mammoth is the thirdmost depicted animal inice age art, after horses and bison, and these images wereproduced between 35,000 and 11,500 years ago. To-

day, more than five hundred depictions of woolly mam-moths are known, in media ranging from cave paintingsand engravings on the walls of 46 caves in Russia, Franceand Spain to engravings and sculptures (termed "portableart") made from ivory, antler, stone and bone. Cave paint-ings of woolly mammoths exist in several styles and sizes.The French Rouffignac cave has most depictions, 159,and some of the drawings are more than 2 metres (6.5ft) in length. Other notable caves with mammoth de-pictions are the Chauvet Cave, Les Combarelles Cave,and Font-de-Gaume.[70] A depiction in the Cave of ElCastillo may instead show Palaeoloxodon, the “straight-tusked elephant”.[71]

“Portable art” can be more accurately dated than cave artsince it is found in the same deposits as tools and otherice age artefacts. The largest collection of portable mam-moth art, consisting of 62 depictions on 47 plaques, wasfound in the 1960s at an excavated open-air camp nearGönnersdorf in Germany. There does not seem to be acorrelation between the number of mammoths depictedand the species that were most often hunted, since rein-deer bones are the most frequently found animal remainsat the site. Two spear throwers shaped as woolly mam-moths have also been found in France.[70] Some portablemammoth depictions may not have been produced wherethey were discovered, but could have moved around byancient trading.[71]

6.5.1 Exploitation

Reconstructed bone hut, based on finds in Mezhyrich

Woolly mammoth bones were used as construction ma-terial for dwellings by both Neanderthals and modern hu-mans during the ice age. More than 70 such dwellingsare known, mainly from the Russian Plain. The bases ofthe huts were circular, and ranged from 8 to 24 squaremetres (86 to 258 sq ft). The arrangement of dwellingsvaried, and ranged from 1m (3.3 ft) to 20 m (66 ft) apart,depending on location. Large bones were used as foun-dations for the huts, tusks for the entrances, and the roofswere probably skins held in place by bones or tusks. Somehuts had floors that extended 40 cm (16 in) below ground.Some huts included fireplaces, 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-moths killed by humans, but the state of the bones, andthe fact that bones used to build a single dwelling variedby several thousands of years in age, suggests that theywere collected remains of long-dead animals. Woollymammoth bones were also made into various tools, fur-niture, and musical instruments. Large bones, such asshoulder blades, were also used to cover dead human bod-ies during burial.[72]

The Venus of Brassempouy

Woolly mammoth ivory was used to create art objects.Several Venus figurines, including the Venus of Brassem-pouy and the Venus of Lespugue, were made from thismaterial. Weapons made from ivory, such as daggers,spears, and a boomerang, are also known. To be ableto 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 straight-ened, and it is unknown how this was achieved.[73]

Several woolly mammoth specimens show evidence ofbeing butchered by humans, which is indicated by breaks,cut-marks, and associated stone tools. It is not knownhowmuch prehistoric humans relied on woolly mammothmeat, since there were many other large herbivores avail-able. Many mammoth carcasses may have been scav-enged by humans rather than hunted. Some cave paint-ings show woolly mammoths in structures interpreted aspitfall traps. Few specimens show direct, unambiguousevidence of having been hunted by humans. A Siberianspecimen with a spearhead embedded in its shoulderblade shows that a spear had been thrown at it with greatforce.[74] A specimen from the Mousterian age of Italyshows evidence of spear hunting by Neanderthals.[75] Thejuvenile specimen nicknamed “Yuka” is the first frozenmammoth with evidence of human interaction. It showsevidence of having been killed by a large predator, and ofhaving been scavenged by humans shortly after. Some ofits bones had been removed, and were found nearby.[76]A site near the Yana River in Siberia has revealed sev-eral specimens with evidence of human hunting, but thefinds were interpreted to show that the animals were not

hunted intensively, but perhaps mainly when ivory wasneeded.[77]

6.6 Extinction

16,500 year old mammoth spear thrower from France

Most woolly mammoth populations disappeared duringthe late Pleistocene and early Holocene, alongsidemost ofthe Pleistocene megafauna, during the Quaternary extinc-tion event.[78] Scientists are divided over whether hunt-ing or climate change, which led to the shrinkage of itshabitat, was the main factor that contributed to the ex-tinction of the woolly mammoth, or whether it was dueto a combination of the two. Whatever the cause, largemammals are generallymore vulnerable than smaller onesdue to their smaller population size and low reproduc-tion rates. Different woolly mammoth populations didnot die out simultaneously across their range, but grad-ually went extinct over time. The last mainland popu-lation existed in the Kyttyk Peninsula of Siberia 9,650years ago.[79] A small population of woolly mammothssurvived on St. Paul Island, Alaska, until 6,400 yearsago.[26][80][81] The last known population remained onWrangel Island in the Arctic Ocean until 4,000 yearsago.[82][83][84] Genetic evidence implies the extinction ofthis final population was sudden, rather than the culmi-nation of a gradual decline;[84] the disappearance coin-cides roughly in timewith the first evidence for humans onthe island.[85] The woolly mammoths of eastern Beringia(modern Alaska and Yukon) had similarly died out about13,300 years ago, soon (roughly 1000 years) after the firstappearance of humans in the area, which parallels the fateof all the other late Pleistocene proboscids (mammoths,gomphotheres and mastodons), as well as most of the restof the megafauna, of the Americas.[86] In contrast, theSt. Paul Island mammoth population apparently died outprior to human arrival due to habitat shrinkage resultingfrom the post-ice age sea level rise.[86]

A 2008 study estimated that changes in climateshrank 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 mam-moths survived an even greater loss of habitat at the endof the Saale glaciation 125,000 years ago, and it is likely

6.7. FROZEN SPECIMENS 59

Cast of the “Hebior Mammoth” specimen, which bearstool/butcher marks

that humans hunted the remaining populations to extinc-tion at the end of the last glacial period.[89][90] Studiesof an 11,300–11,000 year old trackway in southwesternCanada showed that M. primigenius was in decline whilecoexisting with humans, since far fewer tracks of juve-niles were identified than would be expected in a normalherd.[46]

A 2010 study suggests that the decline of the woollymammoth could have increased temperatures by up to0.2 °C (0.36 °F) at high latitudes in the northern hemi-sphere. Mammoths frequently ate birch trees, creating agrassland habitat. With the disappearance of mammoths,birch forests, which absorb more sunlight than grasslands,expanded, leading to regional warming.[91]

6.7 Frozen specimens

Early 19th century interpretation of the "Adams mammoth" car-cass prior to excavation

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

fewer finds in the latter. Such remains are mostly foundabove the Arctic Circle, in permafrost. It appears thatsoft tissue was less likely to be preserved between 30,000and 15,000 years ago, perhaps because the climate wasmilder during that period. Most specimens have partiallydegraded prior to discovery, due to exposure or to be-ing scavenged. This "natural mummification" requiredthe animal to have been buried rapidly in liquid or semi-solids such as silt, mud and icy water, which then froze.[92]The presence of undigested food in the stomach and seedpods still in the mouth of many of the specimens sug-gests neither starvation nor exposure are likely. The ma-turity of this ingested vegetation places the time of deathin autumn rather than in spring, when flowers would beexpected.[93] The animals may have fallen through iceinto small ponds or potholes, entombing them. Manyare certainly known to have been killed in rivers, perhapsthrough being swept away by floods. In one location, bythe Berelekh River in Yakutia in Siberia, more than 8,000bones from at least 140 mammoths have been found ina single spot, apparently having been swept there by thecurrent.[94]

Illustration of the “Adams mammoth” skeleton with outwardcurving tusks, 1815

Between 1692 and 1806, only four descriptions of frozenmammoths were published in Europe. None of the re-mains of those five were preserved, and no completeskeleton was recovered during that time.[95] While frozenwoolly mammoth carcasses had been excavated by Euro-peans as early as 1728, the first fully documented spec-imen was discovered near the delta of the Lena Riverin 1799 by Ossip Schumachov, a Siberian hunter.[96]Schumachov let it thaw until he could retrieve the tusksfor sale to the ivory trade. While in Yakutsk in 1806,Michael Friedrich Adams heard about the frozen mam-moth. Upon arrival at the location, Adams discoveredthat wild animals had eaten most of the organs and fleshof the mammoth, including the trunk. He examined thecarcass and realised what was left would still be the mostcomplete mammoth recovered by that time. Adams re-covered the entire skeleton, apart from the tusks, whichShumachov had already sold, and one foreleg, most ofthe skin, and nearly 18 kg (40 lb) of hair. During his re-turn voyage he purchased a pair of tusks that he believed

60 CHAPTER 6. WOOLLY MAMMOTH

were the ones that Shumachov had sold. Adams broughtit to the Zoological Museum of the Zoological Institute ofthe Russian Academy of Sciences, and the task of mount-ing the skeleton was given toWilhelmGottlieb Tilesius.[5]The Kunstkamera, the museum established by Peter theGreat, contained the skeleton of an Indian elephant thatcould be used as reference.[97] This was one of the first at-tempts at reconstructing the skeleton of an extinct animal.Most of the reconstruction is correct, but Tilesius placedeach tusk in the opposite socket, so that they curved out-ward instead of inward. The error was not corrected until1899, and the correct placement of mammoth tusks wasstill a matter of debate into the twentieth century.[98][99]

A third of this model is covered with the skin of the “Berezovkamammoth”, Museum of Zoology, St. Petersburg

The 1901 excavation of the “Berezovka mammoth” is thebest documented of the early finds. It was discovered bythe Berezovka River, and the Russian authorities financedits excavation. Its head was exposed, and the flesh hadbeen scavenged. The animal still had grass between itsteeth and on the tongue, showing that it had died sud-denly. The entire expedition took 10 months, and thespecimen had to be cut to pieces before it could be trans-ported to St. Petersburg. It was identified as a 35–40year old male, which had died 35,000 years ago. One ofits shoulder blades was broken, whichmay have happenedwhen it fell into a crevasse.[93]

By 1929, the remains of 34 mammoths with frozen softtissues (skin, flesh, or organs) had been documented.Only four of them were relatively complete. Since then,about that manymore have been found. Inmost cases, theflesh showed signs of decay before its freezing and laterdesiccation.[100] Since 1860, Russian authorities have of-fered rewards of up to руб.1000 for finds of frozen woollymammoth carcasses. Often such finds were kept secretdue to superstition. Several carcasses have been lost be-cause they were not reported, and one was fed to dogs.In more recent years, scientific expeditions have beendevoted to finding carcasses instead of relying solely onchance encounters.[101]

In 1977, the well-preserved carcass of a seven- to eight-month-old woolly mammoth calf named “Dima” was dis-covered. This carcass was recovered near a tributary of

“Dima”, a frozen calf about seven months old

the Kolyma River in northeastern Siberia. This spec-imen weighed approximately 100 kg (220 lb) at deathand was 104 cm (41 in) high and 115 cm (45 in) long.Radiocarbon dating determined that “Dima” died about40,000 years ago. Its internal organs are similar to thoseof modern elephants, but its ears are only one-tenth thesize of those of an African elephant of similar age. A lesscomplete juvenile, nicknamed “Mascha”, was found onthe Yamal Peninsula in 1988. It was 3–4 months old, anda laceration on its right foot may have been the cause ofdeath. It is the westernmost frozen mammoth found.[102]

In 1997, a piece of mammoth tusk was discovered pro-truding from the tundra of the Taymyr Peninsula inSiberia, Russia. In 1999, this 20,380 year old carcass and25 tons of surrounding sediment were transported by anMi-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 thiscave began with the use of hairdryers to keep the hair andother 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 hadbeen buried for 41,800 years.[52][105] By cutting a sectionthrough a molar and analysing its growth lines, they foundthat the animal had died at the age of one month.[59] Themummified calf weighed 50 kg (110 lb), was 85 cm (33in) high and 130 cm (51 in) in length.[106][107] At the timeof discovery, its eyes and trunk were intact and some furremained on its body. Its organs and skin are very well

6.7. FROZEN SPECIMENS 61

preserved.[108] “Lyuba” is believed to have been suffo-cated by mud in a river that its herd was crossing.[52][109]After death, its body may have been colonised by bacte-ria that produce lactic acid, which “pickled” it, preservingthe mammoth in a nearly pristine state.[52]

Skull and mandible of the frozen calf “Yuka”

In 2012, a juvenile was found in Siberia, which had man-made cut marks. Scientists estimated its age at death tobe 2.5 years, and nicknamed it “Yuka”. Its skull andpelvis had been removed prior to discovery, but werefound nearby.[110][76] Another mammoth discovery wasreported in October 2012, when it was excavated on theTaymyr Peninsula. It was dated to 30,000 years old. Itwas named “Zhenya” after the boy who found it.[111]

In 2013, a well preserved carcass was found on MalyLyakhovsky Island, one of the islands in the New SiberianIslands archipelago, a female between 50 and 60 years oldat the time of death. The carcass containedwell preservedmuscular tissue. When it was extracted from the ice, liq-uid blood spilled from the abdominal cavity. The find-ers interpreted this as indicating woolly mammoth bloodpossessed anti-freezing properties.[112]

6.7.1 Recreating the species

The existence of preserved soft tissue remains and DNAof woolly mammoths has led to the idea that the speciescould be recreated by scientific means. Two methodshave been proposed to achieve this. The first is cloning,which would involve removal of the DNA-containingnucleus of the egg cell of a female elephant, and replace-ment with a nucleus from woolly mammoth tissue. Thecell would then be stimulated into dividing, and insertedback into a female elephant. The resulting calf wouldhave the genes of the woolly mammoth, although its fetalenvironment would be different. To date, even the mostintact mammoths have had little usable DNA because oftheir conditions of preservation. There is not enough toguide the production of an embryo.[113]

The second method involves artificially inseminating anelephant egg cell with sperm cells from a frozen woollymammoth carcass. The resulting offspring would be an

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

elephant–mammoth hybrid, and the process would haveto be repeated so more hybrids could be used in breeding.After several generations of cross-breeding these hybrids,an almost pure woollymammothwould be produced. Thefact that sperm cells of modern mammals are potent for15 years at most after deep-freezing is a hindrance tothis method.[113] In one case, an Asian elephant and anAfrican elephant produced a live calf named Motty, butit died of defects at less than two weeks old.[114]

In 2008, a Japanese team found usable DNA in thebrains of mice that had been frozen for 16 years. Theyhope to use similar methods to find usable mammothDNA.[115] In 2009, the Pyrenean Ibex (a subspecies ofthe Spanish ibex) was the first extinct animal to be clonedback to life; the clone lived for only seven minutes be-fore dying of lung defects.[116] As the woolly mammothgenome has been mapped, it may be possible to recre-ate a complete set of woolly mammoth chromosomes inthe future by adding mammoth-only sequences to ele-phant chromosomes.[117] If the process is ever successful,there are plans to introduce cloned woolly mammoths toPleistocene Park, a wildlife reserve in Siberia.[118]

Mammoth expert Adrian Lister questions the ethics ofsuch recreation attempts. In addition to the technicalproblems, he notes that there is not much habitat left thatwould be suitable for woolly mammoths. Because thespecies was social and gregarious, creating a few spec-imens would not be ideal. He also notes that the timeand resources required would be enormous, and that thescientific benefits would be unclear; he suggests these re-sources should instead be used to preserve extant elephantspecies which are endangered.[113] A 2014 article aboutpotential cloning also questioned the ethics of using ele-phants as surrogate mothers, as most embryos would notsurvive, and noted that it would be impossible to knowthe exact needs of a resurrected calf.[119]

62 CHAPTER 6. WOOLLY MAMMOTH

6.8 Cultural significance

Peter III of Russia carved in mammoth ivory

The woolly mammoth has remained culturally significantlong after its extinction. Indigenous peoples of Siberiahad long found what are now known to be woolly mam-moth remains, collecting their tusks for the ivory trade.Native Siberians believed woolly mammoth remains to bethose of giant mole-like animals that lived undergroundand died when burrowing to the surface.[6] Woolly mam-moth tusks had been articles of trade in Asia long be-fore Europeans became acquainted with them. Güyük,the 13th-century Khan of the Mongols, is reputed to havesat on a throne made from mammoth ivory.[95] Siberianmammoth ivory is reported to have been exported to Rus-sia and Europe in the 10th century. The first Siberianivory to reach western Europe was brought to London in1611.[120]

When Russia occupied Siberia, the ivory trade grewand became a widely exported commodity, with hugeamounts being excavated for this market. From the 19thcentury and onwards, woolly mammoth ivory became ahighly prized commodity, used as raw material for manydifferent products. Today it is still in great demand as areplacement for the now-banned export of elephant ivory,and has been referred to as “white gold”. Local dealersestimate that there are 10 million mammoths still frozenin Siberia, and conservationists have suggested that thiscould help save the living species of elephants from ex-tinction. Elephants are hunted by poachers for their ivory,but if this could instead be supplied by the already ex-tinct mammoths, the demand could instead be met bythese. Trade in elephant ivory has been forbidden inmost places following the 1989 LausanneConference, butdealers have been known to label it as mammoth ivory

to get it through customs. Mammoth ivory looks simi-lar to elephant ivory, but the former is browner and theSchreger lines are coarser in texture.[121] In the 21st cen-tury, global warming has made access to Siberian tuskseasier, since the permafrost thaws more quickly, expos-ing the mammoths embedded within it.[122]

Le Mammouth by Paul Jamin, 1885

Stories abound about frozen woolly mammoth meat thatwas consumed once defrosted, especially that of the“Berezovka mammoth”, but most of these are considereddubious. The carcasses were in most cases decayed, andthe stench so unbearable that only wild scavengers andthe dogs accompanying the finders showed any interestin the flesh. It appears that such meat was once recom-mended against illness in China, and Siberian natives haveoccasionally cooked the meat of frozen carcasses theydiscovered.[123]

6.8.1 Cryptozoology

There have been occasional claims that the woolly mam-moth is not extinct, and that small isolated herds mightsurvive in the vast and sparsely inhabited tundra of theNorthern Hemisphere. In the 19th century, several re-ports of “large shaggy beasts” were passed on to the Rus-sian authorities by Siberian tribesmen, but no scientificproof ever surfaced. A French chargé d'affaires work-ing in Vladivostok, M. Gallon, said in 1946 that in 1920he had met a Russian fur-trapper who claimed to haveseen living giant, furry “elephants” deep into the taiga.Gallon added that the fur-trapper had not heard of mam-moths before.[124] Due to the large area of Siberia, it can-

6.9. REFERENCES 63

not be completely ruled out that woolly mammoths sur-vived into more recent times, but all evidence indicatesthat they went extinct thousands of years ago. It is likelythat these natives had gained their knowledge of woollymammoths from carcasses they encountered, and that thisis the source for their legends of the animal.[125]

Legends from several Native American tribes have alsobeen interpreted by some as indicating folk memory ofextinct elephants.[126][127][128] In the late 19th century,there were persistent rumours about surviving mammothsin Alaska.[124] In October 1899, Henry Tukeman detailedhis killing of a mammoth in Alaska and his subsequentdonation of the specimen to the Smithsonian Institutionin Washington, D.C. The museum denied the existenceof any mammoth corpse; the story was a hoax.[129] BengtSjögren believed that the myth began when the Americanbiologist Charles Haskins Townsend travelled in Alaska,saw Eskimos trading mammoth tusks, asked if there stillwere livingmammoths in Alaska, and provided themwitha drawing of the animal.[124]

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[2] Switek, B. (2010). Written in Stone: Evolution, the FossilRecord, and Our Place in Nature. Bellevue Literary Press.pp. 174–180. ISBN 978-1-934137-29-1.

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[4] Sloane, H. (1727–1728). “Of Fossile Teeth and Bones ofElephants. Part the Second”. Philosophical Transactions35 (399–406): 497–514. Bibcode:1727RSPT...35..497S.doi:10.1098/rstl.1727.0048.

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[7] Cuvier, G. (1796). “Mémoire sur les épèces d'elephanstant vivantes que fossils, lu à la séance publique del'Institut National le 15 germinal, an IV”. Magasin en-cyclopédique, 2e anée (in French): 440–445.

[8] Reich, M.; Gehler, A. (2008). “Giants’ Bones and Uni-corn Horns Ice Age Elephants Offer 21st Century In-sights”. Collections - Wisdom, Insight, Innovation 8: 44–50.

[9] Brookes, J. (1828). A catalogue of the anatomical & zo-ological museum of Joshua Brookes 1. London: RichardTaylor. p. 73.

[10] “Mammoth entry in Oxford English Dictionary”. 2000.

[11] Lister, 2007. p. 49

[12] Simpson, J. (2009). "Word Stories: Mammoth.” OxfordEnglish Dictionary Online, Oxford University Press. Ac-cessed 5 June 2009.

[13] Lister, 2007. pp. 18–21

[14] Shoshani, J.; Tassy, P. (2005). “Advances in pro-boscidean taxonomy & classification, anatomy & phys-iology, and ecology & behavior”. Quaternary Inter-national. 126–128: 5. Bibcode:2005QuInt.126....5S.doi:10.1016/j.quaint.2004.04.011.

[15] Gross, L. (2006). “Reading the EvolutionaryHistory of the Woolly Mammoth in Its Mito-chondrial Genome”. PLoS Biology 4 (3): e74.doi:10.1371/journal.pbio.0040074. PMC 1360100.PMID 20076539.

[16] Krause, J.; Dear, P. H.; Pollack, J. L.; Slatkin, M.;Spriggs, H.; Barnes, I.; Lister, A. M.; Ebersberger, I.;Pääbo, S.; Hofreiter, M. (2005). “Multiplex amplifica-tion of the mammoth mitochondrial genome and the evo-lution of Elephantidae”. Nature 439 (7077): 724–727.doi:10.1038/nature04432. PMID 16362058.

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6.10 Bibliography• Data related to Mammuthus primigenius at Wik-ispecies

• Media related to Mammuthus primigenius at Wiki-media Commons

• Lister, A.; Bahn, P. (2007). Mammoths - Giants ofthe Ice Age (3 ed.). London: Frances Lincoln. ISBN978-0-520-26160-0.

Chapter 7

Woolly rhinoceros

The woolly rhinoceros (Coelodonta antiquitatis) is anextinct species of rhinoceros that was common through-out Europe and northern Asia[1] during the Pleistoceneepoch and survived the last glacial period. The genusname Coelodonta means “cavity tooth”. The woollyrhinoceros was a member of the Pleistocene megafauna.

7.1 Evolution

Chauvet cave art depicting a woolly rhino

As the last and most derived member of the Pleistocenerhinoceros lineage, the woolly rhinoceros was welladapted to its environment. Stocky limbs and thickwoolly pelage made it well suited to the steppe-tundraenvironment prevalent across the Palearctic ecozone dur-ing the Pleistocene glaciations. Like the vast majorityof rhinoceroses, the body plan of the woolly rhinocerosadhered to a conservative morphology, like the firstrhinoceroses seen in the late Eocene.A study of 40-70.000 year old DNA samples showed itsclosest extant relative is the Sumatran rhinoceros.[2]

7.2 Description

The external appearance of woolly rhinos is known frommummified individuals from Siberia as well as cavepaintings.[3] An adult woolly rhinoceros was typicallyaround 3 to 3.8 metres (10 to 12.5 feet) in length, with

Cast of the mummified Starunia specimen, Natural History Mu-seum, London

an estimated weight of around 2,721–3,175 kg (5,999–7,000 lb).[1] The woolly rhinoceros could grow to be 2m (6.6 ft) tall;[1] the body size was thus comparable, orslightly larger than, the extant White rhinoceros.[4] Twohorns on the skull were made of keratin, the anterior hornbeing 61 cm (24 in) in length,[5] with a smaller horn be-tween its eyes.[6] It had thick, long fur, small ears, short,thick legs, and a stocky body. Cave paintings suggesta wide dark band between the front and hind legs, butthe feature is not universal, and identification of picturedrhinoceroses as woolly rhinoceros is uncertain.Its shape was known only from prehistoric cave drawingsuntil a completely preserved specimen (missing only thefur and hooves) was discovered in a tar pit in Starunia,Poland. The specimen, an adult female, is now on displayin the Polish Academy of Sciences' Museum of NaturalHistory in Kraków. Several frozen specimens have alsobeen found in Siberia, the latest in 2007.[7]

7.3 Behavior and habitat

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

68

7.4. EXTINCTION 69

Restoration by Charles R. Knight

posed Doggerland and much of Northern Europe andwas common in the cold, arid desert that is southernEngland[9] and the North Sea today. Its geographi-cal range expanded and contracted with the alternatingcold and warm cycles, forcing populations to migrate asglaciers receded. The woolly rhinoceros co-existed withwoolly mammoths and several other extinct larger mam-mals of the Pleistocene megafauna. A close relative,Elasmotherium, had a more southern range.In 2011, the oldest known woolly rhinoceros fossil wasdiscovered from 3.6 million years in the Himalayas on thecold Tibetan Plateau, suggesting it existed there during aperiod of general climate warmth around the earth. It isbelieved that they migrated from there to northern Asiaand Europe when the Ice Age began.[10]

Females gave birth to one or two calves.[11]

7.3.1 Diet

Frozen head, leg and horn from Siberia, 1849

Controversy has long surrounded the precise dietary pref-erence of Coelodonta as past investigations have foundboth grazing and browsing modes of life to be plausi-ble. The palaeodiet of the woolly rhinoceros has been re-

constructed using several lines of evidence. Climatic re-constructions indicate the preferred environment to havebeen cold and arid steppe-tundra, with large herbivoresforming an important part of the feedback cycle. Pollenanalysis shows a prevalence of grasses and sedges withina more complicated vegetation mosaic.A strain vector biomechanical investigation of the skull,mandible and teeth of a well-preserved last cold stage in-dividual recovered from Whitemoor Haye, Staffordshire,revealed musculature and dental characteristics that sup-port a grazing feeding preference. In particular, the en-largement of the temporalis and neck muscles is consis-tent with that required to resist the large tugging forcesgenerated when taking large mouthfuls of fodder fromthe ground. The presence of a large diastema supportsthis theory.Comparisons with extant perissodactyls confirm thatCoelodonta was a hindgut fermentor with a single stom-ach, and as such would have grazed upon cellulose-rich,protein-poor fodder. This method of digestion wouldhave required a large throughput of food and thus linksthe large mouthful size to the low nutritive content of thechosen grasses and sedges.[12]

Recent evidence suggests that woolly rhinos alive in theArctic during the Last Glacial Maximum consumedapproximately equal volumes of forbs, such as Artemisia,and graminoids.[13]

7.4 Extinction

Main article: Pleistocene megafaunaMany species of Pleistocene megafauna, like the woolly

Woolly rhinoceros and other Ice Age mammals in late Pleistocenenorthern Spain, by Mauricio Antón

rhinoceros, became extinct around the same time period.Human and Neanderthal hunting is often cited as onecause.[14] Other theories for the cause of the extinctionsare climate change associated with the receding Ice ageand the hyperdisease hypothesis (q.v. Quaternary extinc-tion event).[15]

Recent radiocarbon dating indicates that populations sur-vived as recently as 8,000 BC in western Siberia. How-ever, the accuracy of this date is uncertain, as several ra-

70 CHAPTER 7. WOOLLY RHINOCEROS

diocarbon plateaus exist around this time. The extinctiondoes not coincide with the end of the last ice age but doescoincide with a minor yet severe climatic reversal thatlasted for about 1,000–1,250 years, the Younger Dryas(GS1 - Greenland Stadial 1), characterized by glacialreadvances and severe cooling globally, a brief interludein the continuing warming subsequent to the terminationof the last major ice age (GS2), thought to have been dueto a shutdown of the thermohaline circulation in the oceandue to huge influxes of cold fresh water from the preced-ing sustained glacial melting during the warmer Intersta-dial (GI1 - Greenland Interstadial 1 - ca. 16,000–11,45014C years B.P.).The Pinhole CaveMan is a late Paleolithic figure of amanengraved on a rib bone of the Woolly rhinoceros, foundat 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.; Hänni, C. (2003). “Ancient DNAanalysis reveals woolly rhino evolutionary relation-ships”. Molecular Phylogenetics and Evolution 28 (3):485–499. doi:10.1016/S1055-7903(03)00023-X. PMID12927133.

[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”. Thecomplete guide to prehistoric life (First ed.). Buffalo, N.Y.:Firefly Books. p. 203. ISBN 978-1-55407-181-4.

[6] Fortelius, Mikael (1983). “The morphology and pa-leobiological significance of the horns ofCoelodontaantiquitatis(Mammalia: Rhinocerotidae)". Jour-nal of Vertebrate Paleontology 3 (2): 125–135.doi:10.1080/02724634.1983.10011964. ISSN 0272-4634.

[7] Boeskorov, G. G. (2012). “Some specific mor-phological and ecological features of the fossilwoolly rhinoceros (Coelodonta antiquitatis Blumen-bach 1799)". Biology Bulletin 39 (8): 692–707.doi:10.1134/S106235901208002X.

[8] Jacobi, Roger M.; Rose, James; MacLeod, Alison;Higham, Thomas F.G. (2009). “Revised radiocar-bon ages on woolly rhinoceros (Coelodonta antiqui-tatis) from western central Scotland: significance fortiming the extinction of woolly rhinoceros in Britainand the onset of the LGM in central Scotland”.Quaternary Science Reviews 28 (25–26): 2551–2556.doi:10.1016/j.quascirev.2009.08.010. ISSN 0277-3791.

[9] Ian Rolfe, W. D. (1966). “Woolly rhinoceros from theScottish Pleistocene”. Scottish Journal of Geology 2 (3):253. doi:10.1144/sjg02030253.

[10] “Ice Age giants may have evolved in Tibet”. CNN. 1September 2011. Retrieved 2 September 2011.

[11] Walker, Matt (6 December 2012). “Prehistoric rhino re-veals secrets”. BBC News.

[12] SAS Bulletin, Volume 26, number 3/4, Winter 2003 fromthe Society for Archaeological Sciences

[13] Willerslev E, Davison J, Moora M, Zobel M, Coissac E,Edwards ME, Lorenzen ED, Vestergård M, GussarovaG, Haile J, Craine J, Gielly L, Boessenkool S, Epp LS,Pearman PB, Cheddadi R, Murray D, Bråthen KA, Yoc-coz N, Binney H, Cruaud C, Wincker P, Goslar T, Al-sos IG, Bellemain E, Brysting AK, Elven R, Sønstebø JH,Murton J, Sher A, Rasmussen M, Rønn R, Mourier T,Cooper A, Austin J, Möller P, Froese D, Zazula G, Pom-panon F, Rioux D, Niderkorn V, Tikhonov A, SavvinovG, Roberts RG, MacPhee RD, Gilbert MT, Kjær KH,Orlando L, Brochmann C, Taberlet P. (2014). “Fiftythousand years of Arctic vegetation and megafaunal diet”.Nature 506 (7486): 47–51. doi:10.1038/nature12921.PMID 24499916.

[14] Diamond, Jared (1997). Guns, Germs and Steel. NewYork: Vintage. ISBN 0-09-930278-0.

[15] Grayson, D. K.; Meltzer, D. J. (2003). “A requiemfor North American overkill”. Journal of Archaeo-logical Science 30 (5): 585–593. doi:10.1016/S0305-4403(02)00205-4.

[16] “engraved bone/antler”. British Museum.

• Parker, Steve. Dinosaurus: The Complete Guide toDinosaurs. Firefly Books Inc, 2003. Pg. 422.

7.7 External links• More pictures of the fully preserved tar pit whollyrhinoceros that was found in Poland (text in Polish)

• Fossil skull of a woolly rhinoceros from Belgium

• Fossil skull of a woolly rhinoceros from Germany

• International Rhino Foundation: Woolly Rhino

7.8. TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES 71

7.8 Text and image sources, contributors, and licenses

7.8.1 Text• Ice age Source: http://en.wikipedia.org/wiki/Ice%20age?oldid=637031735 Contributors: AxelBoldt, Paul Drye, Mav, Bryan Derksen, The

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 fjättrade ankan, Glenn, MichaelShields, Silthor, Nikai, Andres, Lee M, The Tom, Charles Matthews, Timwi, Steinsky, Hao2lian, Dragons flight, 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, Zig-ger, 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,TrevorMacInnis, Lacrimosus, Atrian, Rfl, 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 Ap-pleyard, 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, MG-Tom, Tckma, MONGO, Kelisi, Dzordzm, GregorB, Jugger90, Rotten, Prothonotar, Kmontgom, Marudubshinki, Dysepsion, Lusitana, Gra-ham87, 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, Spu-riousQ, 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,Allens, Bluezy, Jonathan.s.kt, Mjroots, Airconswitch, Mejor Los Indios, CIreland, Knowledgeum, Luk, SmackBot, Unschool, Haymaker,Smitz, Steve carlson, Reedy, Tarret, KnowledgeOfSelf, Ma8thew, Hydrogen Iodide, CyclePat, Prototime, Vald, Anastrophe, Frymaster,ZeroEgo, Kintetsubuffalo, Vodkasim, Xaosflux, Cool3, Skizzik, TRosenbaum, Squiddy, NormStephens, Saros136, Master Jay, RDBrown,Tito4000, Master of Puppets, Raymond arritt, Tasty monster, Miquonranger03, Los3, SchfiftyThree, Hibernian, Moshe Constantine Has-san Al-Silverburg, Bazonka, Y2kbird, Epastore, Darth Panda, Royboycrashfan, Squilibob, Jennica, Rsm99833, Japeo, Kcordina, Mr.Z-man, The tooth, Krich, Patrickbowman, Nakon, John D. 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dream of horses, Arctic Night, Hyattdj, Tom.Reding, Avidmosh, Tinton5, Île flottante, VinnyXY, Ice Age lover, Random-StringOfCharacters, 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, Kpuffer-

72 CHAPTER 7. WOOLLY RHINOCEROS

fish, Pditlev, Jakaloke, ScottyBerg, RA0808, Poopysnoop, Heljqfy, Beatleben5, Slightsmile, TuHan-Bot, Saiguy96, Emmick4, Mithrandi-rAgain, Alcart, Appaloosacakes, Xuiolhcarlos, 1234r00t, Lockon23, ZephyrWindOlympus, Frostyrollie, Wayne Slam, Getsagotta, Jay-Sebastos, 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, Mar-tialartistx, 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, Moon-dyne, 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, Smack-Bot, Francisco Valverde, Drn8, Chris the speller, Deanmo19, J. Spencer, Jwillbur, Adrian fine, PiMaster3, GumTree, Richard001, ThePIPE, Bcasterline, Soap, Tktktk, Mgiganteus1, A. Parrot, JHunterJ, Neddyseagoon, Arctic-Editor, Kurtle, Peter Horn, Scorpios, Irides-cent, Kaarel, J Milburn, CRGreathouse, Rwflammang, 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, Jeff G., Philip Trueman, TXiKiBoT, MrChupon, Ethel Aardvark, MeegsC, Theway, 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, Materialsci-entist, 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, PeterM. Brown, 1nv151b13.b0b, Ego White Tray, Orin04, ClueBot NG, Megaherbivore, Jonathan Fernando, Dream of Nyx, Oslep11, Help-ful 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, Prae-monitus, Fafnir1, Monkbot, Filedelinkerbot and Anonymous: 366

• Pleistocene Source: http://en.wikipedia.org/wiki/Pleistocene?oldid=636705203 Contributors: Bryan Derksen, Zundark, Berek, Andre En-gels, Vignaux, Rmhermen, Edward, Michael Hardy, Glenn, Susurrus, Evercat, Smack, Emperorbma, David Newton, Reddi, Tarosan,The Anomebot, Dragons flight, 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, Jeffrey 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, Anomalo-caris, Dysmorodrepanis, Patrick MMA Bringmans, SFC9394, Bota47, Botteville, Leptictidium, Poppy, Deville, Barryob, Pádraic MacUid-hir, 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, BenMoore, 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, An-tiVandalBot, Luna Santin, Seaphoto, Mikenorton, Deflective, Pedro, VoABot II, Fang 23, E104421, Warren Dew, Gwern, Ugajin, R'n'B,CommonsDelinker, J.delanoy, UBeR, KenSharp, ElinWhitneySmith, Crocadog, Johnbod, Austin512, Janet1983, Rosenknospe, Soczy-czi, Root7, STBotD, Tygrrr, Squids and Chips, VolkovBot, Thisisborin9, BlazeTheMovieFan, A4bot, Zybez, Cbrettin, Raven rs, UnaSmith, 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, Ice-man63976, 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, Fluffer-nutter, Ka Faraq Gatri, LaaknorBot, 37ophiuchi, AndersBot, Spike the Dingo, Tyw7, Tide rolls, Lightbot, ,ماني Solrac1993, Legobot,Luckas-bot, Yobot, Ptbotgourou, MTWEmperor, Shore3, KamikazeBot, Eric-Wester, AnomieBOT, Lebanonman19, Richardlord50, Hun-njazal, 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, ClueBotNG, 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, Davi-dLeighEllis, 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, Wave-length, 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, Tar-quin, 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

Mic, Ixfd64, Delirium, Paul A, Todd, Looxix, Den fjättrade ankan, Glenn, Marteau, Harry Potter, Mxn, Pizza Puzzle, Reddi, The Anome-bot, DJ Clayworth, Tpbradbury, Itai, Nv8200p, Phoebe, Penfold, Owen, Jni, Branddobbe, Robbot, RedWolf, Arkuat, Mayooranathan, Aca-demic Challenger, Ojigiri, Sunray, UtherSRG, Alan Liefting, Marc Venot, Gtrmp, Jyril, Tom harrison, Aphaia, Chowbok, Gadfium, Andy-cjp, DocSigma, Antandrus, Beland, Kusunose, Adamsan, Rdsmith4, Maximaximax, Bodnotbod, Icairns, Karl-Henner, J0m1eisler, Lu-midek, Neutrality, Okapi, Robin klein, Ratiocinate, Demiurge, Zondor, Trevor MacInnis, Eisnel, Mike Rosoft, EugeneZelenko, Discospin-ster, Rhobite, Mazi, Vsmith, Lulu of the Lotus-Eaters, Dbachmann, Grutter, Stbalbach, ESkog, Kbh3rd, Fenice, Eric Forste, Ben Webber,RoyBoy, Syats, Bobo192, Comtebenoit, Janna Isabot, Stesmo, Smalljim, Func, BrokenSegue, R. S. 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B. Mann, MacsBug, SmackBot,Haymaker, Vald, Bomac, Delldot, Yamaguchi , Gilliam, Kevinalewis, Rmosler2100, Wigren, Chris the speller, Tito4000, MK8, Snori,SchfiftyThree, Hongooi, Oatmeal batman, Scwlong, Javier Arambel, Can't sleep, clown will eat me, Sephiroth BCR, Yidisheryid, Rizzi,Rrburke, Addshore, KnowledgeLord, Smooth O, Underbar dk, Dreadstar, Yom, Evlekis, Kahuroa, Risker, Risssa, ArglebargleIV, Serein(renamed because of SUL), Rukario639, John, Euchiasmus, Scientizzle, Kipala, Heimstern, Thanos5150, Jimd, Jas131, IronGargoyle,Judge Howarth, A. 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DragonBot, Excirial, Eeekster, Jayantanth, Nownownow, Ember of Light,Tahmasp, Moberg, Thingg, Leungkh, Ranjithsutari, Versus22, Starlemusique, Johnuniq, SoxBot III, DumZiBoT, BarretB, ChickenFURY,BodhisattvaBot, Stickee, Rror, Shoeofdeathisadouche, Avoided, Skarebo, NellieBly, Badgernet, Noctibus, ZooFari, Bobliang345, StephenPoppitt, King Pickle, Addbot, ConCompS, Freakmighty, DOI bot, Tcncv, DaughterofSun, Astraydagger, Kristinamwood, Fieldday-sunday,Hoboday, CanadianLinuxUser, Fluffernutter, BabelStone, Download, EhsanQ, ChenzwBot, LinkFA-Bot, West.andrew.g, 5 albert square,Numbo3-bot, Tide rolls, Bluebusy, Albert galiza, Megaman enm, Legobot, PlankBot, Luckas-bot, Pink!Teen, Tohd8BohaithuGh1, Sprach-pfleger, Fraggle81, Ojay123, Gobbleswoggler, Timothyhouse1, Inemanja, IW.HG, Eric-Wester, JungleJoe, Magog the Ogre, Daveosaurus,AnomieBOT, Andrewrp, 1exec1, ThaddeusB, Jim1138, Piano non troppo, AdjustShift, Thewanger, Nick UA, Cillian flood, Csigabi, Flewis,Materialscientist, ImperatorExercitus, Citation bot, Gilderoy8, Nika 243, Bob Burkhardt, Yelloeyes, Maxis ftw, Frankenpuppy, Ober-sachsebot, MauritsBot, Xqbot, Timir2, Intelati, Cureden, Addihockey10, Capricorn42, Sungmanitu, Thermoproteus, Ultimation, J04n,GrouchoBot, Abce2, Brandon5485, Arch27, Doulos Christos, GhalyBot, Halubihalubi, Shadowjams, E0steven, Captain Weirdo the Great,Fortdj33, LucienBOT, Dger, Sebastiangarth, Bukovets, JohnL.Weber, Robo37, HamburgerRadio, Citation bot 1, Harleh, Pinethicket, Idream of horses, Haaqfun, Jonesey95, MJ94, Calmer Waters, Hoo man, Hantzen, SpaceFlight89, Île flottante, Ronald0216, VinnyXY,Newgrounder, Tim1357, , Abc518, Gamewizard71, FoxBot, TrickyM, TobeBot, Jonkerz, Comet Tuttle, Pclaplante, Kmw2700,Defender of torch, Cowlibob, Joodeak, Mttcmbs, Xrmach, Suffusion of Yellow, PleaseStand, DARTH SIDIOUS 2, Fellisha123, Bmath-ews96, Onel5969, TheRealSimmonds, RjwilmsiBot, TjBot, Shiftyfifty, Hajatvrc, DASHBot, EmausBot, John of Reading, 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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, Ubiq-uity, Jimfbleak, Raven in Orbit, Adam Bishop, Timwi, Tpbradbury, AnonMoos, Wetman, Twang, Stephan Schulz, Auric, UtherSRG,Michael Devore, Ezhiki, Mboverload, JoJan, DragonflySixtyseven, TJSwoboda, Discospinster, Rich Farmbrough, Rama, Vsmith, Wefa,Mikkel, Xezbeth, WegianWarrior, Bender235, Malkin, Bobo192, DCEdwards1966, Hesperian, Supersexyspacemonkey, Anthony Ap-pleyard, 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, Jameshfisher, Michael-ritchie200, DVdm, Royal Scottish, Sceptre, Hairy Dude, Rtkat3, Peter G Werner, RussBot, CambridgeBayWeather, Pseudomonas, Nawl-inWiki, 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, EnlilNinlil, KnowledgeOfSelf, Hydrogen Iodide, Pmaas, Jrockley, Unfinite, Drkarthi, Edgar181, Gaff, Ian Rose, Gilliam, Hmains, Chris thespeller, 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. 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• 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, Ben-der235, Kbh3rd, Leperflesh, Summer Song, Adambro, Jonathan Drain, Circeus, Supersexyspacemonkey, A2Kafir, 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, MoeEpsilon, 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, Mahar-ishi yogi, SieBot, BotMultichill, Orna82, FunkMonk, Helioseus, Graminophile, ClueBot, The Famous White Wolf, Hazabaza3, Trivialist,JeffreyW75, Bingodile, Abrech, Mammoth lover, Wkharrisjr, XLinkBot, Addbot, Ronhjones, MrOllie, Glane23, Lightbot, First Light, Zor-robot, , Luckas-bot, Yobot, Hohenloh, AnomieBOT, Jim1138, Graywords, Hunnjazal, ImperatorExercitus, Citation bot, ArthurBot,Xqbot, J04n, GrouchoBot, Brambleshire, Philip72, Jca123456, LucienBOT, Archaeodontosaurus, Pinethicket, Zvn, Gatormax15, Emaus-Bot, WikitanvirBot, Tolly4bolly, ChuispastonBot, ClueBot NG, Jnorton7558, Satellizer, John P. Harrison, Antiqueight, Helpful PixieBot, 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 Images• File:Abyssal_Brachiopod_00148.jpg Source: http://upload.wikimedia.org/wikipedia/commons/5/5b/Abyssal_Brachiopod_00148.jpg

License: Public domain Contributors: Own work Original artist: Myself• File:AntarcticaDomeCSnow.jpg Source: http://upload.wikimedia.org/wikipedia/commons/b/bd/AntarcticaDomeCSnow.jpg License:

CC-BY-2.5 Contributors: Own work Original artist: Stephen Hudson

7.8. TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES 75

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• File:Awashrivermap.png Source: http://upload.wikimedia.org/wikipedia/commons/9/94/Awashrivermap.png License: CC-BY-SA-3.0Contributors: Own work, Elevation data from SRTM, drainage basin from GTOPO [1], all other features from Vector Map. Rand McNally“New International Atlas” (1993) used as reference. Original artist: Kmusser

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• File:Distribution_of_woolly_mammoth.png Source: http://upload.wikimedia.org/wikipedia/commons/8/8b/Distribution_of_woolly_mammoth.png License: CC-BY-SA-3.0 Contributors: Own work based on: Diego J. Álvarez-Lao, Ralf-Dietrich Kahlke, Nuria García,and Dick Mol: The Padul mammoth finds — On the southernmost record of Mammuthus primigenius in Europe and its southern spreadduring the Late Pleistocene. Palaeogeography, Palaeoclimatology, Palaeoecology. 278, 2009, S. 57–70 Original artist: DagdaMor

• File:Dolmenmontebubbonia.jpg Source: http://upload.wikimedia.org/wikipedia/commons/d/dc/Dolmenmontebubbonia.jpg License:CC-BY-3.0 Contributors: Transferred from it.wikipedia; transfer was stated to be made by User:Memorato. Original artist: Originaluploader was Spiccolo at it.wikipedia

• File:EisrandlagenNorddeutschland.png Source: http://upload.wikimedia.org/wikipedia/commons/f/f8/EisrandlagenNorddeutschland.png License: Public domain Contributors: Originally from de.wikipedia; description page is/was here. Original artist: Original uploader wasBotaurus at de.wikipedia

• File:Elephant_near_ndutu.jpg Source: http://upload.wikimedia.org/wikipedia/commons/d/dc/Elephant_near_ndutu.jpg License: CC-BY-SA-2.0 Contributors: Originally from en.wikipedia; description page is/was here. Original artist: The author is nickandmel2006 onflickr

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• File:Folder_Hexagonal_Icon.svg Source: http://upload.wikimedia.org/wikipedia/en/4/48/Folder_Hexagonal_Icon.svg License: Cc-by-sa-3.0 Contributors: ? Original artist: ?

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• File:Frozen_Coelodonta.jpg Source: http://upload.wikimedia.org/wikipedia/commons/6/6d/Frozen_Coelodonta.jpg License: Pub-lic domain Contributors: http://books.google.dk/books?id=WHAhAQAAMAAJ&lpg=PA413&ots=4SVGCqW4qh&dq=speciminis+Wiluiensis&pg=PA408&redir_esc=y#v=onepage&q&f=false Original artist: Brandt’s artist.

• File:Ggantija_Temples_(1).jpg Source: http://upload.wikimedia.org/wikipedia/commons/1/12/Ggantija_Temples_%281%29.jpg Li-cense: CC-BY-SA-3.0 Contributors: Transferred from en.wikipedia; transferred to Commons by User:Dmitri Lytov using CommonsHelper.Original artist: Original uploader was Norum at en.wikipedia

• File:GlaciationsinEarthExistancelicenced_annotated.jpg Source: http://upload.wikimedia.org/wikipedia/commons/8/85/GlaciationsinEarthExistancelicenced_annotated.jpg License: CC-BY-SA-3.0 Contributors: Own work Original artist: William M.Connolley

76 CHAPTER 7. WOOLLY RHINOCEROS

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• File:MammothVsMastodon.jpg Source: http://upload.wikimedia.org/wikipedia/commons/0/0a/MammothVsMastodon.jpgLicense: CC-BY-SA-3.0 Contributors: Transferred from en.wikipedia, same as http://dantheman9758.deviantart.com/art/Mammoth-photoshop-breakdown-54022652 and http://dantheman9758.deviantart.com/art/Mammut-americanum-201147765 Originalartist: Original uploader was Dantheman9758 at en.wikipedia. “I created this image myself with Adobe Photoshop. I simply ask that youdo not drastically alter this image. There are no other available links to this image.”

• File:Mammoth_House_(Replica).JPG Source: http://upload.wikimedia.org/wikipedia/commons/3/3f/Mammoth_House_%28Replica%29.JPG License: CC-BY-SA-3.0 Contributors: Own work Original artist: Nandaro

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• File:Mammuthus_primigenius_fraasi.JPG Source: http://upload.wikimedia.org/wikipedia/commons/0/0b/Mammuthus_primigenius_fraasi.JPG License: CC-BY-SA-3.0 Contributors: Own work Original artist: Ghedoghedo

• File:Mammuthus_primigenius_infant_skeleton.JPG Source: http://upload.wikimedia.org/wikipedia/commons/7/79/Mammuthus_primigenius_infant_skeleton.JPG License: CC-BY-SA-3.0 Contributors: Own work Original artist: Ninjatacoshell

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• File:Peter_III_of_Russia_(mammuthus)2.jpg Source: http://upload.wikimedia.org/wikipedia/commons/f/f2/Peter_III_of_Russia_%28mammuthus%292.jpg License: CC-BY-SA-3.0 Contributors: Own work Original artist: shakko

• File:Pleistocene_SA.jpg Source: http://upload.wikimedia.org/wikipedia/commons/b/b8/Pleistocene_SA.jpg License: CC-BY-SA-3.0Contributors: Transferred from en.wikipedia Original artist: Original uploader was DiBgd at en.wikipedia

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• File:Raedera.png Source: http://upload.wikimedia.org/wikipedia/commons/e/ed/Raedera.png License: CC-BY-SA-2.5 Contributors:Own work Original artist: José-Manuel Benito

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• File:Siegsdorfer_Mammut.jpg Source: http://upload.wikimedia.org/wikipedia/commons/a/ad/Siegsdorfer_Mammut.jpg License: Pub-lic domain Contributors: Own work Original artist: Lou.gruber

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• File:Smithsonian_woolly_mammoth.jpg Source: http://upload.wikimedia.org/wikipedia/commons/6/61/Smithsonian_woolly_mammoth.jpg License: CC-BY-SA-2.0 Contributors: Smithsonian Museum of Natural History Original artist: Kevin Burkett fromPhiladelphia, Pa., USA

• File:Spear_thrower_carved_as_a_mammothDSCF6961.jpg Source: http://upload.wikimedia.org/wikipedia/commons/8/82/Spear_thrower_carved_as_a_mammothDSCF6961.jpg License: CC-BY-SA-3.0 Contributors: Own work Original artist: Johnbod

• File:SummerSolstice65N-future.png Source: http://upload.wikimedia.org/wikipedia/commons/9/92/SummerSolstice65N-future.pngLicense: Public domain Contributors: Own work Original artist: Incredio

• File:Venus_of_Brassempouy.jpg Source: http://upload.wikimedia.org/wikipedia/commons/6/6b/Venus_of_Brassempouy.jpg License:Public domain Contributors: Photograph : Jean-Gilles Berizzi. Upload : Elapied (talk · contribs) Original artist: ?

• File:Vostok_Petit_data.svg Source: http://upload.wikimedia.org/wikipedia/commons/b/b8/Vostok_Petit_data.svg License: CC-BY-SA-3.0 Contributors:

• Vostok-ice-core-petit.png Original artist: Vostok-ice-core-petit.png: NOAA• File:Wiki_letter_w_cropped.svg Source: http://upload.wikimedia.org/wikipedia/commons/1/1c/Wiki_letter_w_cropped.svg License:

CC-BY-SA-3.0 Contributors:

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Contributors:Original artist: Nicholas Moreau

• File:Woolly_mammoth.jpg Source: http://upload.wikimedia.org/wikipedia/commons/6/65/Woolly_mammoth.jpg License: CC-BY-SA-2.0 Contributors: Mammut Original artist: Flying Puffin

• File:Wooly_Mammoths.jpg Source: http://upload.wikimedia.org/wikipedia/commons/9/93/Wooly_Mammoths.jpg License: Public do-main Contributors: http://io9.com/5891441/celebrating-charles-r-knight-the-artist-who-first-brought-dinosaurs-and-megafauna-to-lifeOriginal artist: Charles R. Knight

• File:Wooly_Rhino_at_the_Natural_History_Museum.jpg Source: http://upload.wikimedia.org/wikipedia/commons/5/50/Wooly_Rhino_at_the_Natural_History_Museum.jpg License: CC-BY-2.0 Contributors: Wooly Rhino at the Natural History Museum Originalartist: Paul Hudson from United Kingdom

• File:Wooly_rhinoceros.jpg Source: http://upload.wikimedia.org/wikipedia/commons/d/d3/Wooly_rhinoceros.jpg License: Public do-main Contributors: http://archive.org/stream/menofoldstoneage00osborich#page/210/mode/2up Original artist: Charles R. Knight, H. Os-born, 'Men of the Old Stone Age'

• File:Yuka_(Mammuthus_primigenius)_-_Skull_and_Lower_Jaw_A.JPG Source: http://upload.wikimedia.org/wikipedia/commons/7/76/Yuka_%28Mammuthus_primigenius%29_-_Skull_and_Lower_Jaw_A.JPG License: CC-BY-SA-3.0 Contributors: Ownwork Original artist: Nandaro

• File:Yukagir_mammoth_head.jpg Source: http://upload.wikimedia.org/wikipedia/commons/1/17/Yukagir_mammoth_head.jpg Li-cense: CC-BY-2.0 Contributors: Flickr: Left side of skull 4.jpg Original artist: synchroswimr

• File:Yukagir_mammoth_leg.jpg Source: http://upload.wikimedia.org/wikipedia/commons/d/d4/Yukagir_mammoth_leg.jpg License:CC-BY-2.0 Contributors: Flickr: Leg.jpg Original artist: synchroswimr

78 CHAPTER 7. WOOLLY RHINOCEROS

• File:Каменный_век_(1).jpg Source: http://upload.wikimedia.org/wikipedia/commons/f/fa/%D0%9A%D0%B0%D0%BC%D0%B5%D0%BD%D0%BD%D1%8B%D0%B9_%D0%B2%D0%B5%D0%BA_%281%29.jpg License: Public domain Contributors:http://www.picture.art-catalog.ru/picture.php?id_picture=3316 Original artist: Viktor M. Vasnetsov

• File:Мамонт.jpg Source: http://upload.wikimedia.org/wikipedia/commons/f/f4/%D0%9C%D0%B0%D0%BC%D0%BE%D0%BD%D1%82.jpg License: CC-BY-SA-3.0 Contributors: Own work Original artist: This picture has been taken by Andrew Butko. Contact e-mail: abutko@gmail.com. Do not copy this image illegally by ignoring the terms of the СС-BY-SA or GNU FDL licenses, as it is not inthe public domain. Other photos see here.

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