Ice Age Megafauna and Time Notes Contents

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Ice Age megafauna and time notes Contents

1 Ice age 1 1.1 Origin of ice age theory ...... 1 1.2 Evidence for ice ages ...... 2 1.3 Major ice ages ...... 3 1.4 Glacials and interglacials ...... 4 1.5 Positive and negative feedback in glacial periods ...... 5 1.5.1 Positive feedback processes ...... 5 1.5.2 Negative feedback processes ...... 5 1.6 Causes of ice ages ...... 5 1.6.1 Changes in Earth’s atmosphere ...... 6 1.6.2 Position of the continents ...... 6 1.6.3 Fluctuations in ocean currents ...... 7 1.6.4 Uplift of the Tibetan plateau and surrounding mountain areas above the snowline ...... 7 1.6.5 Variations in Earth’s orbit (Milankovitch cycles) ...... 7 1.6.6 Variations in the Sun’s energy output ...... 8 1.6.7 Volcanism ...... 8 1.7 Recent glacial and interglacial phases ...... 8 1.7.1 Glacial stages in North America ...... 8 1.7.2 Last Glacial Period in the semiarid Andes around Aconcagua and Tupungato ...... 9 1.8 Eﬀects of glaciation ...... 9 1.9 See also ...... 10 1.10 References ...... 10 1.11 External links ...... 13

2 Megafauna 14 2.1 Ecological strategy ...... 14 2.2 Evolution of large body size ...... 14 2.2.1 In terrestrial mammals ...... 15 2.2.2 In marine mammals ...... 15 2.2.3 In ﬂightless birds ...... 15 2.3 Megafaunal mass extinctions ...... 16 2.3.1 Timing and possible causes ...... 16 2.3.2 Consequences of depletion of megafauna ...... 17

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2.4 Examples ...... 17 2.5 Gallery ...... 21 2.5.1 Extinct ...... 21 2.5.2 Living ...... 22 2.6 See also ...... 23 2.7 Notes ...... 23 2.8 References ...... 23 2.9 External links ...... 26

3 Pleistocene 27 3.1 Dating ...... 27 3.2 Paleogeography and climate ...... 28 3.2.1 Glacial features ...... 28 3.2.2 Major events ...... 28 3.2.3 Palaeocycles ...... 29 3.3 Fauna ...... 30 3.4 Humans ...... 30 3.5 Deposits ...... 31 3.6 See also ...... 31 3.7 References ...... 31 3.8 External links ...... 32

4 Prehistoric mammal 33 4.1 List of prehistoric mammals ...... 33 4.2 See also ...... 33

5 Stone Age 34 5.1 Historical signiﬁcance ...... 34 5.2 The Stone Age in archaeology ...... 34 5.2.1 Beginning of the Stone Age ...... 34 5.2.2 End of the Stone Age ...... 35 5.2.3 The concept of Stone Age ...... 35 5.2.4 The three-stage system ...... 36 5.2.5 The problem of the transitions ...... 37 5.3 Chronology ...... 37 5.3.1 Three-age chronology ...... 37 5.3.2 Three-stage chronology ...... 43 5.4 Material culture ...... 43 5.4.1 Tools ...... 43 5.4.2 Food and drink ...... 43 5.4.3 Shelter and habitat ...... 44 5.4.4 Art ...... 44 CONTENTS iii

5.4.5 Stone Age rituals and beliefs ...... 45 5.5 Modern popular culture and the Stone Age ...... 45 5.6 See also ...... 45 5.7 Notes ...... 46 5.8 References ...... 47 5.9 Further reading ...... 47 5.10 External links ...... 47

6 Woolly mammoth 49 6.1 Taxonomy ...... 49 6.1.1 Etymology ...... 50 6.1.2 Evolution ...... 50 6.2 Description ...... 51 6.2.1 Coat ...... 52 6.2.2 Dentition ...... 53 6.3 Palaeobiology ...... 54 6.3.1 Diet ...... 54 6.3.2 Growth and reproduction ...... 55 6.4 Distribution and habitat ...... 56 6.5 Relationship with humans ...... 57 6.5.1 Exploitation ...... 57 6.6 Extinction ...... 58 6.7 Frozen specimens ...... 59 6.7.1 Recreating the species ...... 61 6.8 Cultural signiﬁcance ...... 62 6.8.1 Cryptozoology ...... 62 6.9 References ...... 63 6.10 Bibliography ...... 67

7 Woolly rhinoceros 68 7.1 Evolution ...... 68 7.2 Description ...... 68 7.3 Behavior and habitat ...... 68 7.3.1 Diet ...... 69 7.4 Extinction ...... 69 7.5 See also ...... 70 7.6 References ...... 70 7.7 External links ...... 70 7.8 Text and image sources, contributors, and licenses ...... 71 7.8.1 Text ...... 71 7.8.2 Images ...... 74 7.8.3 Content license ...... 78 Chapter 1

Ice age

4 Temperature variation (ΔT) This article is about a generic geological period of tem- 2 0 perature reduction. For the most recent glacial period −2 °C −4 commonly referred to as the Ice Age, see Last glacial pe- −6 −8 riod. For other uses, see Ice age (disambiguation). −10 400 350 300 250 200 150 100 50 0 An ice age is a period of long-term reduction in the 300 Carbon Dioxide 280

260

240 ppmv 220

200

180 400 350 300 250 200 150 100 50 0

1.8 1.6 Dust concentration 1.4 1.2 1

ppm 0.8 0.6 0.4 0.2 0 400 350 300 250 200 150 100 50 0 Thousands of years ago

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

temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Within a long-term ice age, individual pulses of cold climate are termed "glacial periods" (or alternatively “glacials” or “glaciations” or col- loquially as “ice age”), and intermittent warm periods are called "interglacials". Glaciologically, ice age implies the presence of extensive ice sheets in the northern and south- An artist’s impression of ice age Earth at glacial maximum. ern hemispheres.[1] By this deﬁnition, we are in an inter- Based on: Crowley, T.J. (1995). “Ice age terrestrial carbon glacial period—the holocene—of the ice age that began changes revisited”. Global Biogeochemical Cycles 9 (3): 377– 2.6 million years ago at the start of the Pleistocene epoch, 389. Bibcode:1995GBioC...9..377C. doi:10.1029/95GB01107. because the Greenland, Arctic, and Antarctic ice sheets still exist.[2]

1.1 Origin of ice age theory

In 1742 Pierre Martel (1706–1767), an engineer and ge- ographer living in Geneva, visited the valley of Chamonix in the Alps of Savoy.[3][4] Two years later he published an account of his journey. He reported that the inhab- itants of that valley attributed the dispersal of erratic boulders to the fact that the glaciers had once extended much farther.[5][6] Later similar explanations were reported from other regions of the Alps. In 1815 the carpenter and chamois hunter Jean-Pierre Perraudin (1767– 1858) explained erratic boulders in the Val de Bagnes in The Antarctic ice sheet. Ice sheets expand during an ice age. the Swiss canton of Valais as being due to glaciers previ-

1 2 CHAPTER 1. ICE AGE

ously extending further.[7] An unknown woodcutter from pentier presented his paper before the Schweizerische Meiringen in the Bernese Oberland advocated a simi- Naturforschende Gesellschaft.[20] In the meantime, the lar idea in a discussion with the Swiss-German geologist German botanist Karl Friedrich Schimper (1803–1867) Jean de Charpentier (1786–1855) in 1834.[8] Compara- was studying mosses which were growing on erratic boul- ble explanations are also known from the Val de Ferret ders in the alpine upland of Bavaria. He began to wonder in the Valais and the Seeland in western Switzerland[9] where such masses of stone had come from. During the and in Goethe's Scientific Work.[10] Such explanations summer of 1835 he made some excursions to the Bavar- could also be found in other parts of the world. When the ian Alps. Schimper came to the conclusion that ice must Bavarian naturalist Ernst von Bibra (1806–1878) visited have been the means of transport for the boulders in the the Chilean Andes in 1849–1850 the natives attributed alpine upland. In the winter of 1835 to 1836 he held fossil moraines to the former action of glaciers.[11] some lectures in Munich. Schimper then assumed that Meanwhile, European scholars had begun to wonder what there must have been global times of obliteration (“Verö- dungszeiten”) with a cold climate and frozen water.[21] had caused the dispersal of erratic material. From the middle of the 18th century some discussed ice as a means Schimper spent the summer months of 1836 at Devens, of transport. The Swedish mining expert Daniel Tilas near Bex, in the Swiss Alps with his former university (1712–1772) was, in 1742, the first person to suggest friend Louis Agassiz (1801–1873) and Jean de Charp- drifting sea ice in order to explain the presence of erratic entier. Schimper, de Charpentier and possibly Venetz boulders in the Scandinavian and Baltic regions.[12] In convinced Agassiz that there had been a time of glacia- 1795, the Scottish philosopher and gentleman naturalist, tion. During Winter 1836/7 Agassiz and Schimper devel- James Hutton (1726–1797), explained erratic boulders in oped the theory of a sequence of glaciations. They mainly the Alps with the action of glaciers.[13] Two decades later, drew upon the preceding works of Venetz, de Charpen- in 1818, the Swedish botanist Göran Wahlenberg (1780– tier and on their own fieldwork. There are indications that 1851) published his theory of a glaciation of the Scandi- Agassiz was already familiar with Bernhardi’s paper at that time.[22] At the beginning of 1837 Schimper coined navian peninsula. He regarded glaciation as a regional [23] phenomenon.[14] Only a few years later, the Danish- the term ice age (“Eiszeit”). In July 1837 Agassiz pre- Norwegian Geologist Jens Esmark (1762–1839) argued sented their synthesis before the annual meeting of the a sequence of worldwide ice ages. In a paper published in Schweizerische Naturforschende Gesellschaft at Neuchâ- 1824, Esmark proposed changes in climate as the cause tel. The audience was very critical or even opposed the of those glaciations. He attempted to show that they orig- new theory because it contradicted the established opin- inated from changes in Earth’s orbit.[15] During the fol- ions on climatic history. Most contemporary scientists thought that the earth had been gradually cooling down lowing years, Esmark’s ideas were discussed and taken [24] over in parts by Swedish, Scottish and German scien- since its birth as a molten globe. tists. At the University of Edinburgh Robert Jameson In order to overcome this rejection, Agassiz embarked (1774–1854) seemed to be relatively open to Esmark’s on geological fieldwork. He published his book Study on ideas, as reviewed by Norwegian professor of glaciol- glaciers ("Études sur les glaciers”) in 1840.[25] De Char- ogy Bjørn G. Andersen (1992).[16] Jameson’s remarks pentier was put out by this as he had also been preparing about ancient glaciers in Scotland were most probably a book about the glaciation of the Alps. De Charpentier prompted by Esmark.[17] In Germany, Albrecht Rein- felt that Agassiz should have given him precedence as it hard Bernhardi (1797–1849), a geologist and professor was he who had introduced Agassiz to in-depth glacial of forestry at an academy in Dreissigacker, since incor- research.[26] Besides that, Agassiz had, as a result of per- porated in the southern Thuringian city of Meiningen, sonal quarrels, omitted any mention of Schimper in his adopted Esmark’s theory. In a paper published in 1832, book.[27] Bernhardi speculated about former polar ice caps reach- [18] All together, it took several decades until the ice age the- ing as far as the temperate zones of the globe. ory was fully accepted. This happened on an international Independently of these debates, the Swiss civil engineer scale in the second half of the 1870s following the work Ignaz Venetz (1788–1859) in 1829, explained the dis- of James Croll including the publication of Climate and persal of erratic boulders in the Alps, the nearby Jura Time, in Their Geological Relations in 1875 which pro- Mountains and the North German Plain as being due vided a credible explanation for the causes of ice ages.[28] to huge glaciers. When he read his paper before the Schweizerische Naturforschende Gesellschaft, most scientists remained sceptical.[19] Finally, Venetz managed to convince his friend Jean de Charpentier. De Char- 1.2 Evidence for ice ages pentier transformed Venetz’s idea into a theory with a glaciation limited to the Alps. His thoughts resembled There are three main types of evidence for ice ages: ge- Wahlenberg’s theory. In fact, both men shared the same ological, chemical, and paleontological. volcanistic, or in de Charpentier’s case rather plutonistic assumptions, about the earth’s history. In 1834, de Char- Geological evidence for ice ages comes in various forms, including rock scouring and scratching, glacial moraines, 1.3. MAJOR ICE AGES 3 drumlins, valley cutting, and the deposition of till or tillites and glacial erratics. Successive glaciations tend to distort and erase the geological evidence, making it difficult to interpret. Furthermore, this evidence was difficult to date exactly; early theories assumed that the glacials were short compared to the long interglacials. The advent of sediment and ice cores revealed the true situation: glacials are long, interglacials short. It took some time for the current theory to be worked out. The chemical evidence mainly consists of variations in the ratios of isotopes in fossils present in sediments and sedimentary rocks and ocean sediment cores. For the most recent glacial periods ice cores provide climate proxies from their ice, and atmospheric samples from included bubbles of air. Because water containing heavier isotopes has a higher heat of evaporation, its proportion Ice age map of northern Germany and its northern neighbours. decreases with colder conditions.[29] This allows a tem- Red: maximum limit of Weichselian glacial; yellow: Saale perature record to be constructed. However, this evi- glacial at maximum (Drenthe stage); blue: Elster glacial max- dence can be confounded by other factors recorded by imum glaciation. isotope ratios. The paleontological evidence consists of changes in the geographical distribution of fossils. During a glacial period cold-adapted organisms spread into lower latitudes, Timeline of glaciations, shown in blue. and organisms that prefer warmer conditions become extinct or are squeezed into lower latitudes. This evidence is also difficult to interpret because it requires (1) sequences tending from near Sault Ste. Marie to Sudbury, north- of sediments covering a long period of time, over a wide east of Lake Huron, with giant layers of now-lithified till range of latitudes and which are easily correlated; (2) an- beds, dropstones, varves, outwash, and scoured basement cient organisms which survive for several million years rocks. Correlative Huronian deposits have been found without change and whose temperature preferences are near Marquette, Michigan, and correlation has been made easily diagnosed; and (3) the finding of the relevant fos- with Paleoproterozoic glacial deposits from Western Aus- sils. tralia. Despite the difficulties, analysis of ice core and ocean The next well-documented ice age, and probably the most sediment cores[30] has shown periods of glacials and in- severe of the last billion years, occurred from 850 to 630 terglacials over the past few million years. These also million years ago (the Cryogenian period) and may have confirm the linkage between ice ages and continental produced a Snowball Earth in which glacial ice sheets crust phenomena such as glacial moraines, drumlins, and reached the equator,[33] possibly being ended by the ac- glacial erratics. Hence the continental crust phenomena cumulation of greenhouse gases such as CO are accepted as good evidence of earlier ice ages when 2 produced by volcanoes. “The presence of ice on the they are found in layers created much earlier than the time continents and pack ice on the oceans would inhibit both range for which ice cores and ocean sediment cores are silicate weathering and photosynthesis, which are the two available. major sinks for CO 2 at present.”[34] It has been suggested that the end of this ice age was responsible for the subsequent Ediacaran and 1.3 Major ice ages Cambrian Explosion, though this model is recent and controversial. There have been at least five major ice ages in the earth’s The Andean-Saharan occurred from 460 to 420 million past (the Huronian, Cryogenian, Andean-Saharan, Karoo years ago, during the Late Ordovician and the Silurian Ice Age and the Quaternary glaciation). Outside these period. ages, the Earth seems to have been ice-free even in high [31][32] The evolution of land plants at the onset of the Devonian latitudes. period caused a long term increase in planetary oxygen Rocks from the earliest well established ice age, called levels and reduction of CO the Huronian, formed around 2.4 to 2.1 Ga (billion years) 2 levels, which resulted in the Karoo Ice Age. It is named ago during the early Proterozoic Eon. Several hundreds after the glacial tills found in the Karoo region of South of km of the Huronian Supergroup are exposed 10– Africa, where evidence for this ice age was first clearly 100 km north of the north shore of Lake Huron ex- identified. There were extensive polar ice caps at intervals 4 CHAPTER 1. ICE AGE

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

2 2 41 kyr cycle 100 kyr cycle 0 2.5

-2 3 -4 Five Million Years of 3.5 Equivalent 18

-6 Climate Change δ O Benthic Vostok Δ T (°C) 4 -8 From Sediment Cores Carbonate (per mil) 4.5 Millions of Years Ago

Sediment records showing the fluctuating sequences of glacials Shows the pattern of temperature and ice volume changes asso- and interglacials during the last several million years. ciated with recent glacials and interglacials The current ice age, the Pliocene-Quaternary glaciation, started about 2.58 million years ago during the late Pliocene, when the spread of ice sheets in the Northern Hemisphere began. Since then, the world has seen cycles of glaciation with ice sheets advancing and retreating on 40,000- and 100,000-year time scales called glacial periods, glacials or glacial advances, and interglacial periods, interglacials or glacial retreats. The earth is currently in an interglacial, and the last glacial period ended about 10,000 years ago. All that remains of the continental ice sheets are the Greenland and Antarctic ice sheets and smaller glaciers such as on Baffin Island. Ice ages can be further divided by location and time; for example, the names Riss (180,000–130,000 years bp) and Würm (70,000–10,000 years bp) refer specifically to glaciation in the Alpine region. The maximum extent of the ice is not maintained for the full interval. The scouring action of each glaciation tends to remove most of the evidence of prior ice sheets almost completely, except in regions where the later sheet does not achieve full cover- age. Minimum (interglacial, black) and maximum (glacial, grey) glaciation of the northern hemisphere 1.4 Glacials and interglacials

See also: Glacial period and Interglacial Within the ice ages (or at least within the current one), more temperate and more severe periods occur. The changes in Earth’s orbit called Milankovitch cycles. colder periods are called glacial periods, the warmer pe- The earth has been in an interglacial period known as riods interglacials, such as the Eemian Stage. the Holocene for more than 11,000 years. It was con- Glacials are characterized by cooler and drier climates ventional wisdom that the typical interglacial period lasts over most of the earth and large land and sea ice masses about 12,000 years, but this has been called into ques- extending outward from the poles. Mountain glaciers in tion recently. For example, an article in Nature[35] ar- otherwise unglaciated areas extend to lower elevations gues that the current interglacial might be most analo- due to a lower snow line. Sea levels drop due to the re- gous to a previous interglacial that lasted 28,000 years. moval of large volumes of water above sea level in the Predicted changes in orbital forcing suggest that the next icecaps. There is evidence that ocean circulation patterns glacial period would begin at least 50,000 years from now, are disrupted by glaciations. Since the earth has signiﬁ- even in absence of human-made global warming[36] (see cant continental glaciation in the Arctic and Antarctic, we Milankovitch cycles). Moreover, anthropogenic forc- are currently in a glacial minimum of a glaciation. Such ing from increased greenhouse gases might outweigh or- a period between glacial maxima is known as an inter- bital forcing for as long as intensive use of fossil fuels glacial. The glacials and interglacials also coincided with continues.[37] 1.6. CAUSES OF ICE AGES 5

rent projected consequences of global warming include a largely ice-free Arctic Ocean within 5–20 years, see Arctic shrinkage.) Additional fresh water flowing into the North Atlantic during a warming cycle may also reduce the global ocean water circulation (see Shutdown of thermohaline circulation). Such a reduction (by reducing the effects of the Gulf Stream) would have a cooling effect on northern Europe, which in turn would lead to increased low-latitude snow retention during the summer. It has also been suggested that during an extensive glacial, glaciers may move through the Gulf of Saint Lawrence, extending into the North Atlantic ocean far enough to block the Gulf Stream.

1.5.2 Negative feedback processes

Ice sheets that form during glaciations cause erosion of the land beneath them. After some time, this will re- Minimum (interglacial, black) and maximum (glacial, grey) duce land above sea level and thus diminish the amount glaciation of the southern hemisphere of space on which ice sheets can form. This mitigates the albedo feedback, as does the lowering in sea level that 1.5 Positive and negative feedback accompanies the formation of ice sheets. in glacial periods Another factor is the increased aridity occurring with glacial maxima, which reduces the precipitation available to maintain glaciation. The glacial retreat induced by this Each glacial period is subject to positive feedback which or any other process can be amplified by similar inverse makes it more severe and negative feedback which miti- positive feedbacks as for glacial advances. gates and (in all cases so far) eventually ends it. According to research published in Nature Geoscience, human emissions of carbon dioxide will defer the next ice 1.5.1 Positive feedback processes age. Researchers used data on Earth’s orbit to find the historical warm interglacial period that looks most like the current one and from this have predicted that the next ice Ice and snow increase Earth’s albedo, i.e. they make it age would usually begin within 1,500 years. They go on reflect more of the sun’s energy and absorb less. Hence, to say that emissions have been so high that it will not.[39] when the air temperature decreases, ice and snow fields grow, and this continues until competition with a negative feedback mechanism forces the system to an equilibrium. Also, the reduction in forests caused by the ice’s expan- 1.6 Causes of ice ages sion increases albedo. Another theory proposed by Ewing and Donn in 1956[38] The causes of ice ages are not fully understood for either hypothesized that an ice-free Arctic Ocean leads to in- the large-scale ice age periods or the smaller ebb and flow creased snowfall at high latitudes. When low-temperature of glacial–interglacial periods within an ice age. The con- ice covers the Arctic Ocean there is little evaporation or sensus is that several factors are important: atmospheric sublimation and the polar regions are quite dry in terms composition, such as the concentrations of carbon diox- of precipitation, comparable to the amount found in ide and methane (the specific levels of the previously mid-latitude deserts. This low precipitation allows high- mentioned gases are now able to be seen with the new ice core samples from EPICA Dome C in Antarctica over latitude snowfalls to melt during the summer. An ice- [40] free Arctic Ocean absorbs solar radiation during the long the past 800,000 years ); changes in the earth’s orbit summer days, and evaporates more water into the Arctic around the Sun known as Milankovitch cycles; the mo- atmosphere. With higher precipitation, portions of this tion of tectonic plates resulting in changes in the rela- snow may not melt during the summer and so glacial ice tive location and amount of continental and oceanic crust can form at lower altitudes and more southerly latitudes, on the earth’s surface, which affect wind and ocean cur- reducing the temperatures over land by increased albedo rents; variations in solar output; the orbital dynamics as noted above. Furthermore, under this hypothesis the of the Earth-Moon system; and the impact of relatively lack of oceanic pack ice allows increased exchange of wa- large meteorites, and volcanism including eruptions of ters between the Arctic and the North Atlantic Oceans, supervolcanoes. warming the Arctic and cooling the North Atlantic. (Cur- Some of these factors influence each other. For exam- 6 CHAPTER 1. ICE AGE

ple, changes in Earth’s atmospheric composition (espe- to have a significant global impact on the earth’s climate cially the concentrations of greenhouse gases) may alter and ecosystems, did not begin in the 18th century with the the climate, while climate change itself can change the advent of the Industrial Era, but dates back to 8,000 years atmospheric composition (for example by changing the ago, due to intense farming activities of our early agrarian rate at which weathering removes CO ancestors. It was at that time that atmospheric greenhouse 2). gas concentrations stopped following the periodic pattern Maureen Raymo, William Ruddiman and others propose of the Milankovitch cycles. In his overdue-glaciation hy- that the Tibetan and Colorado Plateaus are immense CO pothesis Ruddiman states that an incipient glacial would probably have begun several thousand years ago, but the 2 “scrubbers” with a capacity to remove enough CO 2 from the global atmosphere to be a significant causal arrival of that scheduled glacial was forestalled by the activities of early farmers.[45] factor of the 40 million year Cenozoic Cooling trend. They further claim that approximately half of their up- At a meeting of the American Geophysical Union (De- lift (and CO cember 17, 2008), scientists detailed evidence in sup- 2 “scrubbing” capacity) occurred in the past 10 million port of the controversial idea that the introduction of years.[41][42] large-scale rice agriculture in Asia, coupled with extensive deforestation in Europe began to alter world climate by pumping significant amounts of greenhouse gases 1.6.1 Changes in Earth’s atmosphere into the atmosphere over the last 1,000 years. In turn, a warmer atmosphere heated the oceans making them much less efficient storehouses of carbon dioxide and re- There is considerable evidence that over the very recent inforcing global warming, possibly forestalling the onset period of the last 100–1000 years, the sharp increases of a new glacial age.[46] in human activity, especially the burning of fossil fuels, has caused the parallel sharp and accelerating increase in atmospheric greenhouse gases which trap the sun’s heat. 1.6.2 Position of the continents The consensus theory of the scientific community is that the resulting greenhouse effect is a principal cause of The geological record appears to show that ice ages start the increase in global warming which has occurred over when the continents are in positions which block or re- the same period, and a chief contributor to the acceler- duce the flow of warm water from the equator to the poles ated melting of the remaining glaciers and polar ice.A and thus allow ice sheets to form. The ice sheets increase 2012 investigation finds that dinosaurs released methane Earth’s reflectivity and thus reduce the absorption of solar through digestion in a similar amount to humanity’s cur- radiation. With less radiation absorbed the atmosphere rent methane release, which “could have been a key fac- cools; the cooling allows the ice sheets to grow, which tor” to the very warm climate 150 million years ago.[43] further increases reflectivity in a positive feedback loop. There is evidence that greenhouse gas levels fell at the The ice age continues until the reduction in weathering start of ice ages and rose during the retreat of the ice causes an increase in the greenhouse effect. sheets, but it is difficult to establish cause and effect (see There are three known configurations of the continents the notes above on the role of weathering). Greenhouse which block or reduce the flow of warm water from the gas levels may also have been affected by other factors equator to the poles: which have been proposed as causes of ice ages, such as the movement of continents and volcanism. • A continent sits on top of a pole, as Antarctica does The Snowball Earth hypothesis maintains that the severe today. freezing in the late Proterozoic was ended by an increase in CO • A polar sea is almost land-locked, as the Arctic 2 levels in the atmosphere, and some supporters of Snow- Ocean is today. ball Earth argue that it was caused by a reduction in atmospheric CO • A supercontinent covers most of the equator, as 2. The hypothesis also warns of future Snowball Earths. Rodinia did during the Cryogenian period. In 2009, further evidence was provided that changes in solar insolation provide the initial trigger for the earth to Since today’s Earth has a continent over the South Pole warm after an Ice Age, with secondary factors like in- and an almost land-locked ocean over the North Pole, creases in greenhouse gases accounting for the magnitude geologists believe that Earth will continue to experience of the change.[44] glacial periods in the geologically near future. William Ruddiman has proposed the early anthropocene Some scientists believe that the Himalayas are a major hypothesis, according to which the anthropocene era, as factor in the current ice age, because these mountains some people call the most recent period in the earth’s his- have increased Earth’s total rainfall and therefore the rate tory when the activities of the human species first began at which carbon dioxide is washed out of the atmosphere, 1.6. CAUSES OF ICE AGES 7

decreasing the greenhouse effect.[42] The Himalayas’ for- year cycle of radiation changes due to variations in Earth’s mation started about 70 million years ago when the Indo- orbit. This comparatively insignificant warming, when Australian Plate collided with the Eurasian Plate, and the combined with the lowering of the Nordic inland ice areas Himalayas are still rising by about 5 mm per year because and Tibet due to the weight of the superimposed ice-load, the Indo-Australian plate is still moving at 67 mm/year. has led to the repeated complete thawing of the inland ice The history of the Himalayas broadly fits the long-term areas.[49][50][51][52] decrease in Earth’s average temperature since the mid- Eocene, 40 million years ago. 1.6.5 Variations in Earth’s orbit (Mi- lankovitch cycles) 1.6.3 Fluctuations in ocean currents The Milankovitch cycles are a set of cyclic variations in Another important contribution to ancient climate characteristics of the Earth’s orbit around the Sun. Each regimes is the variation of ocean currents, which are cycle has a different length, so at some times their effects modified by continent position, sea levels and salinity, reinforce each other and at other times they (partially) as well as other factors. They have the ability to cool cancel each other. (e.g. aiding the creation of Antarctic ice) and the ability to warm (e.g. giving the British Isles a temperate as opposed to a boreal climate). The closing of the Isthmus of Panama about 3 million years ago may have ushered in the present period of strong glaciation over North Amer- ica by ending the exchange of water between the tropical Atlantic and Pacific Oceans.[47] Past and future of daily average insolation at top of the atmosphere on the day of the summer solstice, at 65 N latitude. Analyses suggest that ocean current fluctuations can ade- quately account for recent glacial oscillations. During the There is strong evidence that the Milankovitch cycles last glacial period the sea-level has fluctuated 20–30 m as affect the occurrence of glacial and interglacial periods water was sequestered, primarily in the northern hemi- within an ice age. The present ice age is the most stud- sphere ice sheets. When ice collected and the sea level ied and best understood, particularly the last 400,000 dropped sufficiently, flow through the Bering Strait (the years, since this is the period covered by ice cores that narrow strait between Siberia and Alaska is ~50 m deep record atmospheric composition and proxies for temper- today) was reduced, resulting in increased flow from the ature and ice volume. Within this period, the match of North Atlantic. This realigned the thermohaline circu- glacial/interglacial frequencies to the Milanković orbital lation in the Atlantic, increasing heat transport into the forcing periods is so close that orbital forcing is gener- Arctic, which melted the polar ice accumulation and really accepted. The combined effects of the changing dis- duced other continental ice sheets. The release of water tance to the Sun, the precession of the Earth’s axis, and raised sea levels again, restoring the ingress of colder wa- the changing tilt of the Earth’s axis redistribute the sun- ter from the Pacific with an accompanying shift to north- light received by the Earth. Of particular importance are ern hemisphere ice accumulation.[48] changes 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 much 1.6.4 Uplift of the Tibetan plateau and sur- as 22% (from 450 W/m² to 550 W/m²). It is widely be- rounding mountain areas above the lieved that ice sheets advance when summers become too snowline cool to melt all of the accumulated snowfall from the previous winter. Some workers believe that the strength of Matthias Kuhle’s geological theory of Ice Age develop- the orbital forcing is too small to trigger glaciations, but ment was suggested by the existence of an ice sheet cover- feedback mechanisms like CO ing the Tibetan plateau during the Ice Ages (Last Glacial 2 may explain this mismatch. Maximum?). According to Kuhle, the plate-tectonic up- While Milankovitch forcing predicts that cyclic changes lift of Tibet past the snow-line has led to a surface of c. in the Earth’s orbital elements can be expressed in the 2,400,000 square kilometres (930,000 sq mi) changing glaciation record, additional explanations are necessary to from bare land to ice with a 70% greater albedo. The re- explain which cycles are observed to be most important flection of energy into space resulted in a global cooling, in the timing of glacial–interglacial periods. In partic- triggering the Pleistocene Ice Age. Because this highland ular, during the last 800,000 years, the dominant period is at a subtropical latitude, with 4 to 5 times the insolation of glacial–interglacial oscillation has been 100,000 years, of high-latitude areas, what would be Earth’s strongest which corresponds to changes in Earth’s orbital eccentric- heating surface has turned into a cooling surface. ity and orbital inclination. Yet this is by far the weakest Kuhle explains the interglacial periods by the 100,000- of the three frequencies predicted by Milankovitch. Dur- 8 CHAPTER 1. ICE AGE

ing the period 3.0–0.8 million years ago, the dominant which occurred during the coldest part of the Little pattern of glaciation corresponded to the 41,000-year pe- Ice Age. riod of changes in Earth’s obliquity (tilt of the axis). The reasons for dominance of one frequency versus another The long-term increase in the Sun’s output cannot be a are poorly understood and an active area of current re- cause of ice ages. search, but the answer probably relates to some form of resonance in the Earth’s climate system. 1.6.7 Volcanism The “traditional” Milankovitch explanation struggles to explain the dominance of the 100,000-year cycle over the Volcanic eruptions may have contributed to the incep- last 8 cycles. Richard A. Muller, Gordon J. F. MacDon- tion and/or the end of ice age periods. At times dur- ald,[53][54][55] and others have pointed out that those cal- ing the paleoclimate, carbon dioxide levels were two or culations are for a two-dimensional orbit of Earth but the three times greater than today. Volcanoes and move- three-dimensional orbit also has a 100,000-year cycle of ments in continental plates contributed to high amounts orbital inclination. They proposed that these variations of CO2 in the atmosphere. Carbon dioxide from volca- in orbital inclination lead to variations in insolation, as noes probably contributed to periods with highest over- the Earth moves in and out of known dust bands in the all temperatures.[60] One suggested explanation of the solar system. Although this is a diﬀerent mechanism to Paleocene-Eocene Thermal Maximum is that undersea the traditional view, the “predicted” periods over the last volcanoes released methane from clathrates and thus 400,000 years are nearly the same. The Muller and Mac- caused a large and rapid increase in the greenhouse ef- Donald theory, in turn, has been challenged by Jose An- fect. There appears to be no geological evidence for such tonio Rial.[56] eruptions at the right time, but this does not prove they Another worker, William Ruddiman, has suggested did not happen. a model that explains the 100,000-year cycle by the modulating eﬀect of eccentricity (weak 100,000-year cycle) on precession (26,000-year cycle) combined with 1.7 Recent glacial and interglacial greenhouse gas feedbacks in the 41,000- and 26,000- year cycles. Yet another theory has been advanced by phases Peter Huybers who argued that the 41,000-year cycle has always been dominant, but that the Earth has entered a mode of climate behavior where only the second or third cycle triggers an ice age. This would imply that the 100,000-year periodicity is really an illusion created by averaging together cycles lasting 80,000 and 120,000 years.[57] This theory is consistent with a simple empirical multi-state model proposed by Didier Paillard.[58] Pail- lard suggests that the late Pleistocene glacial cycles can be seen as jumps between three quasi-stable climate states. The jumps are induced by the orbital forcing, while in the early Pleistocene the 41,000-year glacial cycles resulted from jumps between only two climate states. A dynamical model explaining this behavior was proposed by Peter Ditlevsen.[59] This is in support of the suggestion that the late Pleistocene glacial cycles are not due to the Northern hemisphere glaciation during the last ice ages. The set weak 100,000-year eccentricity cycle, but a non-linear re- up of 3 to 4 km thick ice sheets caused a sea level lowering of sponse to mainly the 41,000-year obliquity cycle. about 120 m.

Main article: Timeline of glaciation 1.6.6 Variations in the Sun’s energy output

There are at least two types of variation in the Sun’s energy output 1.7.1 Glacial stages in North America

• In the very long term, astrophysicists believe that the The major glacial stages of the current ice age in North Sun’s output increases by about 7% every one billion America are the Illinoian, Sangamonian and Wisconsin (109) years. stages. The use of the Nebraskan, Afton, Kansan, and Yarmouthian (Yarmouth) stages to subdivide the ice • Shorter-term variations such as sunspot cycles, and age in North America have been discontinued by Qua- longer episodes such as the Maunder minimum, ternary geologists and geomorphologists. These stages 1.8. EFFECTS OF GLACIATION 9 have all been merged into the Pre-Illinoian Stage in the 8.4°C— here was an increase in precipitation. Accord- 1980s.[61][62][63] ingly, at glacial times the humid climatic belt that today is situated several latitude degrees further to the S, was During the most recent North American glaciation, dur- [67][68] ing the latter part of the Wisconsin Stage (26,000 to shifted much further to the N. 13,300 years ago), ice sheets extended to about 45 degrees north latitude. These sheets were 3 to 4 km thick.[62] 1.8 Effects of glaciation This Wisconsin glaciation left widespread impacts on the North American landscape. The Great Lakes and the Finger Lakes were carved by ice deepening old val- leys. Most of the lakes in Minnesota and Wisconsin were gouged out by glaciers and later filled with glacial meltwa- ters. The old Teays River drainage system was radically altered and largely reshaped into the Ohio River drainage system. Other rivers were dammed and diverted to new channels, such as the Niagara, which formed a dramatic waterfall and gorge, when the waterflow encountered a limestone escarpment. Another similar waterfall, at the present Clark Reservation State Park near Syracuse, New York, is now dry. The area from Long Island to Nantucket was formed from glacial till, and the plethora of lakes on the Canadian Shield in northern Canada can be almost entirely attributed to the action of the ice. As the ice retreated and the rock dust dried, winds carried the material hundreds of miles, forming beds of loess many dozens of feet thick in the Missouri Valley. Isostatic rebound continues to re- shape the Great Lakes and other areas formerly under the weight of the ice sheets. The Driftless Zone, a portion of western and southwestern Wisconsin along with parts of adjacent Minnesota, Iowa, and Illinois, was not covered by glaciers. Scandinavia exhibits some of the typical effects of ice age glaciation such as fjords and lakes. See also: Glacial history of Minnesota See also: Glacial landforms

1.7.2 Last Glacial Period in the semi- Although the last glacial period ended more than 8,000 years ago, its effects can still be felt today. For exam- arid Andes around Aconcagua and ple, the moving ice carved out the landscape in Canada Tupungato (See Canadian Arctic Archipelago), Greenland, northern Eurasia and Antarctica. The erratic boulders, till, A specially interesting climatic change during glacial drumlins, eskers, fjords, kettle lakes, moraines, cirques, times has taken place in the semi-arid Andes. Be- horns, etc., are typical features left behind by the glaciers. side the expected cooling down in comparison with the current climate, a significant precipitation is concerned The weight of the ice sheets was so great that they de- here. So, researches in the presently semiarid subtropic formed the Earth’s crust and mantle. After the ice sheets Aconcagua-massif (6,962 m) have shown an unexpect- melted, the ice-covered land rebounded. Due to the high edly extensive glacial glaciation of the type “ice stream viscosity of the Earth’s mantle, the flow of mantle rocks network”.[64][65][66][67][68] The connected valley glaciers which controls the rebound process is very slow—at a rate exceeding 100 km in length, flowed down on the East- of about 1 cm/year near the center of rebound area today. side of this section of the Andes at 32–34°S and 69– During glaciation, water was taken from the oceans to 71°W as far as a height of 2,060 m and on the western form the ice at high latitudes, thus global sea level luff-side still clearly deeper.[68][69] Where current glaciers dropped by about 110 meters, exposing the continental scarcely reach 10 km in length, the snowline (ELA) runs shelves and forming land-bridges between land-masses at a height of 4,600 m and at that time was lowered to for animals to migrate. During deglaciation, the melted 3,200 m asl, i.e. about 1,400 m. From this follows that— ice-water returned to the oceans, causing sea level to rise. beside of an annual depression of temperature about c. This process can cause sudden shifts in coastlines and 10 CHAPTER 1. ICE AGE hydration systems resulting in newly submerged lands, [2] Gribbin, J.R. (1982). Future Weather: Carbon Diox- emerging lands, collapsed ice dams resulting in salination ide, Climate and the Greenhouse Effect. Penguin. ISBN of lakes, new ice dams creating vast areas of freshwater, 0140224599. and a general alteration in regional weather patterns on [3] Rémis, F.; Testus, L.; Testut (2006). “Mais com- a large but temporary scale. It can even cause tempo- ment s’écoule donc un glacier ? Aperçu his- rary reglaciation. This type of chaotic pattern of rapidly torique” (PDF). C. R. Geoscience (in French) 338 changing land, ice, saltwater and freshwater has been pro- (5): 368–385. Bibcode:2006CRGeo.338..368R. posed as the likely model for the Baltic and Scandinavian doi:10.1016/j.crte.2006.02.004. Note: p. 374 regions, as well as much of central North America at the end of the last glacial maximum, with the present-day [4] Montgomery 2010 coastlines only being achieved in the last few millennia [5] Martel, Pierre (1898). “Appendix: Martel, P. (1744) An of prehistory. Also, the effect of elevation on Scandi- account of the glacieres or ice alps in Savoy, in two letters, navia submerged a vast continental plain that had existed one from an English gentleman to his friend at Geneva ; under much of what is now the North Sea, connecting the the other from Pierre Martel , engineer, to the said English British Isles to Continental Europe.[70] gentleman”. In Mathews, C.E. The annals of Mont Blanc. London: Unwin. p. 327. See (Montgomery 2010) for a The redistribution of ice-water on the surface of the full bibliography Earth and the flow of mantle rocks causes changes in the gravitational field as well as changes to the distribution [6] Krüger, Tobias (2013). Discovering the Ice Ages. Interna- of the moment of inertia of the Earth. These changes to tional Reception and Consequences for a Historical Under- the moment of inertia result in a change in the angular standing of Climate (German editon: Basel 2008). Leiden. velocity, axis, and wobble of the Earth’s rotation. p. 47. ISBN 978-90-04-24169-5. The weight of the redistributed surface mass loaded the [7] Krüger 2013, pp. 78-83 lithosphere, caused it to flex and also induced stress within the Earth. The presence of the glaciers generally sup- [8] Krüger 2013, p. 150 [71][72][73] pressed the movement of faults below. How- [9] Krüger 2013, pp. 83, 151 ever, during deglaciation, the faults experience acceler- ated slip triggering earthquakes. Earthquakes triggered [10] Goethe, Johann Wolfgang von: Geologische Probleme near the ice margin may in turn accelerate ice calving and und Versuch ihrer Auflösung, Mineralogie und Geologie may account for the Heinrich events.[74] As more ice is rein Goethes Werke, Weimar 1892, ISBN 3-423-05946-X, moved near the ice margin, more intraplate earthquakes book 73 (WA II,9), p. 253, 254. are induced and this positive feedback may explain the [11] Krüger 2013, p. 83 fast collapse of ice sheets. [12] Krüger 2013, p. 38 In Europe, glacial erosion and isostatic sinking from weight of ice made the Baltic Sea, which before the Ice [13] Krüger 2013, pp. 61-2 Age was all land drained by the Eridanos River. [14] Krüger 2013, pp. 88–90

[15] Krüger 2013, pp. 91-6 1.9 See also [16] Andersen, Bjørn G. (1992). "Jens Esmark—a pioneer in glacial geology” 21. Boreas. pp. 97–102. • Global cooling [17] Davies, Gordon L. (1969). The Earth in Decay. A His- • International Union for Quaternary Research tory of British Geomorphology 1578–1878. London. pp. • Irish Sea Glacier 267f. Cunningham, Frank F. (1990). James David Forbes. Pio- • Late Glacial Maximum neer Scottish Glaciologist. Edinburgh: Scottish Academic Press. p. 15. ISBN 0707303206. • Little Ice Age [18] Krüger 2013, pp. 142-47 • Post-glacial rebound [19] Krüger 2013, pp. 104–05 • Timeline of glaciation [20] Krüger 2013, pp. 150–53

[21] Krüger 2013, pp. 155–59 1.10 References [22] Krüger 2013, pp. 167-70

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1.11 External links

• Cracking the Ice Age from PBS

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

• Raymo, M. (July 2011). “Overview of the Uplift- Weathering Hypothesis”. • Eduard Y. Osipov ., Oleg M. Khlystov. Glaciers and meltwater ﬂux to Lake Baikal during the Last Glacial Maximum.

• Black, R. (9 January 2012). “Carbon emissions 'will defer Ice Age'". BBC News: Science and Environ- ment. Chapter 2

Megafauna

This article is about large land animals. For naked-eye Eurasia, the Americas and Australia became extinct as visible bottom-dwelling animals, see Macrobenthos. For recently as 10,000–40,000 years ago.[4] It is also com- giant animals in mythology, see Megafauna (mythology). monly used for the largest extant wild land animals, espe- In terrestrial zoology, megafauna (Ancient Greek megas cially elephants, giraﬀes, hippopotamuses, rhinoceroses, and large bovines. Megafauna may be subcategorized by their trophic position into megaherbivores (e.g., elk), megacarnivores (e.g., lions), and, more rarely, megaom- nivores (e.g., bears). Other common uses are for giant aquatic species, especially whales, any larger wild or domesticated land animals such as larger antelope and cattle, as well as numerous dinosaurs and other extinct giant reptilians. The term is also sometimes applied to animals (usually extinct) of great size relative to a more common or surviving type of the animal, for example the 1 m (3 ft) dragonﬂies of the Carboniferous period.

2.1 Ecological strategy

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

“large” + New Latin fauna “animal”) are large or giant 2.2 Evolution of large body size animals. The most common thresholds used are 45 kilo- [1][2] [2][3] grams (100 lb) or 100 kilograms (220 lb). This One observation that has been made about the evolution thus includes many species not popularly thought of as of larger body size is that rapid rates of increase that are overly large, such as white-tailed deer, red kangaroo, and often seen over relatively short time intervals are not sus- humans. tainable over much longer time periods. In an exam- In practice, the most common usage encountered in aca- ination of mammal body mass changes over time, the demic and popular writing describes land animals roughly maximum increase possible in a given time interval was larger than a human that are not (solely) domesticated. found to scale with the interval length raised to the 0.25 The term is especially associated with the Pleistocene power.[5] This is thought to reﬂect the emergence, dur- megafauna – the land animals often larger than modern ing a trend of increasing maximum body size, of a se- counterparts considered archetypical of the last ice age, ries of anatomical, physiological, environmental, genetic such as mammoths, the majority of which in northern and 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- been predicted based on the metabolic rate of mammals, sible. A strikingly faster rate of change was found for the energetic cost of obtaining prey, and the maximum es- large decreases in body mass, such as may be associated timated rate coefficient of prey intake.[9] It has also been with the phenomenon of insular dwarfism. When nor- suggested that maximum size for mammalian carnivores malized to generation length, the maximum rate of body is constrained by the stress the humerus can withstand at mass decrease was found to be over 30 times greater than top running speed.[8] the maximum rate of body mass increase for a ten-fold [5] Analysis of the variation of maximum body size over the change. last 40 Ma suggests that decreasing temperature and increasing continental land area are associated with increas- 2.2.1 In terrestrial mammals ing maximum body size. The former correlation would be consistent with Bergmann’s rule,[10] and might be re- Subsequent to the Cretaceous–Paleogene extinction event lated to the thermoregulatory advantage of large body [6] that eliminated the non-avian dinosaurs about 66 Ma ago, mass in cool climates, better ability of larger organ- [10] terrestrial mammals underwent a nearly exponential in- isms to cope with seasonality in food supply, or other [10] crease in body size as they diversified to occupy the eco- factors; the latter correlation could be explainable in [6] logical niches left vacant.[6] Starting from just a few kg terms of range and resource limitations. However, the before the event, maximum size had reached ~50 kg a two parameters are interrelated (due to sea level drops ac- few million years later, and ~750 kg by the end of the companying increased glaciation), making the driver of [6] Paleocene. This trend of increasing body mass appears the trends in maximum size more difficult to identify. to level off about 40 Ma ago (in the late Eocene), suggesting that physiological or ecological constraints had been reached, after an increase in body mass of over three or- 2.2.2 In marine mammals ders of magnitude.[6] However, when considered from the The ancestors of cetaceans are believed to have been the standpoint of rate of size increase per generation, the ex- semiaquatic pakicetids, no larger than wolves, of about ponential increase is found to have continued until the 53 million years (Ma) ago.[11] By 40 Ma ago, cetaceans appearance of Indricotherium 30 Ma ago. (Since gener- had attained a length of 20 m or more in Basilosaurus, ation time scales with body mass0.259, increasing genera- an elongated, serpentine whale that differed from mod- tion times with increasing size cause the log mass vs. time ern whales in many respects and was not ancestral to plot to curve downward from a linear fit.)[5] them. Following this, the evolution of large body size Megaherbivores eventually attained a body mass of over in cetaceans appears to have come to a temporary halt, 10 000 kg. The largest of these, indricotheres and and then to have backtracked, although the available fossil proboscids, have been hindgut fermenters, which are be- records are limited. However, in the period from 31 Ma lieved to have an advantage over foregut fermenters in ago (in the Oligocene) to the present, cetaceans under- terms of being able to accelerate gastrointestinal tran- went a significantly more rapid sustained increase in body sit in order to accommodate very large food intakes.[7] mass (a rate of increase in body mass0.259 of a factor of 3.2 A similar trend emerges when rates of increase of max- per million years) than achieved by any group of terres- imum body mass per generation for different mam- trial mammals.[5] This trend led to the largest animal of malian clades are compared (using rates averaged over all time, the modern blue whale. Several reasons for the macroevolutionary time scales). Among terrestrial mam- more rapid evolution of large body size in cetaceans are mals, the fastest rates of increase of body mass0.259 vs. possible. Fewer biomechanical constraints on increases time (in Ma) occurred in perissodactyls (a slope of 2.1), in body size may be associated with suspension in water followed by rodents (1.2) and proboscids (1.1),[5] all of as opposed to standing against the force of gravity, and which are hindgut fermenters. The rate of increase for with swimming movements as opposed to terrestrial lo- artiodactyls (0.74) was about a third that of perisso- comotion. Also, the greater heat capacity and thermal dactyls. The rate for carnivorans (0.65) was slightly lower conductivity of water compared to air may increase the yet, while primates, perhaps constrained by their arboreal thermoregulatory advantage of large body size in marine habits, had the lowest rate (0.39) among the mammalian endotherms, although diminishing returns apply.[5] groups studied.[5] Cetaceans are not the only marine mammals to reach Terrestrial mammalian carnivores from several eutherian unprecedented size in the modern era. The largest groups (the mesonychid Andrewsarchus, the creodonts carnivoran of all time is the mostly aquatic modern Megistotherium and Sarkastodon, and the carnivorans southern elephant seal. Amphicyon and Arctodus) all reached a maximum size of about 1000 kg[6] (the carnivoran Arctotherium apparently actually got somewhat larger). The largest known 2.2.3 In flightless birds metatherian carnivore, Proborhyaena gigantea, apparently reached 600 kg, also close to this limit.[8] A similar Because of the small initial size of all mammals follow- theoretical maximum size for mammalian carnivores has ing the extinction of the dinosaurs, nonmammalian ver- 16 CHAPTER 2. MEGAFAUNA

tebrates had a roughly ten million year long window of of megafaunal extinction pulses that have occurred dur- opportunity (during the Paleocene) for evolution of gi- ing the last 50,000 years over much of the Earth’s surface, gantism without much competition.[12] During this inter- with Africa and southern Asia being largely spared. The val, apex predator niches were often occupied by reptiles, latter areas did suffer a gradual attrition of megafauna, such as terrestrial crocodilians (e.g. Pristichampsus), particularly of the slower-moving species (a class of vul- large snakes (e.g. Titanoboa) or varanid lizards, or by nerable megafauna epitomized by giant tortoises), over flightless birds[6] (e.g. Gastornis in Europe and North the last several million years.[20][21] America, Paleopsilopterus in South America). This is Outside the mainland of Afro-Eurasia, these megafau- also the period when flightless herbivorous paleognath nal extinctions followed a highly distinctive landmass-by- birds evolved to large size on a number of Gondwanan landmass pattern that closely parallels the spread of hu- land masses. These birds, termed ratites, have tradi- mans into previously uninhabited regions of the world, tionally been viewed as representing a lineage separate and which shows no correlation with climatic history from that of their small flighted relatives, the Neotropic (which can be visualized with plots over recent geolog- tinamous. However, recent genetic studies have found ical time periods of climate markers such as marine oxy- that tinamous nest well within the ratite tree, and are the [22][23] [13][12][14] gen isotopes or atmospheric carbon dioxide levels). sister group of the extinct moa of New Zealand. Australia was struck first around 45,000 years ago,[24] Similarly, the small kiwi of New Zealand have been found followed by Tasmania about 41,000 years ago (after to be the sister group of the extinct elephant birds of [12] formation of a land bridge to Australia about 43,000 Madagascar. These findings indicate that flightlessness years ago),[25][26][27] Japan apparently about 30,000 years and gigantism arose independently multiple times among ago,[28] North America 13,000 years ago, South Amer- ratites via parallel evolution. ica about 500 years later,[29][30] Cyprus 10,000 years In the northern continents, large predatory birds were dis- ago,[31][32] the Antilles 6000 years ago,[33] New Caledo- placed when large eutherian carnivores evolved. In iso- nia[34] and nearby islands[35] 3000 years ago, Madagascar lated South America, the phorusrhacids could not be out- 2000 years ago,[36] New Zealand 700 years ago,[37] the competed by the local metatherian sparassodonts and re- Mascarenes 400 years ago,[38] and the Commander Is- mained dominant until advanced eutherian predators ar- lands 250 years ago.[39] Nearly all of the world’s iso- rived from North America (as part of the Great Amer- lated islands could furnish similar examples of extinc- ican Interchange) during the Pliocene. However, none tions occurring shortly after the arrival of Homo sapi- of the largest predatory (Brontornis), possibly omnivo- ens, though most of these islands, such as the Hawaiian rous (Dromornis[15]) or herbivorous (Aepyornis) flightless Islands, never had terrestrial megafauna, so their extinct birds of the Cenozoic ever grew to masses much above fauna were smaller.[22][23] 500 kg, and thus never attained the size of the largest An analysis of Sporormiella fungal spores (which de- mammalian carnivores, let alone that of the largest mam- rive mainly from the dung of megaherbivores) in swamp malian herbivores. It has been suggested that the increas- sediment cores spanning the last 130,000 years from ing thickness of avian eggshells in proportion to egg mass Lynch’s Crater in Queensland, Australia showed that with increasing egg size places an upper limit on the size [16][note 1] the megafauna of that region virtually disappeared about of birds. The largest species of Dromornis, D. 41,000 years ago, at a time when climate changes were stirtoni, may have gone extinct after it attained the max- minimal; the change was accompanied by an increase imum avian body mass and was then outcompeted by in charcoal, and was followed by a transition from rain- marsupial diprotodonts that evolved to sizes several times [19] forest to fire-tolerant sclerophyll vegetation. The high- larger. resolution chronology of the changes supports the hypothesis that human hunting alone eliminated the megafauna, and that the subsequent change in flora was most likely 2.3 Megafaunal mass extinctions a consequence of the elimination of browsers and an increase in fire.[40][41][42] The increase in fire lagged the disappearance of megafauna by about a century, and most 2.3.1 Timing and possible causes likely resulted from accumulation of fuel once browsing stopped. Over the next several centuries grass in- A well-known mass extinction of megafauna, the creased; sclerophyll vegetation increased with a lag of Holocene extinction (see also Quaternary extinction another century, and a sclerophyll forest developed af- event), occurred at the end of the last ice age glacial peter about another thousand years.[42] During two periods riod (a.k.a. the Würm glaciation) and wiped out many of climate change about 120 and 75 thousand years ago, giant ice age animals, such as woolly mammoths, in sclerophyll vegetation had also increased at the site in re- the Americas and northern Eurasia. Various theories sponse to a shift to cooler, drier conditions; neither of have attributed the wave of extinctions to human hunt- these episodes had a significant impact on megafaunal ing, climate change, disease, a putative extraterrestrial abundance.[42] Similar conclusions regarding the culpa- impact, or other causes. However, this extinction pulse bility of human hunters in the disappearance of Pleis- near the end of the Pleistocene was just one of a series 2.4. EXAMPLES 17

tocene megafauna were obtained via an analysis of a large of the time (up to 10 C warmer than at present).[50][51] collection of eggshell fragments of the flightless Aus- This large emission follows from the enormous estimated tralian bird Genyornis newtoni[43] and from analysis of biomass of sauropods, and because methane production Sporormiella fungal spores from a lake in eastern North of individual herbivores is believed to be almost propor- America.[44][45] tional to their mass.[50] Continuing human hunting and environmental distur- Recent studies have indicated that the extinction of bance has led to additional megafaunal extinctions in the megafaunal herbivores may have caused a reduction recent past, and has created a serious danger of further in atmospheric methane. This hypothesis is relatively extinctions in the near future (see examples below). new.[52] One study examined the methane emissions from A number of other mass extinctions occurred earlier in the bison that occupied the Great Plains of North Amer- Earth’s geologic history, in which some or all of the ica before contact with European settlers. The study estimated that the removal of the bison caused a decrease megafauna of the time also died out. Famously, in the [53] Cretaceous–Paleogene extinction event the dinosaurs and of as much as 2.2 million tons per year. Another study most other giant reptilians were eliminated. However, the examined the change in the methane concentration in the earlier mass extinctions were more global and not so se- atmosphere at the end of the Pleistocene epoch after the lective for megafauna; i.e., many species of other types, extinction of megafauna in the Americas. After early hu- including plants, marine invertebrates[46] and plankton, mans migrated to the Americas about 13,000 BP, their went extinct as well. Thus, the earlier events must have hunting and other associated ecological impacts led to been caused by more generalized types of disturbances to the extinction of many megafaunal species there. Cal- the biosphere. culations suggest that this extinction decreased methane production by about 9.6 million tons per year. This suggests that the absence of megafaunal methane emissions 2.3.2 Consequences of depletion of may have contributed to the abrupt climatic cooling at the [52] megafauna onset of the Younger Dryas. The decrease in atmospheric methane that occurred at that time, as recorded in ice cores, was 2-4 times more rapid than any other de- Effect on nutrient transport crease in the last half million years, suggesting that an unusual mechanism was at work.[52] Megafauna play a significant role in the lateral transport of mineral nutrients in an ecosystem, tending to translocate them from areas of high to those of lower abundance. They do so by their movement between the 2.4 Examples time they consume the nutrient and the time they release it through elimination (or, to a much lesser extent, The following are some notable examples of animals of- through decomposition after death).[47] In South Amer- ten considered as megafauna (in the sense of the “large ica’s Amazon Basin, it is estimated that such lateral dif- animal” definition). This list is not intended to be ex- fusion was reduced over 98% following the megafaunal haustive: extinctions that occurred roughly 12,500 years ago.[48][49] Given that phosphorus availability is thought to limit pro- • Clade Synapsida ductivity in much of the region, the decrease in its trans- • Class Mammalia (phylogenetically, a clade port from the western part of the basin and from flood- within Therapsida; see below) plains (both of which derive their supply from the uplift • Infraclass Metatheria of the Andes) to other areas is thought to have signifi- • cantly impacted the region’s ecology, and the effects may Order Diprotodontia • not yet have reached their limits.[49] The red kangaroo (Macropus ru- fus) is the largest living Australian mammal and marsupial at a weight Effect on methane emissions of up to 85 kg (187 lb). How- ever, its extinct relative, the giant Large populations of megaherbivores have the potential short-faced kangaroo Procoptodon to contribute greatly to the atmospheric concentration of goliah reached 230 kg (510 lb), methane, which is an important greenhouse gas. Mod- while extinct diprotodonts attained ern ruminant herbivores produce methane as a byprod- the largest size of any marsupial in uct of foregut fermentation in digestion, and release it history, up to an estimated 2,750 kg through belching. Today, around 20% of annual methane (6,060 lb). The extinct marsupial emissions come from livestock methane release. In the lion (Thylacleo carnifex), at up to Mesozoic, it has been estimated that sauropods could 160 kg (350 lb) was much larger have emitted 520 million tons of methane to the atmo- than any extant carnivorous marsu- sphere annually,[50] contributing to the warmer climate pial. 18 CHAPTER 2. MEGAFAUNA

• Infraclass Eutheria north (e.g., Megalonyx). The • Superorder Afrotheria largest genera, Megatherium and • Order Proboscidea Eremotherium, reached sizes comparable to elephants.[54] • Elephants are the largest living • land animals. They and their rel- Superorder Euarchontoglires atives arose in Africa, but until • Order Primates recently had a nearly worldwide • The largest living primate, at up distribution. The African bush to 266 kg (586 lb), is the gorilla elephant (Loxodonta africana) has (Gorilla beringei and Gorilla go- a shoulder height of up to 4.3 rilla, with three of four sub- m (14 ft) and weighs up to 13 species being critically endan- tons. Among recently extinct gered). The extinct Malagasy proboscideans, mammoths (Mam- sloth lemur Archaeoindris reached muthus) were close relatives of ele- a similar size, while the ex- phants, while mastodons (Mam- tinct Gigantopithecus blacki of mut) were much more distantly Southeast Asia is believed to have related. The steppe mammoth been several times larger. Some (M. trogontherii) is estimated to populations of archaic Homo were have commonly weighed around significantly larger than recent 10 tonnes, making it possibly the Homo sapiens;[55][56] for example, largest proboscid, which would Homo heidelbergensis in south- make it the second largest land ern Africa may have commonly mammal after indricotherines. reached 7 feet (2.1 m) in height,[57] • Order Sirenia while Neanderthals were about [58] • The largest sirenian at up to 1500 30% more massive. kg is the West Indian manatee • Order Rodentia (Trichechus manatus). Steller’s • The extant capybara (Hydro- sea cow (Hydrodamalis gigas) was choerus hydrochaeris) of South probably around five times as mas- America, the largest living rodent, sive, but was exterminated by hu- weighs up to 65 kg (143 lb). mans within 27 years of its dis- Several recently extinct North covery off the remote Commander American forms were larger: the Islands in 1741. In prehistoric capybara Neochoerus pinckneyi times this sea cow also lived along (another neotropic migrant) was the coasts of northeastern Asia about 40% heavier; the giant and northwestern North America; beaver (Castoroides ohioensis) was it was apparently eliminated from similar. The extinct blunt-toothed these more accessible locations by giant hutia (Amblyrhiza inundata) aboriginal hunters. of several Caribbean islands may • Superorder Xenarthra have been larger still. However, • Order Cingulata several million years ago South • The glyptodonts were a group of America harbored much more large, heavily armored ankylosaur- massive rodents. Phoberomys pat- like xenarthrans related to liv- tersoni, known from a nearly full ing armadillos. They originated skeleton, probably reached 700 kg in South America, invaded North (1,500 lb). Fragmentary remains America during the Great Amer- suggest that Josephoartigasia ican Interchange, and went ex- monesi grew to upwards of 1,000 tinct at the end of the Pleistocene kg (2,200 lb). epoch.[54] • Superorder Laurasiatheria • Order Pilosa • Order Carnivora • Ground sloths were another group • Big cats include the tiger (Pan- of slow, terrestrial xenarthrans, thera tigris) and lion (Panthera related to modern tree sloths. leo). The largest subspecies, at up They had a similar history, al- to 306 kg (675 lb), is the Siberian though they reached North Amer- tiger (P. tigris altaica), in accord ica earlier, and spread farther with Bergmann’s rule. Members 2.4. EXAMPLES 19

of Panthera are distinguished by Rhinoceroses are odd-toed ungu- morphological features which en- lates with horns made of keratin, able them to roar. Larger ex- the same type of protein compos- tinct felids include the American ing hair. They are among the lion (Panthera leo atrox) and the largest living land mammals af- South American saber-toothed cat ter elephants (hippos attain a sim- Smilodon populator. ilar size). Three of five extant • Bears are large carnivorans of the species are critically endangered. caniform suborder. The largest Their extinct central Asian rela- living forms are the polar bear (Ur- tives the indricotherines were the sus maritimus), with a body weight largest terrestrial mammals of all of up to 680 kg (1,500 lb), and time. the similarly sized Kodiak bear • Order Artiodactyla (or cladistically, (Ursus arctos middendorffi), again Cetartiodactyla) consistent with Bergmann’s rule. • Giraffes (Giraffa camelopardalis) Arctotherium augustans, an ex- are the tallest living land animals, tinct short-faced bear from South reaching heights of up to nearly 6 America, was the largest predatory m (20 ft). land mammal ever with an esti- • Bovine ungulates include the mated average weight of 1,600 kg largest surviving land animals of (3,500 lb).[59] Europe and North America. The • Seals, sea lions, and walruses water buffalo (Bubalis arnee), are amphibious marine carnivo- bison (Bison bison and B. bona- rans that evolved from bearlike sus), and gaur (Bos gaurus) can all ancestors. The southern ele- grow to weights of over 900 kg phant seal (Mirounga leonina) of (2,000 lb). Antarctic and subantarctic waters • is the largest carnivoran of all time, The semiaquatic hippopotamus with bull males reaching a maxi- (Hippopotamus amphibius) is the mum length of 6–7 m (20–23 ft) heaviest living even-toed ungulate; and maximum weight of 5,000 kg it and the critically endangered (11,000 lb). pygmy hippo (Choeropsis liberien- sis) are believed to be the closest • Order Perissodactyla extant relatives of cetaceans. • Tapirs are browsing animals, with Hippos are among the megafau- a short prehensile snout and pig- nal species most dangerous to like form that appears to have humans.[60] changed little in 20 million years. • Order Cetacea (or cladistically, They inhabit tropical forests of Cetartiodactyla) Southeast Asia and South and • Central America, and include the Whales, dolphins, and porpoises largest surviving land animals of are marine mammals. The blue the latter two regions. There are whale (Balaenoptera musculus) is four species. the largest baleen whale and the largest animal that has ever lived, at 30 metres (98 ft)[61] in length and 170 tonnes (190 short tons)[62] or more in weight. The sperm whale (Physeter macrocephalus) is the largest toothed whale, as well as the planet’s loudest and brainiest animal (with a brain about five times as massive as a human’s). The killer whale (Orcinus orca) is the largest dolphin. • Order Pelycosauria (traditional; paraphyletic) • • Cotylorhynchus was a large, big-clawed, Rhinoceros, from Dürer’s woodcut herbivorous caseid of Early Permian 20 CHAPTER 2. MEGAFAUNA

North America, reaching 6 m (20 ft) and • Order Saurischia 2 tonnes. • Saurischian dinosaurs of the Jurassic • Order Therapsida and Cretaceous include sauropods, • Anteosaurus was a headbutting, semi- the longest (at up to 40 m or 130 ft) aquatic, carnivorous dinocephalian of and most massive terrestrial animals Middle Permian South Africa. It reached known (Argentinosaurus reached 80– 5–6 m (16–20 ft) long, and weighed about 100 metric tonnes, or 90–110 tons), 500–600 kg (1,100–1,300 lb).[63] as well as theropods, the largest terrestrial carnivores (Spinosaurus grew • Clade Sauropsida to 7–9 tonnes; the more famous • Class Aves (phylogenetically, a clade within Tyrannosaurus, to 6.8 tonnes). Coelurosauria, a taxon within the order • Order Squamata Saurischia; see below) • While the largest extant lizard, the • Order Struthioniformes Komodo dragon (Varanus komod- • The ratites are an ancient and oensis), another island giant, can diverse group of ﬂightless birds reach 3 m (10 ft) in length, its extinct that are found on fragments of the Australian relative Megalania may former supercontinent Gondwana. have reached more than twice that The largest living bird, the ostrich size. These monitor lizards' marine (Struthio camelus) was surpassed by relatives, the mosasaurs, were apex the extinct Aepyornis of Madagascar, predators in late Cretaceous seas. the heaviest of the group (400 kg • The heaviest extant snake is consid- (880 lb)), and the extinct giant moa ered to be the green anaconda (Eu- (Dinornis) of New Zealand, the nectes murinus), while the reticulated tallest, growing to heights of 3.4 m python (Python reticulatus), at up (11 ft). The latter two are examples to 8.7 m or more, is considered of island gigantism. the longest. An extinct Australian • Order Anseriformes Pliocene species of Liasis, the Bluﬀ • Extinct dromornithids of Australia Downs giant python, reached 10 m, such as Dromornis may have ex- while the Paleocene Titanoboa of ceeded the largest ratites in size. South America reached lengths of (Due to its small size for a continent 12–15 m and an estimated weight of and its isolation, Australia is some- about 1135 kilograms (2500 lb). times viewed as the world’s largest is- • Order Testudines land; thus, these species could also be • The largest turtle is the critically en- considered insular giants.) dangered marine leatherback turtle • Class Reptilia (traditional; paraphyletic) (Dermochelys coriacea), weighing • Order Crocodilia up to 900 kg (2,000 lb). It is dis- • Alligators and crocodiles are large tinguished from other sea turtles by semiaquatic reptiles, the largest its lack of a bony shell. The most of which, the saltwater crocodile massive terrestrial chelonians are (Crocodylus porosus), can grow to the giant tortoises of the Galápagos a weight of 1,360 kg (3,000 lb). Islands (Chelonoidis nigra) and Crocodilians’ distant ancestors and Aldabra Atoll (Aldabrachelys gigan- their kin, the crurotarsans, domi- tea), at up to 300 kg (660 lb). These nated the world in the late Triassic, tortoises are the biggest survivors until the Triassic–Jurassic extinction of an assortment of giant tortoise event allowed dinosaurs to over- species that were widely present take them. They remained diverse on continental landmasses[64][65] during the later Mesozoic, when and additional islands[64] during the crocodyliforms such as Deinosuchus Pleistocene. and Sarcosuchus reached lengths of 12 m. Similarly large crocodil- • Class Amphibia (in the wide, probably paraphyletic, ians, such as Mourasuchus and sense) Purussaurus, were present as recently as the Miocene in South • Order Temnospondyli (relationship to extant America. amphibians is unclear) 2.5. GALLERY 21

• The Permian temnospondyl • Order Rajiformes Prionosuchus, the largest amphibian • The manta ray (Manta birostris) is another known, reached 9 m in length and filter feeder and the largest ray, growing was an aquatic predator resembling a to up to 2300 kg. crocodilian. After the appearance of real crocodilians, temnospondyls such as • Class Placodermi Koolasuchus (5 m long) had retreated to • Order Arthrodira the Antarctic region by the Cretaceous, before going extinct. • The largest armored fish, Dunkleosteus, arose during the late Devonian. At up • Class Actinopterygii to 10 metres (33 ft) in length[66] and 3.6 tonnes (4.0 short tons) in mass,[67] it was a • Order Tetraodontiformes hypercarnivorous apex predator that em- • The largest extant bony fish is the ocean ployed suction feeding.[68][69] Its contem- sunfish (Mola mola), whose average adult porary, Titanichthys, apparently an early weight is 1,000 kg (2,200 lb). While phy- filter feeder, rivaled it in size. The an- logenetically a “bony fish”, its skeleton is throdires were eliminated by the environ- primarily cartilage (which is lighter than mental upheavals of the Late Devonian bone). It has a disk-shaped body, and pro- extinction, after existing for only about 50 pels itself with its long, thin dorsal and million years. anal fins; it feeds primarily on jellyfish. In these three respects (as well as in its • Class Cephalopoda size and diving habits), it resembles a • Order Teuthida leatherback turtle. • A number of deep ocean creatures ex- • Order Acipenseriformes hibit abyssal gigantism. These include • The critically endangered beluga (Eu- the giant squid (Architeuthis) and colossal ropean sturgeon, Huso huso) at up to squid (Mesonychoteuthis hamiltoni); both 1,476 kg (3,254 lb) is the largest sturgeon (although rarely seen) are believed to at- (which are also mostly cartilaginous) and tain lengths of 12 m (39 ft) or more. The is considered the largest anadromous fish. latter is the world’s largest invertebrate, • Order Siluriformes and has the largest eyes of any animal. Both are preyed upon by sperm whales. • The critically endangered Mekong giant catfish (Pangasianodon gigas), at up to • Subphylum Chelicerata 293 kg (646 lb), is often viewed as the • largest freshwater fish. Order Eurypterida • Eurypterids (sea scorpions) were a di- • Class Chondrichthyes verse group of aquatic and possibly amphibious predators that included the most • Order Lamniformes massive arthropods to have existed. They • The largest living predatory fish, the great survived over 200 million years, but fi- white shark (Carcharodon carcharias), nally died out in the Permian–Triassic ex- reaches weights up to 2,240 kg (4,940 tinction event along with trilobites and lb). Its extinct relative C. megalodon (the most other forms of life present at the disputed genus being either Carcharodon time, including most of the dominant ter- or Carcharocles) was more than an order restrial therapsids. The Early Devonian of magnitude larger, and is the largest Jaekelopterus reached an estimated length predatory shark or fish of all time (and of 2.5 m (8.2 ft), not including its possibly the largest predator in vertebrate raptorial chelicerae, and is thought to history); it preyed on whales and other have been a freshwater species. marine mammals. • Order Orectolobiformes • The largest extant shark, cartilaginous 2.5 Gallery fish, and fish overall is the whale shark (Rhincodon typus), which reaches weights 2.5.1 Extinct in excess of 21.5 tonnes (47,000 pounds). Like baleen whales, it is a filter feeder and • Some Paleozoic sea scorpions (Eurypterus shown) primarily consumes plankton. were larger than a man. 22 CHAPTER 2. MEGAFAUNA

• Dunkleosteus was a 10 m (33 ft) long toothless 2.5.2 Living armored predatory Devonian placoderm fish. • The gorilla is the largest and one of the most • Sail-backed pelycosaur Dimetrodon and endangered primates on the planet. temnospondyl Eryops from North America’s • Siberian tigers are the biggest living cats, exempli- Permian. fying Bergmann’s rule. • Pliosaur Pliosaurus (right) harassing the filter feeder • Polar bears, the largest bears and semiaquatic carni- fish Leedsichthys during the Jurassic. vores, are vulnerable to global warming.

• Macronarian sauropods; from left, Camarasaurus, • The critically endangered black rhinoceros, up to 14 Brachiosaurus, Giraffatitan, Euhelopus. feet (4.3 m) long, is threatened by poaching. • Wild Bactrian camels are critically endangered. • Tyrannosaurus was a 12.3 m (40 ft) long theropod Their ancestors originated in North America. dinosaur, an apex predator of west North America. • Unlike woolly rhinos and mammoths, muskoxen • Indricotheres, the land mammals closest to narrowly survived the Quaternary extinctions.[1] sauropods in size and lifestyle, were Asian rhinos. • Hippos, the heaviest and most aquatic even-toed un- • The Late Miocene teratorn Argentavis of South gulates, are whales' closest living relatives. America had a 7 m (23 ft) wingspan. • A filter feeder up to 33 m (108 ft) long, the blue whale is the largest animal of all time. • C. megalodon (two possible sizes) with a whale shark, great white shark and human for scale. • The orca, the largest dolphin and pack predator, is highly intelligent and lives in complex societies. • Deinotherium had downward-curving tusks and • ranged widely over Afro-Eurasia. The ostrich is the largest ratite, the heaviest living bird and, at 70 km/h,[2] the fastest running [3][note 1] • Titanis walleri, the only terror bird known to have biped. invaded North America, was 2.5 m (8 ft 2 in) tall. • The saltwater crocodile is the largest living reptile and a dangerous predator of humans. • Hippo-sized Diprotodon of Australia, the largest marsupial of all time, went extinct 40,000 years ago. • The Komodo dragon, an insular giant, is the largest lizard and has infectious and venomous saliva. • Megalania, a giant carnivorous goanna of Australia, • The green anaconda, an aquatic constrictor, is the might have grown to 7 metres long. heaviest snake, weighing up to 97.5 kg (215 lb). • Elephant-sized Megatherium, from South America’s • The deep-diving ocean sunfish is the largest bony Pleistocene, was the largest sloth.[1] fish, but its skeleton is mostly cartilaginous.

• Toxodon, one of South America’s largest and last • The Nile perch, one of the largest freshwater fish, is [note 2] notoungulates.[1] It had a relative in Mexico. also a damaging invasive species. • The whale shark is the largest extant shark or fish • American lions exceeded extant lions in size and species, growing up to 12.6 m (41 ft) in length. ranged over two continents until 10,000 BP. • The manta, a filter feeder, is the largest ray at up to • Woolly mammoths vanished after humans invaded 7.6 m across, yet can breach clear of the water. their habitat in Eurasia and N. America.[2] • Examination of a 9 m giant squid, an abyssal giant • Haast’s eagle, the largest eagle known, attacking and the second largest cephalopod. moa (which included the tallest bird known). 1. ^ Cite error: The named reference Stuart was invoked but never defined (see the help page). 1. ^ Cite error: The named reference Fari.C3.B1a2013 was invoked but never defined 2. ^ Davies, S.J.J.F. (2003). “Birds I Tinamous and (see the help page). Ratites to Hoatzins”. In Hutchins, Michael. Grz- imek’s Animal Life Encyclopedia 8 (2 ed.). Farm- 2. ^ Cite error: The named reference Stuart was in- ington Hills, MI: Gale Group. pp. 99–101. ISBN voked but never defined (see the help page). 0-7876-5784-0. 2.8. REFERENCES 23

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Pleistocene

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

3.1 Dating Earth during the Pleistocene epoch.

[1] The Pleistocene has been dated from 2.588 million The Pleistocene /ˈplaɪstɵsiːn/ (symbol PS ) is the ge- (±.005) to 11,700 years before present (BP), with the end ological epoch which lasted from about 2,588,000 to date expressed in radiocarbon years as 10,000 carbon-14 11,700 years ago, spanning the world’s recent period of years BP.[4] It covers most of the latest period of repeated repeated glaciations. glaciation, up to and including the Younger Dryas cold Charles Lyell introduced this term in 1839 to describe spell. The end of the Younger Dryas has been dated to strata in Sicily that had at least 70% of their mol- about 9640 BC (11,654 calendar years BP). It was not luscan fauna still living today. This distinguished it until after the development of radiocarbon dating, how- from the older Pliocene Epoch, which Lyell had origi- ever, that Pleistocene archaeological excavations shifted nally thought to be the youngest fossil rock layer. He to stratified caves and rock-shelters as opposed to open- constructed the name “Pleistocene” (“Most New” or air river-terrace sites.[5] “Newest”) from the Greek πλεῖστος, pleīstos, “most”, In 2009 the International Union of Geological Sciences and καινός, kainós (latinized as cænus), “new";[2] this (IUGS) confirmed a change in time period for the Pleis- contrasting with the immediately preceding Pleiocene tocene, changing the start date from 1.806 to 2.588 mil- (“More New” or “Newer”, from πλείων, pleíōn, “more”, lion years BP, and accepted the base of the Gelasian as and kainós; usual spelling: Pliocene), and the imme- the base of the Pleistocene, namely the base of the Monte diately subsequent Holocene (“wholly new” or “entirely San Nicola GSSP.[6] The IUGS has yet to approve a type new”, from ὅλος, hólos, “whole”, and kainós) epoch, section, Global Boundary Stratotype Section and Point which extends to the present time. (GSSP), for the upper Pleistocene/Holocene boundary The Pleistocene is the first epoch of the Quaternary (i.e. the upper boundary). The proposed section is the Period or sixth epoch of the Cenozoic Era.[3] The end of North Greenland Ice Core Project ice core 75° 06' N 42° the Pleistocene corresponds with the end of the last glacial 18' W.[7] The lower boundary of the Pleistocene Series period. It also corresponds with the end of the Paleolithic is formally defined magnetostratigraphically as the base age used in archaeology. In the ICS timescale, the Pleis- of the Matuyama (C2r) chronozone, isotopic stage 103. tocene is divided into four stages or ages, the Gelasian, Above this point there are notable extinctions of the cal- Calabrian, Ionian and Tarantian. All of these stages were careous nanofossils: Discoaster pentaradiatus and Dis- defined in southern Europe. In addition to this interna- coaster surculus.[8][9] tional subdivision, various regional subdivisions are often The Pleistocene covers the recent period of repeated used. glaciations. The name Plio-Pleistocene has in the past Before a change finally confirmed in 2009 by the been used to mean the last ice age. The revised defini- International Union of Geological Sciences, the time tion of the Quaternary, by pushing back the start date of boundary between the Pleistocene and the preceding the Pleistocene to 2.58 Ma, results in the inclusion of all Pliocene was regarded as being at 1.806 million years the recent repeated glaciations within the Pleistocene.

27 28 CHAPTER 3. PLEISTOCENE

mean annual temperature at the edge of the ice was −6 °C (21 °F); at the edge of the permafrost, 0 °C (32 °F). Each glacial advance tied up huge volumes of water in continental ice sheets 1,500 to 3,000 metres (4,900– 9,800 ft) thick, resulting in temporary sea-level drops of 100 metres (300 ft) or more over the entire surface of the Earth. During interglacial times, such as at present, drowned coastlines were common, mitigated by isostatic or other emergent motion of some regions. The effects of glaciation were global. Antarctica was ice- bound throughout the Pleistocene as well as the preceding Pliocene. The Andes were covered in the south by the Patagonian ice cap. There were glaciers in New Zealand and Tasmania. The current decaying glaciers of Mount Kenya, Mount Kilimanjaro, and the Ruwenzori Range in east and central Africa were larger. Glaciers existed in the mountains of Ethiopia and to the west in the Atlas mountains. In the northern hemisphere, many glaciers fused into one. The Cordilleran ice sheet covered the North American northwest; the east was covered by the Laurentide. The Fenno-Scandian ice sheet rested on northern Europe, including Great Britain; the Alpine ice sheet on the Alps. Scattered domes stretched across Siberia and the Arctic shelf. The northern seas were ice-covered. South of the ice sheets large lakes accumulated because outlets were blocked and the cooler air slowed evaporation. When the Laurentide ice sheet retreated, north central North America was totally covered by Lake Agassiz. The maximum extent of glacial ice in the north polar area during Over a hundred basins, now dry or nearly so, were over- the Pleistocene period. flowing in the North American west. Lake Bonneville, for example, stood where Great Salt Lake now does. In 3.2 Paleogeography and climate Eurasia, large lakes developed as a result of the runoff from the glaciers. Rivers were larger, had a more copious flow, and were braided. African lakes were fuller, appar- The modern continents were essentially at their present ently from decreased evaporation. Deserts on the other positions during the Pleistocene, the plates upon which hand were drier and more extensive. Rainfall was lower they sit probably having moved no more than 100 km rel- because of the decrease in oceanic and other evaporation. ative to each other since the beginning of the period. According to Mark Lynas (through collected data), the Pleistocene’s overall climate could be characterized as 3.2.2 Major events a continuous El Niño with trade winds in the south Pacific weakening or heading east, warm air rising near Further information: Timeline of glaciation Peru, warm water spreading from the west Pacific and Over 11 major glacial events have been identified, as the Indian Ocean to the east Pacific, and other El Niño well as many minor glacial events.[11] A major glacial markers.[10] event is a general glacial excursion, termed a “glacial.” Glacials are separated by “interglacials”. During a glacial, 3.2.1 Glacial features the glacier experiences minor advances and retreats. The minor excursion is a “stadial"; times between stadials are Pleistocene climate was marked by repeated glacial cycles “interstadials”. in which continental glaciers pushed to the 40th parallel These events are defined differently in different regions in some places. It is estimated that, at maximum glacial of the glacial range, which have their own glacial history extent, 30% of the Earth’s surface was covered by ice. In depending on latitude, terrain and climate. There is a gen- addition, a zone of permafrost stretched southward from eral correspondence between glacials in different regions. the edge of the glacial sheet, a few hundred kilometres Investigators often interchange the names if the glacial in North America, and several hundred in Eurasia. The geology of a region is in the process of being defined. 3.2. PALEOGEOGRAPHY AND CLIMATE 29

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

Milankovitch cycles

Main article: Milankovitch cycles

Glaciation in the Pleistocene was a series of glacials and interglacials, stadials and interstadials, mirroring periodic changes in climate. The main factor at work in climate

Ice ages as reflected in atmospheric CO2, stored in bubbles from cycling is now believed to be Milankovitch cycles. These glacial ice of Antarctica. are periodic variations in regional and planetary solar radiation reaching the Earth caused by several repeating changes in the Earth’s motion. However, it is generally incorrect to apply the name of a glacial in one region to another. Milankovitch cycles cannot be the sole factor responsible for the variations in climate since they explain neither For most of the 20th century only a few regions had been the long term cooling trend over the Plio-Pleistocene, nor studied and the names were relatively few. Today the ge- the millennial variations in the Greenland Ice Cores. Mi- ologists of different nations are taking more of an interest lankovitch pacing seems to best explain glaciation events in Pleistocene glaciology. As a consequence, the num- with periodicity of 100,000, 40,000, and 20,000 years. ber of names is expanding rapidly and will continue to Such a pattern seems to fit the information on climate expand. Many of the advances and stadials remain un- change found in oxygen isotope cores. The timing of named. Also, the terrestrial evidence for some of them our present interglacial interval (known as the Holocene, has been erased or obscured by larger ones, but evidence Postglacial, or the Present Interglacial) to that of the pre- remains from the study of cyclical climate changes. vious interglacial, beginning about 130,000 years ago The glacials in the following tables show historical us- (The Eemian Interglacial), suggests that the next glacial ages, are a simplification of a much more complex cycle would likely begin in about 3,000 years. of variation in climate and terrain, and are generally no longer used. These names have been abandoned in favor Oxygen isotope ratio cycles of numeric data because many of the correlations were found to be either inexact or incorrect and more than four Main article: Oxygen isotope ratio cycle major glacials have been recognized since the historical terminology was established.[11][12][13] In oxygen isotope ratio analysis, variations in the ratio of Corresponding to the terms glacial and interglacial, the 18 terms pluvial and interpluvial are in use (Latin: pluvia, O to 16 rain). A pluvial is a warmer period of increased rainfall; O (two isotopes of oxygen) by mass (measured by a mass an interpluvial, of decreased rainfall. Formerly a pluvial spectrometer) present in the calcite of oceanic core sam- was thought to correspond to a glacial in regions not iced, ples is used as a diagnostic of ancient ocean temperature and in some cases it does. Rainfall is cyclical also. Plu- change and therefore of climate change. Cold oceans are vials and interpluvials are widespread. richer in 18 There is no systematic correspondence of pluvials to O, which is included in the tests of the microorganisms glacials, however. Moreover, regional pluvials do not cor- (foraminifera) contributing the calcite. respond to each other globally. For example, some have A more recent version of the sampling process makes use used the term “Riss pluvial” in Egyptian contexts. Any of modern glacial ice cores. Although less rich in 18 coincidence is an accident of regional factors. Only a few O than sea water, the snow that fell on the glacier year by of the names for pluvials in restricted regions have been year nevertheless contained 18 strategraphically defined. O and 16 O in a ratio that depended on the mean annual tempera- 3.2.3 Palaeocycles ture. Temperature and climate change are cyclical when plot- The sum of transient factors acting at the Earth’s surface ted on a graph of temperature versus time. Temperature is cyclical: climate, ocean currents and other movements, coordinates are given in the form of a deviation from to- wind currents, temperature, etc. The waveform response 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 the ratio found in standard mean ocean water (SMOW). The graph in either form appears as a waveform with overtones. One half of a period is a Marine isotopic stage (MIS). It indicates a glacial (below zero) or an interglacial (above zero). Overtones are stadials or interstadials. According to this evidence, Earth experienced 102 MIS stages beginning at about 2.588 Ma BP in the Early Pleis- tocene Gelasian. Early Pleistocene stages were shallow and frequent. The latest were the most intense and most widely spaced.

By convention, stages are numbered from the Holocene, Pleistocene of South America showing Megatherium and two which is MIS1. Glacials receive an even number; inter- Glyptodon. glacials, odd. The ﬁrst major glacial was MIS2-4 at about 85–11 ka BP. The largest glacials were 2, 6, 12, and 16; the warmest interglacials, 1, 5, 9 and 11. For matching had replaced the megafauna and migrated north. of MIS numbers to named stages, see under the articles The extinctions were especially severe in North America for those names. where native horses and camels were eliminated.

• Asian land mammal ages (ALMA) include 3.3 Fauna Zhoukoudianian, Nihewanian, and Yushean. • European land mammal ages (ELMA) include See also: Quaternary extinction event Gelasian (2.5—1.8 Ma).

Both marine and continental faunas were essentially mod- • North American land mammal ages (NALMA) in- ern and many animals, specifically, mammals were much clude Blancan (4.75–1.8), Irvingtonian (1.8–0.24) larger in body form than their modern relatives . and Rancholabrean (0.24–0.01) in millions of years. The Blancan extends significantly back into the Pliocene. • South American land mammal ages (SALMA) include Uquian (2.5–1.5), Ensenadan (1.5–0.3) and Lujanian (0.3–0.01) in millions of years. The Uquian previously extended significantly back into the Pliocene, although the new definition places it entirely within the Pleistocene.

Pleistocene of Northern Spain showing woolly mammoth, cave 3.4 Humans lions eating a reindeer, tarpans, and woolly rhinoceros.

The severe climatic changes during the ice age had ma- Main articles: Human evolution, Paleolithic and Models jor impacts on the fauna and ﬂora. With each advance of of migration to the New World the ice, large areas of the continents became totally de- populated, and plants and animals retreating southward Scientiﬁc evidence[14] indicates that humans evolved into in front of the advancing glacier faced tremendous stress. their present form during the Pleistocene.[15] In the be- The most severe stress resulted from drastic climatic ginning of the Pleistocene Paranthropus species are still changes, reduced living space, and curtailed food supply. present, as well as early human ancestors, but during the A major extinction event of large mammals (megafauna), lower Palaeolithic they disappeared, and the only hominid which included mammoths, mastodons, saber-toothed species found in fossilic records is Homo erectus for much cats, glyptodons, ground sloths, Irish elk, cave bears, and of the Pleistocene. The Middle and late Palaeolithic saw short-faced bears, began late in the Pleistocene and con- the appearance of new types of humans, as well as the tinued into the Holocene. Neanderthals also became ex- development of more elaborate tools than found in previ- tinct during this period. At the end of the last ice age, ous eras. According to mitochondrial timing techniques, cold-blooded animals, smaller mammals like wood mice, modern humans migrated from Africa after the Riss migratory birds, and swifter animals like whitetail deer glaciation in the middle Palaeolithic during the Eemian 3.7. REFERENCES 31

Stage, spreading all over the ice-free world during the late [7] Svensson, A., S. W. Nielsen, S. Kipfstuhl, S. J. Johnsen, Pleistocene.[16][17][18] A 2005 study posits that humans J. P. Steﬀensen, M. Bigler, U. Ruth, and R. Röthlisberger in this migration interbred with archaic human forms al- (2005) “Visual stratigraphy of the North Greenland Ice ready outside of Africa by the late Pleistocene, incorpo- Core Project (NorthGRIP) ice core during the last glacial rating archaic human genetic material into the modern period” Journal of Geophysical Research 110: (D02108) human gene pool.[19] [8] Gradstein, Felix M.; Ogg, James G. and Smith, A. Gilbert (eds.) (2005) A Geologic Time Scale 2004 Cambridge University Press, Cambridge, UK, p. 28, ISBN 0-521- 3.5 Deposits 78142-6

Pleistocene non-marine sediments are found primarily [9] Rio, D.; Sprovieri, R.; Castradori, D. and Di Stefano, E. (1998) “The Gelasian Stage (Upper Pliocene): a new unit in ﬂuvial deposits, lakebeds, slope and loess deposits as of the global standard chronostratigraphic scale” Episodes well as in the large amounts of material moved about by 21: pp. 82-87 glaciers. Less common are cave deposits, travertines and volcanic deposits (lavas, ashes). Pleistocene marine de- [10] National Geographic Channel, Six Degrees Could Change posits are found primarily in shallow marine basins mostly The World, Mark Lynas interview. Retrieved February (but with important exceptions) in areas within a few tens 14, 2008. of kilometers of the modern shoreline. In a few geologically active areas such as the Southern California coast, [11] Richmond, G.M. and D.S. Fullerton, 1986, Summation Pleistocene marine deposits may be found at elevations of Quaternary glaciations in the United States of America. of several hundred meters. 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- 3.6 See also netism of late Cenozoic deposits of the north-central United States, PDF version, 1.2 MB. Geological Society of Amer- • Climate state ica Bulletin.116(1-2): pp. 30-41; doi:10.1130/B25325.1

• Geologic time scale [13] Aber, J.S. (1991) “Glaciations of Kansas” Boreas 20(4): pp. 297-314 - (contains a summary of how and why the • Pleistocene megafauna Nebraskan, Aftonian, Kansan, and Yarmouthian stages were abandoned by modern stratigraphers). • Timeline of glaciation [14] Rogers, A.R. and Jorde, L.B. (1995) “Genetic evidence on modern human origins” Human Biology 67: pp. 1–36 3.7 References [15] Wall, J.D. and Przeworski, M. (2000) “When did the [1] “Geologic Age Symbol Font (StratagemAge)". USGS. human population start increasing?" Genetics 155: pp. 99-430. Retrieved 2011-06-22. 1865–1874

[2] “Pleistocene”. Online Etymology Dictionary. [16] Cann, R.L.; Stoneking, M. and Wilson, A.C.(1987) “Mi- tochondrial DNA and human evolution” Nature 325: pp. [3] Gibbard, P. and van Kolfschoten, T. (2004) “The Pleis- 31–36 tocene and Holocene Epochs” Chapter 22 PDF (3.1 MB) In Gradstein, F. M., Ogg, James G., and Smith, A. Gilbert [17] Stringer, C.B. (1992) “Evolution of early modern hu- (eds.), A Geologic Time Scale 2004 Cambridge University mans” In: Jones, Steve; Martin, R. and Pilbeam, David Press, Cambridge, ISBN 0-521-78142-6 R. (eds.) (1992) The Cambridge encyclopedia of human evolution Cambridge University Press, Cambridge, ISBN [4] For the top of the series, see: Lourens, L., Hilgen, F., 0-521-32370-3, pp. 241–251. Shackleton, N.J., Laskar, J., Wilson, D., (2004) “The Neogene Period”. In: Gradstein, F., Ogg, J., Smith, A.G. [18] Templeton, A. (2002) “Out of Africa again and again” Na- (Eds.), A Geologic Time Scale 2004. Cambridge: Cam- ture 416: p. 45 bridge University Press.

[5] Moore, Mark; Brumm (2007). “Stone artifacts and ho- [19] Eswarana, Vinayak; Harpendingb, Henry and Rogers, minins in island Southeast Asia: New insights from Flo- Alan R. (2005) “Genomics refutes an exclusively African res, eastern Indonesia”. Journal of Human Evolution 52: origin of humans” Journal of Human Evolution 49(1): pp. 88. Retrieved 10 April 2014. 1–18 Abstract

[6] Riccardi, Alberto C. (30 June 2009) “IUGS ratified ICS Recommendation on redefinition of Pleistocene and for- • Ogg, Jim; June, 2004, Overview of Global mal definition of base of Quaternary” International Union Boundary Stratotype Sections and Points (GSSP’s, of Geological Sciences Stratigraphy.org, Accessed April 30, 2006. 32 CHAPTER 3. PLEISTOCENE

3.8 External links

• The SMU-in-Taos Research Publications digital collection of anthropological and archaeological monographs contains Late Pleistocene environments of the southern high plains. • Pleistocene Microfossils: 50+ images of Foraminifera Chapter 4

Prehistoric mammal

4.2 See also

• Cynodont • List of extinct mammals

• Mammaliaformes • Megamammals

• Pleistocene extinctions • Pleistocene megafauna

• Synapsid • Therapsid

An early drawing depicting prehistoric mammals

Prehistoric mammals are groups of mammals that became extinct before humans developed writing. 164 million years ago, in the Jurassic period, Castorocauda lutrasimilis, a mammaliaform (mammal-shaped) animal weighing about 500 grams (1.1 lb), had a full mammalian pelt, with guard hairs and underfur, webbed feet, and scales on the tail like a modern beaver, as well as teeth specialized for catching ﬁsh. Later, about 130 million years ago in the Cretaceous, there existed larger mammals; a fossil of Repenomamus giganticus indicates that the animal was about 1 me- ter (3¼ ft) long. In the stomach of a smaller cousin, Repenomamus robustus at 52 cm (20½ in), the remains of a juvenile dinosaur have been preserved. The lineages of many varieties continued through the Cenozoic era where some reached very large sizes. Most of the very large mammals became extinct in the last ice age, but have smaller descendants.

4.1 List of prehistoric mammals

Main article: List of prehistoric mammals

Prehistoric mammals include:

33 Chapter 5

Stone Age

For other uses, see Stone Age (disambiguation). 5.1 Historical signiﬁcance The Stone Age is a broad prehistoric period during The Stone Age is contemporaneous with the evolution of the genus Homo, the only exception possibly being at the very beginning, when species prior to Homo may have manufactured tools. According to the age and location of the current evidence, the cradle of the genus is the East African Rift System, especially toward the north in Ethiopia, where it is bordered by grasslands. The closest relative among the other living Primates, the genus Pan, represents a branch that continued on in the deep forest, where the primates evolved. The rift served as a conduit for movement into southern Africa and also north down the Nile into North Africa and through the continuation of the rift in the Levant to the vast grasslands of Asia. Starting from about 3 million years ago (mya) a single biome established itself from South Africa through the rift, North Africa, and across Asia to modern China, which has been called “transcontinental 'savannahstan'" recently.[2] Starting in the grasslands of the rift, Homo erectus, the predecessor of modern humans, found an Modern Awash River, Ethiopia, descendant of the Palaeo- ecological niche as a tool-maker and developed a de- Awash, source of the sediments in which the oldest Stone Age pendence on it, becoming a “tool equipped savanna tools have been found dweller.”[3] which stone was widely used to make implements with a sharp edge, a point, or a percussion surface. The period lasted roughly 3.4 million years, and ended between 6000 5.2 The Stone Age in archaeology BCE and 2000 BCE with the advent of metalworking.[1] Stone Age artifacts include tools used by humans and by their predecessor species in the genus Homo, as well as the 5.2.1 Beginning of the Stone Age earlier partly contemporaneous genera Australopithecus and Paranthropus. Bone tools were used during this pe- During 2010, fossilised animal bones bearing marks from riod as well but are rarely preserved in the archaeological stone tools were found in the Lower Awash Valley in record. The Stone Age is further subdivided by the types Ethiopia. Discovered by an international team led by of stone tools in use. Shannon McPherron, at 3.4 million years old they are the oldest evidence of stone tool use ever found anywhere in The Stone Age is the ﬁrst of the three-age system the world.[1] of archaeology, which divides human technological prehistory into three periods: The oldest known stone tools have been excavated from several sites at Gona, Ethiopia, on the sediments of the paleo-Awash River, which serve to date them. All the • The Stone Age tools come from the Busidama Formation, which lies • The Bronze Age above a disconformity, or missing layer, which would have been from 2.9 to 2.7 mya. The oldest sites contain- • The Iron Age ing tools are dated to 2.6–2.55 mya.[4] One of the most

34 5.2. THE STONE AGE IN ARCHAEOLOGY 35

the initial period of the Bronze Age and is unquestionably part of the Age of Metals. The Bronze Age was followed by the Iron Age. During this entire time stone remained in use in parallel with the metals for some objects, including those also used in the Neolithic, such as stone pottery. The transition out of the Stone Age occurred between 6000 BCE and 2500 BCE for much of humanity living in North Africa and Eurasia. The first evidence of human metallurgy dates to between the 5th and 6th millennium BCE in the archaeological sites of Majdanpek, Yarmovac and Pločnik (a copper axe from 5500 BCE belonging to the Vinca culture), though not conventionally considered part of the Chalcolithic or “Copper Age”, this provides the earliest known example of copper metallurgy.[8] and the Rudna Glava mine in Serbia. Ötzi the Iceman, a mummy from about 3300 BCE carried with him a copper axe and a flint knife. In regions such as Subsaharan Africa, the Stone Age was followed directly by the Iron Age. The Middle East and southeastern Asian regions progressed past Stone Age technology around 6000 BCE. Europe, and the rest of Asia became post–Stone Age societies by about 4000 BCE. The proto-Inca cultures of South America continued at a Stone Age level until around 2000 BCE, when Obsidian projectile point gold, copper and silver made their entrance, the rest following later. Australia remained in the Stone Age until the 17th century. Stone tool manufacture continued. In striking circumstances about these sites is that they are Europe and North America, millstones were in use until from the Late Pliocene, where previous to their discovery well into the 20th century, and still are in many parts of tools were thought to have evolved only in the Pleistocene. the world. Rogers and Semaw, excavators at the locality, point out that:[5] 5.2.3 The concept of Stone Age "...the earliest stone tool makers were skilled flintknappers .... The possible reasons behind The terms was never meant to suggest that advancement this seeming abrupt transition from the absence and time periods in prehistory are only measured by the of stone tools to the presence thereof include ... type of tool material, rather than, for example, social or- gaps in the geological record.” ganization, food sources exploited, adaptation to climate, adoption of agriculture, cooking, settlement and religion. The excavators are confident that more tools will be Like pottery, the typology of the stone tools combined found elsewhere from 2.9 mya. The species who with the relative sequence of the types in various regions made the Pliocene tools remains unknown. Fragments provide a chronological framework for the evolution of of Australopithecus garhi, Australopithecus aethiopicus[6] man and society. They serve as diagnostics of date, rather and Homo, possibly Homo habilis, have been found in than characterizing the people or the society. [7] sites near the age of the oldest tools. Lithic analysis is a major and specialised form of archaeological investigation. It involves the measurement 5.2.2 End of the Stone Age of the stone tools to determine their typology, function and the technology involved. It includes scientific study Innovation of the technique of smelting ore ended the of the lithic reduction of the raw materials, examining Stone Age and began the Bronze Age. The first most how the artifacts were made. Much of this study takes significant metal manufactured was bronze, an alloy of place in the laboratory in the presence of various special- copper and tin, each of which was smelted separately. ists. In experimental archaeology, researchers attempt to The transition from the Stone Age to the Bronze Age was create replica tools, to understand how they were made. a period during which modern people could smelt cop- Flintknappers are craftsmen who use sharp tools to re- per, but did not yet manufacture bronze, a time known duce flintstone to flint tool. as the Copper Age, or more technically the Chalcolithic, In addition to lithic analysis, the field prehistorian uti- “copper-stone” age. The Chalcolithic by convention is lizes a wide range of techniques derived from multiple 36 CHAPTER 5. STONE AGE

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

5.2.4 The three-stage system

In the 1920s, South African archaeologists organizing the stone tool collections of that country observed that they A variety of stone tools did not ﬁt the newly detailed Three-Age System. In the words of J. Desmond Clark,[10]

fields. The work of the archaeologist in determining the paleocontext and relative sequence of the layers is supple- “It was early realized that the threefold di- mented by the efforts of the geologic specialist in identi- vision of culture into Stone, Bronze and Iron fying layers of rock over geologic time, of the paleonto- Ages adopted in the nineteenth century for Eu- logical specialist in identifying bones and animals, of the rope had no validity in Africa outside the Nile palynologist in discovering and identifying plant species, valley.” of the physicist and chemist in laboratories determining dates by the carbon-14, potassium-argon and other meth- Consequently they proposed a new system for Africa, the ods. Study of the Stone Age has never been mainly about Three-stage System. Clark regarded the Three-age Sys- stone tools and archaeology, which are only one form of tem as valid for North Africa; in sub-Saharan Africa, the evidence. The chief focus has always been on the society Three-stage System was best.[11] In practice, the failure and the physical people who belonged to it. of African archaeologists either to keep this distinction Useful as it has been, the concept of the Stone Age has its in mind, or to explain which one they mean, contributes limitations. The date range of this period is ambiguous, to the considerable equivocation already present in the lit- disputed, and variable according to the region in ques- erature. There are in effect two Stone Ages, one part of tion. While it is possible to speak of a general 'stone the Three-age and the other constituting the Three-stage. age' period for the whole of humanity, some groups never They refer to one and the same artifacts and the same developed metal-smelting technology, so remained in a technologies, but vary by locality and time. 'stone age' until they encountered technologically devel- The Three-stage System was proposed in 1929 by Astley oped cultures. The term was innovated to describe the John Hilary Goodwin, a professional archaeologist, and archaeological cultures of Europe. It may not always be Clarence van Riet Lowe, a civil engineer and amateur ar- the best in relation to regions such as some parts of the chaeologist, in an article titled “Stone Age Cultures of Indies and Oceania, where farmers or hunter-gatherers South Africa” in the journal Annals of the South African used stone for tools until European colonisation began. Museum. By then, the dates of the Early Stone Age, or The archaeologists of the late 19th and early 20th cen- Paleolithic, and Late Stone Age, or Neolithic (neo = new), turies CE, who adapted the three-age system to their were fairly solid and were regarded by Goodwin as abso- ideas, hoped to combine cultural anthropology and ar- lute. He therefore proposed a relative chronology of peri- chaeology in such a way that a specific contemporaneous ods with floating dates, to be called the Earlier and Later tribe can be used to illustrate the way of life and beliefs Stone Age. The Middle Stone Age would not change its of the people exercising a specific Stone-Age technology. name, but it would not mean Mesolithic.[12] As a description of people living today, the term stone age The duo thus reinvented the Stone Age. In Sub-Saharan is controversial. The Association of Social Anthropolo- [9] Africa, however, it was ended by the intrusion of the Iron gists discourages this use, asserting: Age from the north. The Neolithic and the Bronze Age never occurred. Moreover, the technologies included in “To describe any living group as 'primitive' those 'stages’, as Goodwin called them, were not exactly or 'Stone Age' inevitably implies that they are the same. Since then, the original relative terms have be- living representatives of some earlier stage of come identified with the technologies of the Paleolithic human development that the majority of hu- and Mesolithic, so that they are no longer relative. More- mankind has left behind. For some, this could over, there has been a tendency to drop the comparative be a positive description, implying, for exam- degree in favor of the positive: resulting in two sets of ple, that such groups live in greater harmony Early, Middle and Late Stone Ages of quite different con- with nature .... For others, ... 'primitive' is tent and chronologies. 5.3. CHRONOLOGY 37

By voluntary agreement, archaeologists respect the deci- but were oﬃcially rejected in 1965 (again on an advi- sions of the Pan-African Congress of Prehistory, which sory basis) by Burg Wartenstein Conference #29, Sys- meets every four years to resolve archaeological business tematic Investigation of the African Later Tertiary and brought before it. Delegates are actually international; the Quaternary,[15] a conference in anthropology held by the organization takes its name from the topic. Louis Leakey Wenner-Gren Foundation, at Burg Wartenstein Castle, hosted the ﬁrst one in Nairobi in 1947. It adopted Good- which it then owned in Austria, attended by the same win and Lowe’s 3-stage system at that time, the stages to scholars that attended the Pan African Congress, includ- be called Early, Middle and Later. ing Louis Leakey and Mary Leakey, who was delivering a pilot presentation of her typological analysis of Early Stone Age tools, to be included in her 1971 contribution 5.2.5 The problem of the transitions to Olduvai Gorge, “Excavations in Beds I and II, 1960– 1963.”[16] The problem of the transitions in archaeology is a branch However, although the intermediate periods were gone, of the general philosophic continuity problem, which the search for the transitions continued. examines how discrete objects of any sort that are contiguous in any way can be presumed to have a relationship of any sort. In archaeology, the relationship is 5.3 Chronology one of causality. If Period B can be presumed to descend from Period A, there must be a boundary between A and

B, the A–B boundary. The problem is in the nature of 2 2 41 kyr cycle 100 kyr cycle this boundary. If there is no distinct boundary, then the 0 2.5

population of A suddenly stopped using the customs char- -2 3 -4 Five Million Years of acteristic of A and suddenly started using those of B, an 3.5 Equivalent 18

-6 Climate Change δ O Benthic unlikely scenario in the process of evolution. More real- Vostok Δ T (°C) 4 istically, a distinct border period, the A/B transition, ex- -8 From Sediment Cores Carbonate (per mil) 4.5 isted, in which the customs of A were gradually dropped Millions of Years Ago and those of B acquired. If transitions do not exist, then there is no proof of any continuity between A and B. Time series plot of temperature over the previous 5 million years

The Stone Age of Europe is characteristically in deficit In 1859 Jens Jacob Worsaae first proposed a division of of known transitions. The 19th and early 20th-century the Stone Age into older and younger parts based on his innovators of the modern three-age system recognized work with Danish kitchen middens that began in 1851.[17] the problem of the initial transition, the “gap” between In the subsequent decades this simple distinction devel- the Paleolithic and the Neolithic. Louis Leakey provided oped into the archaeological periods of today. The major something of an answer by proving that man evolved in subdivisions of the Three-age Stone Age cross two epoch Africa. The Stone Age must have begun there to be car- boundaries on the geologic time scale: ried repeatedly to Europe by migrant populations. The different phases of the Stone Age thus could appear there • without transitions. The burden on African archaeolo- The geologic Pliocene–Pleistocene boundary gists became all the greater, because now they must find (highly glaciated climate) the missing transitions in Africa. The problem is difficult • The Paleolithic period of archaeology and ongoing. After its adoption by the First Pan African Congress in • The geologic Pleistocene–Holocene boundary 1947, the Three-Stage Chronology was amended by the (modern climate) Third Congress in 1955 to include a First Intermedi- • ate Period between Early and Middle, to encompass the Mesolithic or Epipaleolithic period of archae- Fauresmith and Sangoan technologies, and the Second In- ology termediate Period between Middle and Later, to encom- • Neolithic period of archaeology pass the Magosian technology and others. The chrono- logic basis for definition was entirely relative. With the The succession of these phases varies enormously from arrival of scientific means of finding an absolute chronol- one region (and culture) to another. 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, 5.3.1 Three-age chronology while Sangoan is a facies of Lupemban.[13] Magosian is [14] “an artificial mix of two different periods.” Main articles: Paleolithic, Human evolution and Three- Once seriously questioned, the intermediates did not wait age system for the next Pan African Congress two years hence, 38 CHAPTER 5. STONE AGE

The Paleolithic or Palaeolithic (from Greek: παλαιός, palaios, “old"; and λίθος, lithos, “stone” lit. “old stone,” coined by archaeologist John Lubbock and published in 1865) is the earliest division of the Stone Age. It covers the greatest portion of 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 the ﬁrst documented use of stone tools by hominans such as Homo habilis, to the end of the Pleistocene around 10,000 BCE.[18] The Paleolithic era ended with the Mesolithic, or in areas with an early neolithisation, the Epipaleolithic.

Lower Paleolithic This is a Mode 1, or Oldowan, stone tool from the western Sa- hara. Main article: Lower Paleolithic

nal stone is called a core; the resultant pieces, flakes. Typ- At sites dating from the Lower Paleolithic Period (about ically, but not necessarily, small pieces are detached from 2,500,000 to 200,000 years ago), simple pebble tools a larger piece, in which case the larger piece may be called have been found in association with the remains of what the core and the smaller pieces the flakes. The prevalent may have been the earliest human ancestors. A somewhat usage, however, is to call all the results flakes, which can more sophisticated Lower Paleolithic tradition, known as be confusing. A split in half is called bipolar flaking. the Chopper chopping-tool industry, is widely distributed in the Eastern Hemisphere. This tradition is thought Consequently the method is often called “core-and- to have been the work of the hominin species named flake”. More recently, the tradition has been called “small Homo erectus. Although no such fossil tools have yet flake” since the flakes were small compared to subsequent been found, it is believed that H. erectus probably made Acheulean tools.[20] tools of wood and bone as well as stone. About 700,000 years ago, a new Lower Paleolithic tool, the hand ax, ap- “The essence of the Oldowan is the making peared. The earliest European hand axes are assigned and often immediate use of small flakes.” to the Abbevillian industry, which developed in northern France in the valley of the Somme River; a later, more re- Another naming scheme is “Pebble Core Technology fined hand-ax tradition is seen in the Acheulian industry, (PBC)":[21] evidence of which has been found in Europe, Africa, the Middle East, and Asia. Some of the earliest known hand “Pebble cores are ... artifacts that have axes were found at Olduvai Gorge (Tanzania) in associ- been shaped by varying amounts of hard- ation with remains of H. erectus. Alongside the hand- hammer percussion.” ax tradition there developed a distinct and very different stone-tool industry, based on flakes of stone: special tools were made from worked (carefully shaped) flakes of Various refinements in the shape have been called chop- flint. In Europe, the Clactonian industry is one example pers, discoids, polyhedrons, subspheroid, etc. To date no [22] of a flake tradition. The early flake industries probably reasons for the variants have been ascertained: contributed to the development of the Middle Paleolithic flake tools of the Mousterian industry, which is associated “From a functional standpoint, pebble with the remains of Neanderthal man.[19] cores seem designed for no specific purpose.”

However, they would not have been manufactured for no Oldowan in Africa Main article: Oldowan purpose:[22]

The earliest documented stone tools were found in East “Pebble cores can be useful in many cut- Africa, manufacturers unknown. They belonged to an ting, scraping or chopping tasks, but ... they industry now known as Oldowan, after the type site of are not particularly more eﬃcient in such tasks Olduvai Gorge in Tanzania; however, sites in Ethiopia than a sharp-edged rock ....” later proved to be older. The tools were formed by knocking pieces oﬀ a river peb- The whole point of their utility is that each is a “sharp- ble, or stones like it, with a hammerstone to obtain large edged rock” in locations where nature has not provided and small pieces with one or more sharp edges. The origi- any. There is additional evidence that Oldowan, or Mode 5.3. CHRONOLOGY 39

1, tools were utilized in “percussion technology"; that is, temporaneously in the same regions H. habilis inherited they were designed to be gripped at the blunt end and the tools around 2.3 mya. At about 1.9 mya H. erectus strike something with the edge, from which use they were came on stage and lived contemporaneously with the oth- given the name of choppers. Modern science has been ers. Mode 1 was now being shared by a number of Hom- able to detect mammalian blood cells on Mode 1 tools inans over the same ranges, presumably subsisting in dif- at Sterkfontein, Member 5 East, in South Africa. As the ferent niches, but the archaeology is not precise enough blood must have come from a fresh kill, the tool users to say which. are likely to have done the killing and used the tools for butchering. Plant residues bonded to the silicon of some tools confirm the use to chop plants.[23] Oldowan out of Africa Tools of the Oldowan tradition first came to archaeological attention in Europe, Although the exact species authoring the tools remains where, being intrusive and not well defined, compared unknown, Mode 1 tools in Africa were manufactured and to the Acheulean, they were puzzling to archaeologists. used predominantly by Homo habilis. They cannot be said The mystery would be elucidated by African archaeol- to have developed these tools or to have contributed the ogy at Olduvai, but meanwhile, in the early 20th century, tradition to technology. They continued a tradition of yet the term “Pre-Acheulean” came into use in climatology. unknown origin. As chimpanzees sometimes naturally C.E.P, Brooks, a British climatologist working in the use percussion to extract or prepare food in the wild, and United States, used the term to describe a “chalky boulder may use either unmodified stones or stones that they have clay” underlying a layer of gravel at Hoxne, central Eng- split, creating an Oldowan tool, the tradition may well be land, where Acheulean tools had been found.[31] Whether far older than its current record. any tools would be found in it and what type was not Towards the end of Oldowan in Africa a new species ap- known. Hugo Obermaier, a contemporary German ar- peared over the range of Homo habilis: Homo erectus. chaeologist working in Spain, quipped: The earliest “unambiguous” evidence is a whole cranium, KNM-ER 3733 (a find identifier) from Koobi Fora in “Unfortunately, the stage of human indus- Kenya, dated to 1.78 mya.[24] An early skull fragment, try which corresponds to these deposits cannot KNM-ER 2598, dated to 1.9 mya, is considered a good be positively identified. All we can say is that candidate also.[25] Transitions in paleoanthropology are it is pre-Acheulean....” always hard to find, if not impossible, but based on the “long-legged” limb morphology shared by H. habilis and H. rudolfensis in East Africa, an evolution from one of This uncertainty was clarified by the subsequent excava- those two has been suggested.[26] tions at Olduvai; nevertheless, the term is still in use for pre-Acheulean contexts, mainly across Eurasia, that are The most immediate cause of the new adjustments ap- yet unspecified or uncertain but with the understanding pears to have been an increasing aridity in the region and that they are or will turn out to be pebble-tool.[32] consequent contraction of parkland savanna, interspersed with trees and groves, in favor of open grassland, dated There are ample associations of Mode 2 with H. erectus in 1.8–1.7 mya. During that transitional period the percent- Eurasia. H. erectus – Mode 1 associations are scantier but age of grazers among the fossil species increased from they do exist, especially in the Far East. One strong piece 15–25% to 45%, dispersing the food supply and requir- of evidence prevents the conclusion that only H. erectus ing a facility among the hunters to travel longer distances reached Eurasia: at Yiron, Israel, Mode 1 tools have been comfortably, which H. erectus obviously had.[27] The ul- found dating to 2.4 mya,[33] about 0.5 my earlier than the timate proof is the “dispersal” of H. erectus “across much known H. erectus finds. If the date is correct, either an- of Africa and Asia, substantially before the development other Hominan preceded H. erectus out of Africa or the of the Mode 2 technology and use of fire ....”[26] H. erectus earliest H. erectus has yet to be found. carried Mode 1 tools over Eurasia. After the initial appearance at Gona in Ethiopia at 2.7 According to the current evidence (which may change at mya, pebble tools date from 2.0 mya at Sterkfontein, any time) Mode 1 tools are documented from about 2.6 Member 5, South Africa, and from 1.8 mya at El Kherba, mya to about 1.5 mya in Africa,[28] and to 0.5 mya outside Algeria, North Africa. The manufacturers had already of it.[29] The genus Homo is known from H. habilis and left pebble tools at Yiron, Israel, at 2.4 mya, Riwat, Pak- H. rudolfensis from 2.3 to 2.0 mya, with the latest habilis istan, at 2.0 mya, and Renzidong, South China, at over 2 being an upper jaw from Koobi Fora, Kenya, from 1.4 mya.[34] The identification of a fossil skull at Mojokerta, mya. H. erectus is dated 1.8–0.6 mya.[30] Pernung Peninsula on Java, dated to 1.8 mya, as H. erectus, suggests that the African finds are not the earliest to According to this chronology Mode 1 was inher- be found in Africa, or that, in fact, erectus did not origi- ited by Homo from unknown Hominans, probably nate in Africa after all but on the plains of Asia.[26] The Australopithecus and Paranthropus, who must have con- outcome of the issue waits for more substantial evidence. tinued on with Mode 1 and then with Mode 2 until their Erectus was found also at Dmanisi, Georgia, from 1.75 extinction no later than 1.1 mya. Meanwhile living con- mya in association with pebble tools. 40 CHAPTER 5. STONE AGE

Pebble tools are found the latest first in southern Europe at Kokiselei, West Turkana, Kenya.[25] At Sterkfontein, and then in northern. They begin in the open areas of Italy South Africa, they are in Member 5 West, 1.7–1.4 and Spain, the earliest dated to 1.6 mya at Pirro Nord, mya.[23] The 1.7 is a fairly certain, fairly standard date. Italy. The mountains of Italy are rising at a rapid rate Mode 2 is often found in association with H. erectus. It in the framework of geologic time; at 1.6 mya they were makes sense that the most advanced tools should have lower and covered with grassland (as much of the high- been innovated by the most advanced Hominan; conse- lands still are). Europe was otherwise mountainous and quently, they are typically given credit for the innovation. covered over with dense forest, a formidable terrain for A Mode 2 tool is a biface consisting of two concave warm-weather savanna dwellers. Similarly there is no ev- surfaces intersecting to form a cutting edge all the way idence that the Mediterranean was passable at Gibraltar around, except in the case of tools intended to feature a or anywhere else to H. erectus or earlier hominans. They point. More work and planning go into the manufacture might have reached Italy and Spain along the coasts. of a Mode 2 tool. The manufacturer hits a slab off a larger In northern Europe pebble tools are found earliest at rock to use as a blank. Then large flakes are struck off the Happisburgh, United Kingdom, from 0.8 mya. The blank and worked them into bifaces by hard-hammer per- last traces are from Kent’s Cavern, dated 0.5 mya. By cussion on an anvil stone. Finally the edge is retouched: that time H. erectus is regarded as having been ex- small flakes are hit off with a bone or wood soft ham- tinct; however, a more modern version apparently had mer to sharpen or resharpen it. The core can be either evolved, Homo heidelbergensis, who must have inherited the blank or another flake. Blanks are ported for manu- the tools.[35] He also explains the last of the Acheulean in facturing supply in places where nature has provided no Germany at 0.4 mya. suitable stone. In the late 19th and early 20th centuries archaeologists Although most Mode 2 tools are easily distinguished from worked on the assumptions that a succession of Homi- Mode 1, there is a close similarity of some Oldowan and nans and cultures prevailed, that one replaced another. some Acheulean, which can lead to confusion. Some Today the presence of multiple hominans living contem- Oldowan tools are more carefully prepared to form a poraneously near each other for long periods is accepted more regular edge. One distinguishing criterion is the as proved true; moreover, by the time the previously as- size of the flakes. In contrast to the Oldowan “small flake” sumed “earliest” culture arrived in northern Europe, the tradition, Acheulean is “large flake:" “The primary tech- rest of Africa and Eurasia had progressed to the Mid- nological distinction remaining between Oldowan and dle and Upper Palaeolithic, so that across the earth all the Acheulean is the preference for large flakes (>10 three were for a time contemporaneous. In any given re- cm) as blanks for making large cutting tools (handaxes gion there was a progression from Oldowan to Acheulean, and cleavers) in the Acheulean.”[36] “Large Cutting Tool Lower to Upper, no doubt. (LCT)" has become part of the standard terminology as well.[22]

Acheulean in Africa Main article: Acheulean In North Africa, the presence of Mode 2 remains a mys- The end of Oldowan in Africa was brought on by the tery, as the oldest finds are from Thomas Quarry in Morocco at 0.9 mya.[34] Archaeological attention, however, shifts to the Jordan Rift Valley, an extension of the East African Rift Valley (the east bank of the Jor- dan is slowly sliding northward as East Africa is thrust away from Africa). Evidence of use of the Nile Valley is in deficit, but Hominans could easily have reached the palaeo-Jordan river from Ethiopia along the shores of the Red Sea, one side or the other. A crossing would not have been necessary, but it is more likely there than over a theoretical but unproven land bridge through either Gibraltar or Sicily. Meanwhile Acheulean went on in Africa past the 1.0 mya mark and also past the extinction of H. erectus there. The last Acheulean in East Africa is at Olorgesailie, Kenya, dated to about 0.9 mya. Its owner was still H. erectus,[34] but in South Africa, Acheulean at Elandsfontein, 1.0– 0.6 mya, is associated with Saldanha man, classified as H. heidelbergensis, a more advanced, but not yet mod- An Acheulean tool, not worked over the entire surface ern, descendant most likely of H. erectus. The Thoman Quarry Hominans in Morocco similarly are most likely appearance of Acheulean, or Mode 2, stone tools. The Homo rhodesiensis,[37] in the same evolutionary status as earliest known instances are in the 1.7–1.6 mya layer 5.3. CHRONOLOGY 41

H. heidelbergensis. itation of archeology, but after 1 mya evidence not available to Movius indicates the prevalence of Acheulean. For example, the Acheulean site at Bose, China, is dated Acheulean out of Africa Mode 2 is first known out 0.803±3K mya.[46] The authors of this chronologically of Africa at 'Ubeidiya, Israel, a site now on the Jordan later East Asian Acheulean remain unknown, as does River, then frequented over the long term (hundreds of whether it evolved in the region or was brought in. thousands of years) by Homo on the shore of a variable- There is no named boundary line between Mode 1 and level palaeo-lake, long since vanished. The geology was Mode 2 on the west; nevertheless, Mode 2 is equally late created by successive “transgression and regression” of in Europe as it is in the Far East. The earliest comes the lake[38] resulting in four cycles of layers. The tools from a rock shelter at Estrecho de Quípar in Spain, dated are located in the first two, Cycles Li (Limnic Inferior) to greater than 0.9 mya. Teeth from an undetermined and Fi (Fluviatile Inferior), but mostly in Fi. The cy- Hominan were found there also.[47] The last Mode 2 in cles represent different ecologies and therefore different Southern Europe is from a deposit at Fontana Ranuccio cross-sections of fauna, which makes it possible to date near Anagni in Italy dated to 0.45 mya, which is generally them. They appear to be the same faunal assemblages as linked to Homo cepranensis, a “late variant of H. erectus,” the Ferenta Faunal Unit in Italy, known from excavations a fragment of whose skull was found at Ceprano nearby, at Selvella and Pieterfitta, dated to 1.6–1.2 mya.[39] dated 0.46 mya.[48] At 'Ubeidiya the marks on the bones of the animal species found there indicate that the manufacturers of the tools butchered the kills of large predators, an activity that has Middle Paleolithic been termed “scavenging.”[40] There are no living floors, nor did they process bones to obtain the marrow. These Main article: Middle Paleolithic activities cannot be understood therefore as the only or even the typical economic activity of Hominans. Their This period is best known as the era during which the interests were selective: they were primarily harvesting Neanderthals lived in Europe and the Near East (c. the meat of Cervids,[41] which is estimated to have been 300,000–28,000 years ago). Their technology is mainly available without spoiling for up to four days after the kill. the Mousterian, but Neanderthal physical characteris- The majority of the animals at the site were of tics have been found also in ambiguous association with “Palaearctic biogeographic origin.”[42] However, these the more recent Châtelperronian archeological culture overlapped in range on 30–60% of “African biogeo- in Western Europe and several local industries like the graphic origin.”[43] The biome was Mediterranean, not Szeletian in Eastern Europe/Eurasia. There is no evi- savanna. The animals were not passing through; there dence for Neanderthals in Africa, Australia or the Amer- was simply an overlap of normal ranges. Of the Homi- icas. nans, H. erectus left several cranial fragments. Teeth of Neanderthals nursed their elderly and practised ritual undetermined species may have been H. ergaster.[44] The burial indicating an organised society. The earliest evi- tools are classified as “Lower Acheulean” and “Developed dence (Mungo Man) of settlement in Australia dates to Oldowan.” The latter is a disputed classification created around 40,000 years ago when modern humans likely by Mary Leakey to describe an Acheulean-like tradition crossed from Asia by island-hopping. Evidence for sym- in Bed II at Olduvai. It is dated 1.53–1.27 mya. The date bolic behavior such as body ornamentation and burial is of the tools therefore probably does not exceed 1.5 mya; ambiguous for the Middle Paleolithic and still subject to 1.4 is often given as a date. This chronology, which is debate. The Bhimbetka rock shelters exhibit the earliest definitely later than in Kenya, supports the “out of Africa” traces of human life in India, some of which are approx- hypothesis for Acheulean, if not for the Hominans. imately 30,000 years old. From Southwest Asia, as the Levant is now called, the Acheulean extended itself more slowly eastward, arriving at Isampur, India, about 1.2 mya. It does not appear in Upper Paleolithic China and Korea until after 1mya and not at all in Indone- sia. There is a discernible boundary marking the furthest Main article: Upper Paleolithic extent of the Acheulean eastward before 1 mya, called the Movius Line, after its proposer, Hallam L. Movius. On From 50,000 to 10,000 years ago in Europe, the Up- the east side of the line the small flake tradition contin- per Paleolithic ends with the end of the Pleistocene and ues, but the tools are additionally worked Mode 1, with onset of the Holocene era (the end of the last ice age). flaking down the sides. In Athirampakkam at Chennai in Modern humans spread out further across the Earth dur- Tamil Nadu the Acheulean age started at 1.51 mya and it [45] ing the period known as the Upper Paleolithic. The Up- is also prior than North India and Europe. per Paleolithic is marked by a relatively rapid succession The cause of the Movius Line remains speculative, of often complex stone artifact technologies and a large whether it represents a real change in technology or a lim- increase in the creation of art and personal ornaments. 42 CHAPTER 5. STONE AGE

During period between 35 and 10 kya evolved: from 38 to 30 kya Châtelperronian, 40–28 Aurignacian, 28–22 Gravettian, 22–17 Solutrean, and 18–10 Magdalenian. All of these industries except the Châtelperronian are associated with anatomically modern humans. Authorship of the Châtelperronian is still the subject of much debate. The Americas were colonised via the Bering land bridge which was exposed during this period by lower sea levels. These people are called the Paleo-Indians, and the earliest accepted dates are those of the Clovis culture sites, some 13,500 years ago. Globally, societies were hunter- gatherers but evidence of regional identities begins to appear in the wide variety of stone tool types being devel- Ġgantija temples, Gozo. Some of the world’s oldest free-standing oped to suit very diﬀerent environments. structures.

Epipaleolithic/Mesolithic

Main articles: Epipaleolithic, Mesolithic

The period starting from the end of the last ice age, 10,000 years ago, to around 6,000 years ago was characterized by rising sea levels and a need to adapt to a changing environment and find new food sources. The development of Mode 5 (microlith) tools began in response to these changes. They were derived from the previous Pale- olithic tools, hence the term Epipaleolithic, or were intermediate between the Paleolithic and the Neolithic, hence the term Mesolithic (Middle Stone Age). The choice of a word depends on exact circumstances and the inclina- Skara Brae, Scotland. Europe’s most complete Neolithic village tion of the archaeologists excavating the site. Microliths were used in the manufacture of more efficient composite tools, resulting in an intensification of hunting and fish- Due to the increased need to harvest and process plants, ing and with increasing social activity the development ground stone and polished stone artifacts became much of more complex settlements, such as Lepenski Vir. Do- more widespread, including tools for grinding, cutting, mestication of the dog as a hunting companion probably and chopping. Skara Brae located on Orkney island dates to this period. off Scotland is one of Europe’s best examples of a Ne- olithic village. The community contains stone beds, The earliest known battle occurred during the Mesolithic shelves and even an indoor toilet linked to a stream. The period at a site in Egypt known as Cemetery 117. first large-scale constructions were built, including settlement towers and walls, e.g., Jericho and ceremonial sites, e.g.: Stonehenge. The Ġgantija temples of Gozo Neolithic in the Maltese archipelago are the oldest surviving free Main article: Neolithic standing structures in the world, erected c. 3600–2500 The Neolithic, New Stone Age, was approximately char- BCE. The earliest evidence for established trade exists in acterized by the adoption of agriculture, the shift from the Neolithic with newly settled people importing exotic food gathering to food producing in itself is one of the goods over distances of many hundreds of miles. most revolutionary changes in human history so-called These facts show that there were sufficient resources and Neolithic Revolution, the development of pottery, pol- co-operation to enable large groups to work on these ished stone tools and more complex, larger settlements projects. To what extent this was a basis for the devel- such as Çatal Hüyük and Jericho. Some of these features opment of elites and social hierarchies is a matter of on- began in certain localities even earlier, in the transitional going debate.[49] Although some late Neolithic societies Mesolithic. The first Neolithic cultures started around formed complex stratified chiefdoms similar to Polyne- 7000 BCE in the fertile crescent and spread concentri- sian societies such as the Ancient Hawaiians, based on the cally to other areas of the world; however, the Near East societies of modern tribesmen at an equivalent technolog- was probably not the only nucleus of agriculture, the cul- ical level, most Neolithic societies were relatively simple tivation of maize in Meso-America and of rice in the Far and egalitarian.[50] A comparison of art in the two ages East being others. leads some theorists to conclude that Neolithic cultures 5.4. MATERIAL CULTURE 43 were noticeably more hierarchical than the Paleolithic years ago.[53] It is considered as an equivalent of Euro- cultures that preceded them.[51] pean Middle Paleolithic.[54] It is associated with anatomically modern or almost modern Homo sapiens. Early physical evidence comes from Omo [55] and Herto,[56] 5.3.2 Three-stage chronology both in Ethiopia and dated respectively at c. 195 ka and at c. 160 ka. The Earlier or Early Stone Age (ESA)

Main articles: Paleolithic and Lower Paleolithic The Later Stone Age (LSA) This period is not to be identiﬁed with “Old Stone Age”, Main article: Later Stone Age

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

5.4 Material culture

5.4.1 Tools

Stone tools were made from a variety of stone. For example, ﬂint and chert were shaped (or chipped) for use as cutting tools and weapons, while basalt and sandstone were used for ground stone tools, such as quern-stones. Wood, bone, shell, antler (deer) and other materials were widely used, as well. During the most recent part of the period, sediments (such as clay) were used to make pottery. Agriculture was developed and certain animals Acheulean biface from Lake Langano area, Ethiopia. were domesticated. Some species of non-Primates are able to use stone tools, a translation of Paleolithic, or with Paleolithic, or with such as the Sea Otter, which breaks Abalone shells with the “Earlier Stone Age” that originally meant what be- them. Primates can both use and manufacture stone tools. came the Paleolithic and Mesolithic. In the initial decades This combination of abilities is more marked in apes and of its deﬁnition by the Pan-African Congress of Prehis- men, but only men, or more generally Hominans, depend tory, it was parallel in Africa to the Upper and Middle on tool use for survival.[57] The key anatomical and be- Paleolithic. However, since then Radiocarbon dating has havioral features required for tool manufacture, which are shown that the Middle Stone Age is in fact contempo- possessed only by Hominans, are the larger thumb and the raneous with the Middle Paleolithic.[52] The Early Stone ability to hold by means of an assortment of grips.[58] Age therefore is contemporaneous with the Lower Pale- olithic and happens to include the same main technologies, Oldowan and Acheulean, which produced Mode 1 5.4.2 Food and drink and Mode 2 stone tools respectively. A distinct regional term is warranted, however, by the location and chronol- Main articles: Paleolithic diet and Paleolithic diet and ogy of the sites and the exact typology. nutrition

Food sources of the Palaeolithic hunter-gatherers were The Middle Stone Age (MSA) wild plants and animals harvested from the environment. They liked animal organ meats, including the livers, Main article: Middle Stone Age kidneys and brains. Large seeded legumes were part of the human diet long before the agricultural revolu- The Middle Stone Age was a period of African prehis- tion, as is evident from archaeobotanical finds from the tory between Early Stone Age and Late Stone Age. It be- Mousterian layers of Kebara Cave, in Israel.[59] More- gan around 300,000 years ago and ended around 50,000 over, recent evidence indicates that humans processed 44 CHAPTER 5. STONE AGE and consumed wild cereal grains as far back as 23,000 Petroglyphs years ago in the Upper Paleolithic.[60] Near the end of the Wisconsin glaciation, 15,000 to 9,000 Main article: Petroglyph years ago, mass extinction of Megafauna such as the Wooly mammoth occurred in Asia, Europe, North Amer- Petroglyphs appeared in the Neolithic. A Petroglyph is an ica and Australia. This was the first Holocene extinc- intaglio abstract or symbolic image engraved on natural tion event. It possibly forced modification in the dietary stone by various methods, usually by prehistoric peoples. habits of the humans of that age and with the emergence They were a dominant form of pre-writing symbols. Pet- of agricultural practices, plant-based foods also became roglyphs have been discovered in different parts of the a regular part of the diet. A number of factors have been world, including Asia (Bhimbetka, India), North Amer- suggested for the extinction: certainly over-hunting, but ica (Death Valley National Park), South America (Cumbe also deforestation and climate change.[61] The net effect Mayo, Peru), and Europe (Finnmark, Norway). was to fragment the vast ranges required by the large animals and extinguish them piecemeal in each fragment. Rock paintings

5.4.3 Shelter and habitat

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

Rock painting at Bhimbetka, India, a World heritage site • A tent-like structure inside a cave near the Grotte du Lazaret, Nice, France. Main article: Cave painting

• A structure with a roof supported with timber, dis- In paleolithic times, mostly animals were painted, in the- covered in Dolni Vestonice, The Czech Republic, ory ones that were used as food or represented strength, dates to around 23,000 BCE. The walls were made such as the rhinoceros or large cats (as in the Chauvet of packed clay blocks and stones. Cave). Signs such as dots were sometimes drawn. Rare human representations include handprints and half- • Many huts made of mammoth bones were found in human/half-animal ﬁgures. The Cave of Chauvet in the Eastern Europe and Siberia. The people who made Ardèche département, France, contains the most impor- these huts were expert mammoth hunters. Exam- tant cave paintings of the paleolithic era, dating from ples have been found along the Dniepr river valley about 31,000 BCE. The Altamira cave paintings in Spain of Ukraine, including near Chernihiv, in Moravia, were done 14,000 to 12,000 BCE and show, among oth- Czech Republic and in southern Poland. ers, bisons. The hall of bulls in Lascaux, Dordogne, France, dates from about 15,000 to 10,000 BCE. • An animal hide tent dated to around 15000 to 10000 The meaning of many of these paintings remains un- BCE, in the Magdalenian, was discovered at Plateau known. They may have been used for seasonal ritu- Parain, France. als. The animals are accompanied by signs that suggest a possible magic use. Arrow-like symbols in Lascaux are sometimes interpreted as calendar or almanac use, but the 5.4.4 Art evidence remains interpretative.[62] Some scenes of the Mesolithic, however, can be typed Prehistoric art is visible in the artifacts. Prehistoric music and therefore, judging from their various modiﬁcations, is inferred from found instruments, while parietal art can are fairly clear. One of these is the battle scene between be found on rocks of any kind. The latter are petroglyphs organized bands of archers. For example, “the marching and rock paintings. The art may or may not have had a Warriors,” a rock painting at Cingle de la Mola, Castellón religious function. in 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 step • Megalithic tombs, multichambered, and dolmens, toward each other, each man carrying a bow in one hand single-chambered, were graves with a huge stone and a fistful of arrows in the other. A file of five men leads slab stacked over other similarly large stone slabs; one band, one of whom is a figure with a “high crowned they have been discovered all across Europe and hat.” In other scenes elsewhere, the men wear head- Asia and were built in the Neolithic and the Bronze dresses and knee ornaments but otherwise fight nude. Age. Some scenes depict the dead and wounded, bristling with arrows.[63] One is reminded of Ötzi the Iceman, a Copper Age mummy revealed by an Alpine melting glacier, who 5.5 Modern popular culture and collapsed from loss of blood due to an arrow wound in the back. the Stone Age

5.4.5 Stone Age rituals and beliefs

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

Imaginative depiction of the Stone Age, by Viktor Vasnetsov

The image of the caveman is commonly associated with the Stone Age. For example, the 2003 documentary series showing the evolution of humans through the Stone Age was called Walking with Cavemen, although only the last programme showed humans living in caves. While the idea that human beings and dinosaurs coexisted is sometimes portrayed in popular culture in cartoons, films Poulnabrone dolmen in County Clare, Ireland and computer games, such as The Flintstones, One Million Years B.C. and Chuck Rock, the notion of hominids and non-avian dinosaurs co-existing is not supported by any scientific evidence. Other depictions of the Stone Age include the best-selling Earth’s Children series of books by Jean M. Auel, which are set in the Paleolithic and are loosely based on archaeological and anthropological findings. The 1981 film Quest for Fire by Jean-Jacques Annaud tells the story of a group of neanderthals searching for their lost fire. A twenty first century series, Chronicles of Ancient Dark- ness by Michelle Paver tells of two New Stone Age children fighting to fulfil a prophecy and save their clan.

[64] Monte Bubbonia dolmen (single-chambered tomb), Sicily 5.6 See also from the Stone Age indicate certain rituals and beliefs of • the people in those prehistoric times. It is now believed Megalith that activities of the Stone Age humans went beyond the • Prehistoric warfare immediate requirements of procuring food, body cover- ings, and shelters. Speciﬁc rites relating to death and • Ice Age burial were practiced, though certainly diﬀering in style and execution between cultures. • Pleistocene 46 CHAPTER 5. STONE AGE

• Homo [21] Shea 2010, p. 49

• Timeline of the Stone Age [22] Shea 2010, p. 50 [23] Barham & Mitchell 2008, p. 132 5.7 Notes [24] Barham & Mitchell 2008, pp. 126–127. [25] Barham & Mitchell 2008, p. 128 [1] http://www.nhm.ac.uk/about-us/news/2010/august/ oldest-tool-use-and-meat-eating-revealed75831.html [26] Barham & Mitchell 2008, p. 145

[2] Barham & Mitchell 2008, p. 106 [27] Barham & Mitchell 2008, p. 146.

[3] Barham & Mitchell 2008, p. 147 [28] Barham & Mitchell 2008, p. 112

[4] Rogers & Semaw 2009, pp. 162–163 [29] Shea 2010, p. 57

[5] Rogers & Semaw 2009, p. 155 [30] Barham & Mitchell 2008, p. 73

[6] As to whether aethiopicus is the genus Australopithecus or [31] Brooks, Charles E.P. (1919), “The Correlation of the the genus Paranthropus, broken out to include the more Quaternary Deposits of the British Isles with Those of robust forms, anthropological opinion is divided and both the Continent of Europe”, Annual Report of the Board of usages occur in the professional sources. Regents of the Smithsonian Institution 1917, Washington: Government Pronting Oﬃce, p. 277 [7] Rogers & Semaw 2009, p. 164 [32] Hugo Obermaier; Christine Matthew; Henry Osborne [8] “Neolithic Vinca was a metallurgical culture”. Archaeo (1924). Fossil Man in Spain. New Haven: Yale University News. Reuters. 17 November 2007. Retrieved 25 Jan- Press for the Hispanic Society of America. p. 272. uary 2011. [33] Barham & Mitchell 2008, pp. 106–107 [9] “ASA Statement on the use of 'primitive' as a descriptor of contemporary human groups”. ASA News (Association of [34] Shea 2010, pp. 55–57 Social Anthropologists of the UK and Commonwealth). 27 August 2007. [35] Barham & Mitchell 2008, p. 24

[10] Clark 1970, p. 22 [36] Barham & Mitchell 2008, p. 130

[11] Clark 1970, pp. 18–19 [37] Jean-Paul Raynal; et al. (2010). “Hominid Cave at Thomas Quarry I (Casablanca, Morocco): Recent ﬁnd- [12] Deacon & Deacon 1999, pp. 5–6 ings and their context”. Quaternary International (223– 224): 369–382. [13] Isaac, Glynn (1982). “The Earliest Archaeological Traces”. In Clark, J. Desmond. The Cambridge History [38] Belmaker 2006, p. 9 of Africa. Volume. I: From the Earliest Times to C. 500 BC. Cambridge: Cambridge University Press. p. 246. [39] Belmaker 2006, pp. 119–120 [40] Belmaker 2006, p. 149 [14] Willoughby, Pamela R. (2007). The evolution of modern humans in Africa: a comprehensive guide. Lanham, MD: [41] Belmaker 2006, p. 147 AltaMira Press. p. 54. [42] Belmaker 2006, p. 67 [15] Barham & Mitchell 2008, p. 477 [43] Belmaker 2006, p. 21 [16] “History: Systematic Investigation of the African Later Tertiary and Quaternary”. The Wenner-Gren Foundation. [44] Belmaker 2006, p. 20 Retrieved 3 March 2011. [45] “Acheulian stone tools discovered near Chennai”. The [17] “Worsaae, Jens Jacob Asmussen”. Encyclopædia Britan- Hindu. nica. [46] “Bose, China”. What Does It Mean to be Human?. Smith- [18] Toth, Nicholas; Schick, Kathy (2007). “21 Overview sonian National Museum of Natural History. of Paleolithic Archaeology”. In Henke, H.C. Winfried; Hardt, Thorolf; Tattersall, Ian. Handbook of Paleoan- [47] Dalton, Rex (2 September 2009). “Europe’s thropology. Volume 3. Berlin; Heidelberg; New York: oldest axes discovered”. Nature News (Nature). Springer-Verlag. p. 1944. ISBN 978-3-540-32474-4 doi:10.1038/news.2009.878.

[19] http://www.britannica.com/EBchecked/topic/439507/ [48] Giovanni Muttoni; et al. (2009). “Pleistocene mag- Paleolithic-Period netochronology of early hominid sites at Ceprano and Fontana Ranuccio, Italy”. Earth and Planetary Science [20] Barham & Mitchell 2008, p. 130. Letters 286: 255–268. doi:10.1016/j.epsl.2009.06.032. 5.8. REFERENCES 47

[49] Kuijt, Ian (2000). “Chapter 13: Near Eastern Neolithic 5.8 References Research: Directions and Trends”. In Kuijt, Ian. Life in Neolithic Farming Communities: Social Organization, • Barham, Lawrence; Mitchell, Peter (2008). The Identity, and diﬀerentiation. Fundamental Issues in Ar- First Africans: African Archaeology from the Earli- chaeology. New York: Kluwer Academic/Plenum Pub- est Toolmakers to Most Recent Foragers. Cambridge lishers. p. 317 World Archaeology. Oxford: Oxford University [50] Boehm, Christopher (2000). “The Origin of Morality as Press. Social Control”. In Katz, Leonard D. Evolutionary Ori- • Belmaker, Miriam (March 2006). Community gins of Morality: Cross-disciplinary Perspectives. Journal of Consciousness Studies Volume 7. Thorverton: Imprint Structure through Time: 'Ubeidiya, a Lower Pleis- Academic. p. 158. ISBN 0-7190-5612-8 tocene Site as a Case Study (Thesis). Paleoanthro- pology Society. [51] Guthrie, Russell Dale (2005). The Nature of Paleolithic • Clark, J. Desmond (1970). The Prehistory of Africa. Art. Chicago: University of Chicago Press. pp. 419–420. ISBN 978-0-226-31126-5. Ancient People and Places, Volume 72. New York; Washington: Praeger Publishers. [52] Clark, J. Desmond (1982). “The Culture of the Middle • Deacon, Hilary John; Deacon, Janette (1999). Hu- Paleolithic/MIddle Stone Age”. In Clark, J. Desmond. man beginnings in South Africa: uncovering the se- The Cambridge History of Africa. Volume. I: From the Earliest Times to C. 500 BC. Cambridge: Cambridge crets of the Stone Age. Walnut Creek, Calif. [u.a.]: University Press. p. 248. Altamira Press. • Piccolo, Salvatore (2013). Ancient Stones: The Pre- [53] McBrearty and Brooks 2000 historic Dolmens of Sicily. Abingdon (UK): Brazen [54] Biological origins of modern humans Head Publishing. • Rogers, Michael J.; Semaw, Sileshi (2009). “From [55] McDougall et al. 2005 Nothing to Something: The Appearance and Con- text of the Earliest Archaeological Record”. In [56] White et al. 2003 Camps i Calbet, Marta; Chauhan, Parth R. Source- [57] Barham & Mitchell 2008, p. 74 book of paleolithic transitions: methods, theories, and interpretations. New York: Springer. [58] Barham & Mitchell 2008, p. 108 • Schick, Kathy D.; Nicholas Toth (1993). Mak- ing Silent Stones Speak: Human Evolution and the [59] Efraim Lev; Mordechai E. Kislev; Ofer Bar-Yosef (March Dawn of Technology. New York: Simon & Schus- 2005). “Mousterian vegetal food in Kebara Cave, Mt. Carmel”. Journal of Archaeological Science 32 (3): 475– ter. ISBN 0-671-69371-9. 484. doi:10.1016/j.jas.2004.11.006. • Shea, John J. (2010). “Stone Age Visiting Cards Re- visited: a Strategic Perspective on the Lithic Tech- [60] Dolores R. Piperno; Ehud Weiss; Irene Holst; Dani Nadel (5 August 2004). “Processing of wild cereal grains in nology of Early Hominin Dispersal”. In Fleagle, the Upper Palaeolithic revealed by starch grain analysis”. John G.; Shea, John J.; Grine, Frederick E.; Boden, Nature 430 (7000): 670–3. doi:10.1038/nature02734. Andrea L.; Leakey, Richard E,. Out of Africa I: the PMID 15295598. First Hominin Colonization of Eurasia. Dordrecht; Heidelberg; London; New York: Springer. pp. 47– [61] Turvey, Samuel T. (2009). “Chapter 2: In the shadow of 64. the megafauna: prehistoric mammal and bird extinctions across the Holocene”. In Turvey, Samuel T. Holocene Ex- tinctions. Oxford Biology. Oxford: Oxford University Press. pp. 16–17 5.9 Further reading

[62] Aczel, Amir D. (2000). The Cave and the Cathedral: How • Scarre, Christopher (ed.) (1988). Past Worlds: The a Real-Life Indiana Jones and a Research Scholar De- Times Atlas of Archaeology. London: Times Books. coded the Ancient Art of Man. Hoboken: John Wiley & ISBN 0-7230-0306-8. Sons Inc. pp. 157–158.

[63] Martínez, Antonio Beltrán (1982) [1979]. Rock art of the 5.10 External links Spanish Levant. The Imprint of Man. Cambridge: Cam- bridge University Press. pp. 48–51. • Giusepi, Robert A. (2000). “The Stone Age”. His- [64] Salvatore Piccolo, Ancient Stones: The Prehistoric Dol- tory World International. Retrieved 22 February mens of Sicily. Abingdon, 2013. 2011. 48 CHAPTER 5. STONE AGE

• Kowalski, D.R. “Stone Age Hand-axes”. Aerobi- ologicalEngineering.com. Retrieved 22 February 2011.

• 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 3 March 2011.

• “The ASA”. Association of Social Anthropologists of the UK and Commonwealth. Chapter 6

Woolly mammoth

The woolly mammoth (Mammuthus primigenius) was a change and consequent shrinkage of its habitat, hunting species of mammoth, the common name for the extinct by humans, or a combination of the two. Isolated popu- elephant genus Mammuthus. The woolly mammoth was lations survived on St. Paul Island until 6,400 years ago one of the last in a line of mammoth species, beginning and Wrangel Island until 4,000 years ago. After its ex- with Mammuthus subplanifrons in the early Pliocene. M. tinction, humans continued using its ivory as a raw mate- primigenius diverged from the steppe mammoth, M. tro- rial, a tradition that continues today. It has been proposed gontherii, about 200,000 years ago in eastern Asia. Its the species could be recreated through cloning, but this closest extant relative is the Asian elephant. method is as yet infeasible because of the degraded state The appearance and behaviour of this species are among of the remaining genetic material. the best studied of any prehistoric animal because of the discovery of frozen carcasses in Siberia and Alaska, as well as skeletons, teeth, stomach contents, dung, and de- 6.1 Taxonomy piction from life in prehistoric cave paintings. Mammoth remains had long been known in Asia before they became known to Europeans in the 17th century. The origin of these remains was long a matter of debate, and often explained as being remains of legendary creatures. The mammoth was identified as an extinct species of elephant by Georges Cuvier in 1796. The woolly mammoth was roughly the same size as modern African elephants. Males reached shoulder heights between 2.7 and 3.4 m (9 and 11 ft) and weighed up to 6 tonnes (6.6 short tons). Females averaged 2.6–2.9 me- A mammoth tusk with Inuit carvings of scenes on the Yukon tres (8.5–9.5 ft) in height and weighed up to 4 tonnes (4.4 River, 19th century. short tons). A newborn calf weighed about 90 kilograms (200 lb). The woolly mammoth was well adapted to the Remains of various extinct elephants were known by cold environment during the last ice age. It was covered Europeans for centuries, but were generally interpreted, in fur, with an outer covering of long guard hairs and a based on biblical accounts, as the remains of legendary shorter undercoat. The colour of the coat varied from creatures such as behemoths or giants. It was also the- dark to light. The ears and tail were short to minimise orised that they were remains of modern elephants that frostbite and heat loss. It had long, curved tusks and four had been brought to Europe during the Roman Repub- molars, which were replaced six times during the life- lic, for example the war elephants of Hannibal the Great time of an individual. Its behaviour was similar to that and Pyrrhus of Epirus, or animals that had wandered of modern elephants, and it used its tusks and trunk for north.[2] The first woolly mammoth remains studied by manipulating objects, fighting, and foraging. The diet of European scientists were examined by Hans Sloane in the woolly mammoth was mainly grass and sedges. In- 1728 and consisted of fossilised teeth and tusks from dividuals could probably reach the age of 60. Its habitat Siberia. Sloane was the first to recognise that the remains was the mammoth steppe, which stretched across north- belonged to elephants.[3] Sloane turned to another bibli- ern Eurasia and North America. cal explanation for the presence of elephants in the Arc- The woolly mammoth coexisted with early humans, who tic, asserting that they had been buried during the Great used its bones and tusks for making art, tools, and Flood, and that Siberia had previously been tropical prior dwellings, and the species was also hunted for food.[1] to a drastic climate change.[4] Others interpreted Sloane’s It disappeared from its mainland range at the end of the conclusion slightly differently, arguing the flood had car- Pleistocene 10,000 years ago, most likely through climate 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- the species, since holotype designation was not practised tered bones collected in Siberia and Britain. He discussed in Blumenbach’s time.[8] In 1828 Joshua Brookes recog- the question of whether or not the remains were from ele- nised the species was distinct enough to warrant a new phants, but drew no conclusions.[5] genus, and reclassified it as Mammuthus primigenius.[9] In 1738, Johann Philipp Breyne argued that mammoth It is unclear where and how the word “mammoth” originated. According to the Oxford English Dictionary, it fossils represented some kind of elephant. He could not [10] explain why a tropical animal would be found in such a comes from an old Vogul word mēmoŋt 'earth-horn'. cold area as Siberia, and suggested that they might have It may be a version of mehemot, the Arabic version of the biblical word “behemoth”. Another possible origin is been transported there by the Great Flood.[6] In 1796, French anatomist Georges Cuvier was the first to identify Estonian, where maa means earth, and mutt means mole. The word was first used in Europe during the early 17th the woolly mammoth remains not as modern elephants transported to the Arctic, but as an entirely new species. century, when referring to maimanto tusks discovered in Siberia.[11] Thomas Jefferson, who had a keen interest He argued this species had gone extinct and no longer existed, a concept that was not widely accepted at the in palaeontology, is partially responsible for transform- mammoth time.[2][7] ing the word from a noun describing the prehistoric elephant to an adjective describing anything of surprisingly large size. The first recorded use of the word 6.1.1 Etymology as an adjective was in a description of a wheel of cheese (the "Cheshire Mammoth Cheese") given to Jefferson in 1802.[12]

6.1.2 Evolution

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

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

In 2005, researchers assembled a complete mitochondrial genome profile of the woolly mammoth, which allowed them to trace the close evolutionary relationship between Cuvier’s 1796 comparison between the mandible of a woolly mammoths and Asian elephants, Elephas maximus.[15] mammoth and an Indian elephant. African elephants, Loxodonta africana, branched away from this clade around 6 million years ago, close to Following Cuvier’s identification, Johann Friedrich Blu- the time of the similar split between chimpanzees and menbach gave the woolly mammoth its scientific name, humans. Before the publication of the Neanderthal Elephas primigenius, in 1799, placing it in the same genus genome, many researchers expected the first fully seas the Asian elephant. This name means “the firstborn quenced nuclear genome of an extinct species would be elephant”. Henry Fairfield Osborn chose a molar from that of the mammoth.[16] A 2010 study confirmed these Blumenbach’s collection as the lectotype specimen for relationships, and suggested the mammoth and Asian ele- 6.2. DESCRIPTION 51

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

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

entered North America. A 2011 genetic study showed that two examined specimens of the Columbian mammoth were grouped within a subclade of woolly mammoths. This suggests that the two populations interbred and produced fertile offspring. A North American form M. jeffersonii, a possible hybrid between Columbian and woolly known as M. jeffersonii may be a hybrid between the two [23] mammoths. species. Individuals and populations showing transitional mor- Since many remains of each species of mammoth are phologies between each of the mammoth species are known from several localities, it is possible to recon- known, and primitive and derived species coexisted as struct the evolutionary history of the genus through well until the former disappeared. The different species morphological studies. Mammoth species can be identi- and their intermediate forms therefore can be termed fied from the number of enamel ridges on their molars; "chronospecies". Many intermediate subspecies have primitive species had few ridges, and the number in- also been proposed, but their validity is uncertain; decreased gradually as new species evolved and replaced pending on author, they are either considered primi- the preceding ones. The crowns of the teeth lengthened tive forms of an advanced species or advanced forms of and the skulls became taller to accommodate this. At the a primitive species.[21] Regional and intermediate sub- same time, the skulls became shorter from front to back species such as M. p. primigenius, M. p. jatzkovi, M. to minimise the weight.[21] These adaptations were ac- p. sibiricus, and M. p. fraasi have been proposed.[24] quired gradually as mammoths turned to more abrasive The St. Paul Island and Wrangel Island populations food items.[22] were described as dwarf varieties, much smaller than the The first known members of the genus Mammuthus are mainland woolly mammoth; the Wrangel Island popu- the African species M. subplanifrons from the Pliocene, lation was also proposed to be a new subspecies, M. p. and M. africanavus from the Pleistocene. The former is vrangeliensis.[25][26] The Wrangel mammoths were iso- thought to be the ancestor of later forms. Mammoths lated for 5000 years, but experienced only a slight loss entered Europe around 3 million years ago. The earli- of genetic variation.[27] est type known from there has been named M. rumanus, which spread across Europe and China. Only its molars are known, which show that it had 8–10 enamel ridges. A population evolved 12–14 ridges, splitting off from and 6.2 Description replacing the earlier type, becoming M. meridionalis. In turn, this species was replaced by the steppe mammoth, The appearance of the woolly mammoth is probably the M. trogontherii, with 18–20 ridges, which evolved in east- best known of any prehistoric animal due to the many ern Asia c. 1 million years ago. Mammoths derived from frozen specimens with preserved soft tissue and depic- M. trogontherii evolved molars with 26 ridges 200,000 tions by contemporary humans in their art. Fully grown years ago in Siberia and became the woolly mammoth, M. males reached shoulder heights between 2.7 and 3.4 m (9 primigenius.[21] The Columbian mammoth, M. columbi, and 11 ft) and weighed up to 6 tonnes (6.6 short tons). evolved from a population of M. trogontherii that had This is almost as large as extant male African elephants, 52 CHAPTER 6. WOOLLY MAMMOTH

Model at the Royal BC Museum Cave art from Les Combarelles, France dle Kolyma mammoth”, which was preserved with a complete trunk tip. Unlike the trunk lobes of modern ele- which commonly reach 3–3.4 m (9.8–11.2 ft), and is phants, the upper “finger” at the tip of the trunk had a long less than the size of the earlier mammoth species M. pointed lobe and was 10 cm (3.9 in) long, while the lower meridionalis and M. trogontherii, and the contemporary “thumb” was 5 cm (2.0 in) and was broader. The trunk M. columbi. The reason for the smaller size is unknown. of “Dima” was 76 cm (2.49 ft) long, whereas the trunk Female woolly mammoths averaged 2.6–2.9 m (8.5–9.5 of the adult “Liakhov mammoth” was 2 metres (6.6 ft) ft) in height and were built more lightly than males, weigh- long.[31] Few frozen specimens have preserved genitals, ing up to 4 tonnes (4.4 short tons). A newborn calf would so the gender is usually determined through examination have weighed about 90 kg (200 lb). These sizes are de- of the skeleton. Males were generally larger and had more duced from comparison with modern elephants of sim- [28] robust skeletons and tusks. The best indication of sex is ilar size. Though the mammoths on Wrangel Island the size of the pelvic girdle, as the birth canal is always were smaller than those of the mainland, their size var- wider in females than in males.[33] ied, and they were not small enough to be considered “dwarves”.[29] It has been claimed that the last woolly mammoth populations decreased in size and increased 6.2.1 Coat their sexual dimorphism, but this was dismissed in a 2012 study.[30] Woolly mammoths had several adaptations to the cold, most noticeably the layer of fur covering all parts of the body. Other adaptations to cold weather include ears that are far smaller than those of modern elephants; they were about 38 cm (15 in) long and 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 reduced heat loss and frostbite, and the tail was short for the same reason, only 36 cm (14 in) long in the “Berezovka mammoth”. The tail contained 21 vertebrae, whereas the tails of modern elephants contain 28–33. Their skin was no thicker than that of present-day elephants, between 1.25 and 2.5 cm (0.49 and 0.98 in). They had a layer of Fur in Naturhistorisches Museum, Vienna fat up to 10 cm (3.9 in) thick under the skin, which helped to keep them warm. Woolly mammoths had broad flaps The coat consisted of an outer layer of long, coarse “guard of skin under their tails which covered the anus; this is [31] hair”, which was 30 cm (12 in) on the upper part of the also seen in modern elephants. body, up to 90 cm (35 in) in length on the flanks and un- Other characteristic features depicted in cave paintings derside, and 0.5 mm (0.020 in) in diameter, and a denser include a large, high, single-domed head and a sloping inner layer of shorter, slightly curly under-wool, up to 8 back with a high shoulder hump, resulting from long cm (3.1 in) long and 0.05 mm (0.0020 in) in diameter. spinous processes on the neck vertebrae. These fea- The hairs on the upper leg were up to 38 cm (15 in) long, tures were not present in juveniles, which had concave and those of the feet were 15 cm (5.9 in) long, reaching backs like African elephants.[32] Another feature shown the toes. The hairs on the head were relatively short, but in cave paintings was confirmed by the discovery of a longer on the underside and the sides of the trunk. The frozen specimen in 1924, an adult nicknamed the “Mid- tail was extended by coarse hairs up to 60 cm (24 in) long, 6.2. DESCRIPTION 53 which were thicker than the guard hairs. It is likely that the woolly mammoth moulted seasonally, and that the heaviest fur was shed during spring. Since mammoth carcasses were more likely to be preserved during autumn, it is possible that only the winter coat has been preserved in frozen specimens. Modern elephants have much less hair, though juveniles have a more extensive covering of hair than adults.[34] Comparison between the over-hairs of woolly mammoths and extant elephants show that they did not differ much in overall morphology.[35] Woolly mammoths 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 the fur a glossy sheen.[36] Preserved woolly mammoth fur is orange-brown, but this is believed to be an artefact from the bleaching of pig- ment during burial. The amount of pigmentation varied from hair to hair and also within each hair.[31] A 2006 study sequenced the Mc1r gene (which influences hair colour in mammals) from woolly mammoth bones. Two alleles were found: a dominant (fully active) and a recessive (partially active) one. In mammals, recessive Mc1r alleles results in light hair. Mammoths born Specimen with asymmetrical tusks, the Smithsonian Museum with at least one copy of the dominant allele would have had dark coats, while those with two copies of the recessive allele would have had light coats.[37] A 2011 study tusks, but there is no fossil evidence that any adult woolly showed that light individuals would have been rare.[38] A mammoths lacked them.[40] 2014 study instead indicated that the colouration of an individual varied from non-pigmented on the overhairs, bi coloured, non-pigmented and mixed red-brown guard hairs, and non-pigmented underhairs, which would give a light overall appearance.[39]

6.2.2 Dentition

Woolly mammoths had very long tusks, which were more curved than those of modern elephants. The largest known male tusk is 4.2 m (14 ft) long and weighs 91 kg Molar from font de Champdamoy, France (201 lb), but 2.4–2.7 m (7.9–8.9 ft) and 45 kg (99 lb) was a more typical size. Female tusks averaged at 1.5–1.8 m Woolly mammoths had four functional molar teeth at a (4.9–5.9 ft) and weighed 9 kg (20 lb). About a quarter time, two in the upper jaw and two in the lower. 23 cm of the length was inside the sockets. The tusks grew spi- (9.1 in) of the crown was within the jaw, and 2.5 cm (1 in) rally in opposite directions from the base and continued was above. The crown was continually pushed forwards in a curve until the tips pointed towards each other. In and up as it wore down, comparable to a conveyor belt. this way, most of the weight would have been close to the The teeth had up to 26 separated ridges of enamel, which skull, and there would be less torque than with straight were themselves covered in “prisms” that were directed tusks. The tusks were usually asymmetrical and showed towards the chewing surface. These were quite wear re- considerable variation, with some tusks curving down in- sistant and kept together by cementum and dentine.A stead of outwards and some being shorter due to break- mammoth had six sets of molars throughout a lifetime, age. Calves developed small milk tusks a few centimetres which were replaced five times, though a few specimens long at six months old, which were replaced by perma- with a seventh set are known. The latter condition could nent tusks a year later. Tusk growth continued throughout extend the lifespan of the individual, unless the tooth only life but became slower as the animal reached adulthood. consisted of a few plates. The first molars were about The tusks grew by 2.5–15 cm (0.98–5.91 in) each year. the size of those of a human, 1.3 cm (0.51 in), the third Some cave paintings show woolly mammoths with small were 15 cm (6 in) 15 cm (5.9 in) long, and the sixth were or no tusks, but it is unknown whether this reflected real- about 30 cm (1 ft) long and weighed 1.8 kg (4 lb). The ity or was artistic license. Female Asian elephants have no molars grew larger and contained more ridges with each 54 CHAPTER 6. WOOLLY MAMMOTH replacement.[41] supported by fossil assemblages and cave paintings show- Distortion in the molars is the most common health prob- ing groups. It is therefore probable that most of their lem found in woolly mammoth fossils. Sometimes the other social behaviour was similar to those of modern replacement was disrupted, and the molars were pushed elephants. Accumulations of modern elephant remains into abnormal positions, but some animals are known have been termed "elephants’ graveyards", as these sites to have survived this. Teeth from Britain showed that were erroneously thought to be where old elephants went 2% of specimens had periodontal disease, with half of to die. Similar accumulations of woolly mammoth bones these containing caries. The teeth also sometimes had have been found; it is thought these are the result of individuals dying near or in the rivers over thousands of cancerous growths.[42] years, and their bones eventually being brought together by the streams, or due to animals being mired in mud. Some accumulations are also thought to be the remains 6.3 Palaeobiology of herds that died together at the same time, perhaps due to flooding.[45]

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

Adult woolly mammoths could effectively defend themselves from predators with their tusks, trunks and size, but Leg and foot of the “Yukagir mammoth” juveniles and weakened adults were vulnerable to pack hunters such as wolves, cave hyenas and large felines. The Trackways made by a woolly mammoth herd 11,300– tusks may also have been used in intra-species fighting, 11,000 years ago, have been found in the St. Mary Reser- such as territorial fights or fights over mates. Because voir in Canada, showing that there were in this case al- of their curvature, the tusks were not suitable for stab- most equal numbers of adults, sub-adults and juveniles. bing, but may have been used for hitting, as indicated The adults had a stride of 2 m (6.6 ft), and the juve- by injuries to some fossil shoulder blades. As in mod- niles ran to keep up.[46] The well-preserved foot of the ern elephants, the sensitive and muscular trunk worked adult male "Yukagir mammoth" shows that the soles of as a limb-like organ with many functions. It was used the feet contained many cracks that would have helped in for manipulating objects, and in social interactions. The gripping surfaces during locomotion. Like modern ele- very long hairs on the tail probably compensated for the phants, woolly mammoths walked on their toes and had shortness of the tail, enabling its use as a flyswatter, simi- large, fleshy pads behind the toes.[31] [43] lar to the tail on modern elephants. As in reindeer and Evidence of several different bone diseases has been musk oxen, the haemoglobin of the woolly mammoth was found in woolly mammoths. The most common of adapted to the cold, with three mutations to improve oxy- these diseases was osteoarthritis, found in 2% of speci- gen delivery around the body and prevent freezing. This mens. One specimen from Switzerland had several fused feature may have helped the mammoths to live in high [44] vertebrae as a result of this condition. The “Yukagir latitudes. mammoth” had suffered from spondylitis in two vertebrae, and osteomyelitis is also known from some specimens. Several specimens have healed bone fractures, showing that the animals had survived these injuries.[47] Parasitic flies and protozoa were identified in the gut of the calf “Dima”.[48]

6.3.1 Diet

Food at various stages of digestion has been found in the Mounted “family group” intestines of several woolly mammoths, giving a good picture of their diet. Woolly mammoths sustained them- Like modern elephants, woolly mammoths were likely selves on plant food, mainly grass and sedges, which were very social and lived in matriarchal family groups. This is supplemented with herbaceous plants, ﬂowering plants, 6.3. PALAEOBIOLOGY 55

Mandibles at various growth stages, Naturalis, Leiden

wards while opening, which made the sharp enamel ridges cut across each other and grind the food. The ridges were wear-resistant to enable the animal to chew large quan- tities of food, which often contained grit. Woolly mammoths may have used their tusks as shovels to clear snow from the ground and reach the vegetation buried below, and to break ice to drink. This is indicated on many pre- The frozen calf “Lyuba” which still had food in its stomach served tusks by flat, polished sections up to 30 centimetres (12 in) long on the part of the surface that would have reached the ground. The tusks were also used for ob- shrubs, mosses, and tree matter. The composition and taining food in other ways, such as digging up plants and exact varieties differed from location to location. Woolly stripping off bark.[54] mammoths needed a varied diet to support their growth, like modern elephants. An adult of six tonnes would need to eat 180 kg (397 lb) daily, and may have foraged as 6.3.2 Growth and reproduction long as twenty hours every day. The two-fingered tip of the trunk was probably adapted for picking up the short grasses of the last ice age (Quaternary glaciation, 2.58 million years ago to present) by wrapping around them, whereas modern elephants curl their trunks around the longer grass of their tropical environments. The trunk could also be used for pulling off large grass tufts, del- icately picking buds and flowers, and tearing off leaves and branches where trees and shrubs were present. The “Yukagir mammoth” had ingested plant matter that contained spores of dung fungus.[49] Isotope analysis shows that woolly mammoths fed mainly on C3 plants, unlike horses and rhinos.[50] Scientists identified milk in the stomach and faecal matter in the intestines of the mammoth calf "Lyuba".[51] The faecal matter may have been eaten by “Lyuba” to Cross sectioned tusk with growth rings promote development of the intestinal microbes necessary for digestion of vegetation, as is the case in modern [52] The age of a woolly mammoth can be determined by elephants. An isotope analysis of woolly mammoths counting the growth rings of its tusks when viewed in from Yukon, Canada, showed that the young nursed for at cross section. Each major line represents a year, and least three years, and were weaned and gradually changed weekly and daily ones can be found in between. Dark to a diet of plants when they were two to three years old. bands correspond to summers, and it is therefore possi- This is later than in modern elephants and may be due to ble to determine the season in which a mammoth died. a higher risk of predator attack or difficulty in obtaining The growth of the tusks slowed when it became harder to food during the long periods of winter darkness in high [53] forage, for example during disease, when a male was ban- latitudes. ished from the herd, and during periods of severe glacia- The molars were adapted to their diet of coarse tundra tion. Woolly mammoths continued growing past adult- grasses, with more enamel plates and a higher crown than hood, like other elephants. Unfused limb bones show that their earlier, southern relatives. The woolly mammoth males grew until they reached the age of 40, and females chewed its food by using its powerful jaw muscles to move grew until they were 25. The frozen calf “Dima” was 90 the mandible forwards and close the mouth, then back- cm (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- stretched across northern Asia, many parts of Europe, cess of erupting, and the first set would be worn out at and the northern part of North America during the last ice 18 months of age. The third set of molars lasted for ten age. It was similar to the grassy steppes of modern Rus- years, and this process was repeated until the final, sixth sia, but the flora was more diverse, abundant, and grew set emerged when the animal was 30 years old. A woolly faster. Grasses, sedges, shrubs, and herbaceous plants mammoth could probably reach the age of 60, like mod- were present, and scattered trees were mainly found in ern elephants of the same size. By then the last set of southern regions. This habitat was not dominated by ice molars would be worn out, the animal would be unable to and snow, as is popularly believed, since these regions chew and feed, and it would die of starvation.[55] are thought to have been high-pressure areas at the time. The habitat of the woolly mammoth also supported other grazing herbivores such as the woolly rhinoceros, wild horses and bison.[60] A 2014 study concluded that forbs (a group of herbaceous plants) were more important in the steppe-tundra than previously acknowledged, and that it was a primary food source for the ice-age megafauna.[61]

Head of the “Yukagir mammoth”

The best preserved head of a frozen adult specimen, that Restoration of a group in late Pleistocene northern Spain, by of a male nicknamed the “Yukagir mammoth”, shows Mauricio Antón that woolly mammoths had temporal glands between the ear and the eye.[56] This feature indicates that, like bull The southernmost woolly mammoth specimen known is from the Shandong province of China, and is 33,000 elephants, male woolly mammoths also entered "musth", [62] a period of heightened aggressiveness. The glands are years old. The southernmost European remains are from the Depression of Granada in Spain and are of used especially by males to produce an oily substance with [63] a strong smell called temporin. Their fur may have helped roughly the same age. DNA studies have helped de- in spreading the scent further.[57] termine the phylogeography of the woolly mammoth. A 2008 DNA study showed there were two distinct groups Examination of preserved calves show that they were of woolly mammoths: one that went extinct 45,000 years all born during spring and summer, and since modern ago and another one that went extinct 12,000 years ago. elephants have gestation periods of 21–22 months, it The two groups are speculated to be divergent enough is probable that the mating season was from summer to be characterised as subspecies. The group that went to autumn.[58] δ15N isotopic analysis of the teeth of extinct earlier stayed in the middle of the high Arctic, “Lyuba” has demonstrated their prenatal development, while the group with the later extinction had a much and indicates its gestation period was similar to that of wider range.[64] Recent stable isotope studies of Siberian a modern elephant, and that it was born in spring.[59] and New World mammoths have shown there were also differences in climatic conditions on either side of the Bering land bridge, with Siberia being more uniformly 6.4 Distribution and habitat cold and dry throughout the Late Pleistocene.[65] Dur- ing the Younger Dryas age, woolly mammoths briefly expanded into north-east Europe, whereafter the mainland populations became extinct.[66] A 2008 genetic study showed that some of the woolly mammoths that entered North America through the Bering land bridge from Asia migrated back about 300,000 years ago and had replaced the previous Asian population by about 40,000 years ago, Possible distribution during the last glacial period, based on lo- not long before the entire species went extinct.[67] Woolly cations of fossil finds mammoths have been found in the same locations as those of the Columbian mammoth in North America, but it is The habitat of the woolly mammoth is known as unknown whether the two species were sympatric. The "mammoth steppe" or “tundra steppe”. This environment woolly mammoth may have entered these southern ar- 6.5. RELATIONSHIP WITH HUMANS 57 eas during times when Columbian mammoth populations day, more than five hundred depictions of woolly mam- were absent.[68] moths are known, in media ranging from cave paintings and engravings on the walls of 46 caves in Russia, France and Spain to engravings and sculptures (termed "portable 6.5 Relationship with humans art") made from ivory, antler, stone and bone. Cave paintings 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 (6.5 ft) in length. Other notable caves with mammoth depictions are the Chauvet Cave, Les Combarelles Cave, and Font-de-Gaume.[70] A depiction in the Cave of El Castillo may instead show Palaeoloxodon, the “straight- tusked elephant”.[71] “Portable art” can be more accurately dated than cave art since it is found in the same deposits as tools and other ice age artefacts. The largest collection of portable mammoth art, consisting of 62 depictions on 47 plaques, was Woolly mammoth carved in ivory, discovered by Édouard Lartet found in the 1960s at an excavated open-air camp near in 1864 Gönnersdorf in Germany. There does not seem to be a correlation between the number of mammoths depicted Modern humans coexisted with woolly mammoths dur- and the species that were most often hunted, since rein- ing the Upper Palaeolithic period when they entered Eu- deer bones are the most frequently found animal remains rope from Africa between 30,000 and 40,000 years ago. at the site. Two spear throwers shaped as woolly mam- Prior to this, Neanderthals had coexisted with mammoths moths have also been found in France.[70] Some portable during the Middle Palaeolithic. Woolly mammoths were mammoth depictions may not have been produced where very important to ice age humans, and human survival they were discovered, but could have moved around by may have depended on the mammoth in some areas. Ev- ancient trading.[71] idence for such coexistence was not recognised until the 19th century. William Buckland published his discovery of the Red Lady of Paviland skeleton in 1823, which was found in a cave alongside woolly mammoth bones, 6.5.1 Exploitation but he mistakenly denied that these were contemporaries. In 1864, Édouard Lartet found an engraving of a woolly mammoth on a piece of mammoth ivory in the Abri de la Madeleine cave in Dordogne, France. This was the first widely accepted evidence for the coexistence of humans with prehistoric extinct animals and is the first contemporary depiction of such a creature known to modern science.[69]

Reconstructed bone hut, based on ﬁnds in Mezhyrich

Woolly mammoth bones were used as construction material for dwellings by both Neanderthals and modern humans during the ice age. More than 70 such dwellings are known, mainly from the Russian Plain. The bases of the huts were circular, and ranged from 8 to 24 square metres (86 to 258 sq ft). The arrangement of dwellings varied, and ranged from 1 m (3.3 ft) to 20 m (66 ft) apart, Cro-Magnon artists painting mammoths in Font-de-Gaume, by depending on location. Large bones were used as foun- Charles R. Knight dations for the huts, tusks for the entrances, and the roofs were probably skins held in place by bones or tusks. Some The woolly mammoth is the third most depicted animal in huts had ﬂoors that extended 40 cm (16 in) below ground. ice age art, after horses and bison, and these images were Some huts included ﬁreplaces, which used bones as fuel, produced between 35,000 and 11,500 years ago. To- probably because wood was scarce. It is possible that 58 CHAPTER 6. WOOLLY MAMMOTH some of the bones used for materials came from mam- hunted intensively, but perhaps mainly when ivory was moths killed by humans, but the state of the bones, and needed.[77] the fact that bones used to build a single dwelling varied by several thousands of years in age, suggests that they were collected remains of long-dead animals. Woolly 6.6 Extinction mammoth bones were also made into various tools, fur- niture, and musical instruments. Large bones, such as shoulder blades, were also used to cover dead human bod- ies during burial.[72]

16,500 year old mammoth spear thrower from France

Most woolly mammoth populations disappeared during the late Pleistocene and early Holocene, alongside most of the Pleistocene megafauna, during the Quaternary extinction event.[78] Scientists are divided over whether hunting or climate change, which led to the shrinkage of its The Venus of Brassempouy habitat, was the main factor that contributed to the extinction of the woolly mammoth, or whether it was due Woolly mammoth ivory was used to create art objects. to a combination of the two. Whatever the cause, large Several Venus figurines, including the Venus of Brassem- mammals are generally more vulnerable than smaller ones pouy and the Venus of Lespugue, were made from this due to their smaller population size and low reproduc- material. Weapons made from ivory, such as daggers, tion rates. Different woolly mammoth populations did spears, and a boomerang, are also known. To be able not die out simultaneously across their range, but grad- to process the ivory, the large tusks had to be chopped, ually went extinct over time. The last mainland popu- chiselled and split into smaller, more manageable pieces. lation existed in the Kyttyk Peninsula of Siberia 9,650 Some ivory artefacts show that tusks had been straight- years ago.[79] A small population of woolly mammoths ened, and it is unknown how this was achieved.[73] survived on St. Paul Island, Alaska, until 6,400 years [26][80][81] Several woolly mammoth specimens show evidence of ago. The last known population remained on Wrangel Island in the Arctic Ocean until 4,000 years being butchered by humans, which is indicated by breaks, [82][83][84] cut-marks, and associated stone tools. It is not known ago. Genetic evidence implies the extinction of this final population was sudden, rather than the culmi- how much prehistoric humans relied on woolly mammoth [84] meat, since there were many other large herbivores avail- nation of a gradual decline; the disappearance coin- cides roughly in time with the first evidence for humans on able. Many mammoth carcasses may have been scav- [85] enged by humans rather than hunted. Some cave paint- the island. The woolly mammoths of eastern Beringia ings show woolly mammoths in structures interpreted as (modern Alaska and Yukon) had similarly died out about pitfall traps. Few specimens show direct, unambiguous 13,300 years ago, soon (roughly 1000 years) after the first evidence of having been hunted by humans. A Siberian appearance of humans in the area, which parallels the fate specimen with a spearhead embedded in its shoulder of all the other late Pleistocene proboscids (mammoths, gomphotheres and mastodons), as well as most of the rest blade shows that a spear had been thrown at it with great [86] force.[74] A specimen from the Mousterian age of Italy of the megafauna, of the Americas. In contrast, the shows evidence of spear hunting by Neanderthals.[75] The St. Paul Island mammoth population apparently died out prior to human arrival due to habitat shrinkage resulting juvenile specimen nicknamed “Yuka” is the first frozen [86] mammoth with evidence of human interaction. It shows from the post-ice age sea level rise. evidence of having been killed by a large predator, and of A 2008 study estimated that changes in climate having been scavenged by humans shortly after. Some of shrank suitable mammoth habitat from 7,700,000 km2 its bones had been removed, and were found nearby.[76] (3,000,000 sq mi) 42,000 years ago to 800,000 km2 A site near the Yana River in Siberia has revealed sev- (310,000 sq mi) 6,000 years ago.[87][88] Woolly mam- eral specimens with evidence of human hunting, but the moths survived an even greater loss of habitat at the end finds were interpreted to show that the animals were not of the Saale glaciation 125,000 years ago, and it is likely 6.7. FROZEN SPECIMENS 59

fewer finds in the latter. Such remains are mostly found above the Arctic Circle, in permafrost. It appears that soft tissue was less likely to be preserved between 30,000 and 15,000 years ago, perhaps because the climate was milder during that period. Most specimens have partially degraded prior to discovery, due to exposure or to being scavenged. This "natural mummification" required the 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 seed pods still in the mouth of many of the specimens suggests neither starvation nor exposure are likely. The ma- Cast of the “Hebior Mammoth” specimen, which bears turity of this ingested vegetation places the time of death tool/butcher marks in autumn rather than in spring, when flowers would be expected.[93] The animals may have fallen through ice that humans hunted the remaining populations to extinc- into small ponds or potholes, entombing them. Many tion at the end of the last glacial period.[89][90] Studies are certainly known to have been killed in rivers, perhaps of an 11,300–11,000 year old trackway in southwestern through being swept away by floods. In one location, by Canada showed that M. primigenius was in decline while the Berelekh River in Yakutia in Siberia, more than 8,000 coexisting with humans, since far fewer tracks of juve- bones from at least 140 mammoths have been found in a single spot, apparently having been swept there by the niles were identified than would be expected in a normal [94] herd.[46] current. A 2010 study suggests that the decline of the woolly mammoth could have increased temperatures by up to 0.2 °C (0.36 °F) at high latitudes in the northern hemisphere. Mammoths frequently ate birch trees, creating a grassland habitat. With the disappearance of mammoths, birch forests, which absorb more sunlight than grasslands, expanded, leading to regional warming.[91]

6.7 Frozen specimens

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

Between 1692 and 1806, only four descriptions of frozen mammoths were published in Europe. None of the remains of those five were preserved, and no complete skeleton was recovered during that time.[95] While frozen woolly mammoth carcasses had been excavated by Euro- peans as early as 1728, the first fully documented specimen was discovered near the delta of the Lena River in 1799 by Ossip Schumachov, a Siberian hunter.[96] Early 19th century interpretation of the "Adams mammoth" car- Schumachov let it thaw until he could retrieve the tusks cass prior to excavation for sale to the ivory trade. While in Yakutsk in 1806, Michael Friedrich Adams heard about the frozen mam- Woolly mammoth fossils have been found in many differ- moth. Upon arrival at the location, Adams discovered ent types of deposits, including former rivers and lakes, that wild animals had eaten most of the organs and flesh and also in "Doggerland" in the North Sea, which was of the mammoth, including the trunk. He examined the dry at times during the ice age. Such fossils are usu- carcass and realised what was left would still be the most ally fragmentary and contain no soft tissue. Apart from complete mammoth recovered by that time. Adams re- frozen remains, the only soft tissue known is from a spec- covered the entire skeleton, apart from the tusks, which imen that was preserved in a petroleum seep in Starunia, Shumachov had already sold, and one foreleg, most of Poland. Frozen remains of woolly mammoths have been the skin, and nearly 18 kg (40 lb) of hair. During his re- found in the northern parts of Siberia and Alaska, with far turn voyage he purchased a pair of tusks that he believed 60 CHAPTER 6. WOOLLY MAMMOTH were the ones that Shumachov had sold. Adams brought it to the Zoological Museum of the Zoological Institute of the Russian Academy of Sciences, and the task of mount- ing the skeleton was given to Wilhelm Gottlieb Tilesius.[5] The Kunstkamera, the museum established by Peter the Great, contained the skeleton of an Indian elephant that could be used as reference.[97] This was one of the first attempts at reconstructing the skeleton of an extinct animal. Most of the reconstruction is correct, but Tilesius placed each tusk in the opposite socket, so that they curved outward instead of inward. The error was not corrected until 1899, and the correct placement of mammoth tusks was still a matter of debate into the twentieth century.[98][99]

“Dima”, a frozen calf about seven months old

the Kolyma River in northeastern Siberia. This specimen weighed approximately 100 kg (220 lb) at death and was 104 cm (41 in) high and 115 cm (45 in) long. Radiocarbon dating determined that “Dima” died about 40,000 years ago. Its internal organs are similar to those of modern elephants, but its ears are only one-tenth the size of those of an African elephant of similar age. A less complete juvenile, nicknamed “Mascha”, was found on the Yamal Peninsula in 1988. It was 3–4 months old, and a laceration on its right foot may have been the cause of A third of this model is covered with the skin of the “Berezovka death. It is the westernmost frozen mammoth found.[102] mammoth”, Museum of Zoology, St. Petersburg In 1997, a piece of mammoth tusk was discovered pro- truding from the tundra of the Taymyr Peninsula in The 1901 excavation of the “Berezovka mammoth” is the Siberia, Russia. In 1999, this 20,380 year old carcass and best documented of the early finds. It was discovered by 25 tons of surrounding sediment were transported by an the Berezovka River, and the Russian authorities financed Mi-26 heavy lift helicopter to an ice cave in Khatanga. its excavation. Its head was exposed, and the flesh had The specimen was nicknamed the “Jarkov mammoth”. been scavenged. The animal still had grass between its In October 2000, the careful defrosting operations in this teeth and on the tongue, showing that it had died sud- cave began with the use of hairdryers to keep the hair and denly. The entire expedition took 10 months, and the other soft tissues intact.[103][104] specimen had to be cut to pieces before it could be transported to St. Petersburg. It was identified as a 35–40 year old male, which had died 35,000 years ago. One of its shoulder blades was broken, which may have happened when it fell into a crevasse.[93] By 1929, the remains of 34 mammoths with frozen soft tissues (skin, flesh, or organs) had been documented. Only four of them were relatively complete. Since then, about that many more have been found. In most cases, the flesh showed signs of decay before its freezing and later desiccation.[100] Since 1860, Russian authorities have of- Frozen calf nicknamed “Mascha” fered rewards of up to руб.1000 for finds of frozen woolly mammoth carcasses. Often such finds were kept secret In 2007, the carcass of a female calf nicknamed “Lyuba” due to superstition. Several carcasses have been lost be- was discovered near the Yuribey River, where it had cause they were not reported, and one was fed to dogs. been buried for 41,800 years.[52][105] By cutting a section In more recent years, scientific expeditions have been through a molar and analysing its growth lines, they found devoted to finding carcasses instead of relying solely on [59] [101] that the animal had died at the age of one month. The chance encounters. mummified calf weighed 50 kg (110 lb), was 85 cm (33 In 1977, the well-preserved carcass of a seven- to eight- in) high and 130 cm (51 in) in length.[106][107] At the time month-old woolly mammoth calf named “Dima” was dis- of discovery, its eyes and trunk were intact and some fur covered. This carcass was recovered near a tributary of remained on its body. Its organs and skin are very well 6.7. FROZEN SPECIMENS 61

preserved.[108] “Lyuba” is believed to have been 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, preserving the mammoth in a nearly pristine state.[52]

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

Skull and mandible of the frozen calf “Yuka”

In 2012, a juvenile was found in Siberia, which had man- made cut marks. Scientists estimated its age at death to elephant–mammoth hybrid, and the process would have be 2.5 years, and nicknamed it “Yuka”. Its skull and to be repeated so more hybrids could be used in breeding. pelvis had been removed prior to discovery, but were After several generations of cross-breeding these hybrids, found nearby.[110][76] Another mammoth discovery was an almost pure woolly mammoth would be produced. The reported in October 2012, when it was excavated on the fact that sperm cells of modern mammals are potent for Taymyr Peninsula. It was dated to 30,000 years old. It 15 years at most after deep-freezing is a hindrance to was named “Zhenya” after the boy who found it.[111] this method.[113] In one case, an Asian elephant and an African elephant produced a live calf named Motty, but In 2013, a well preserved carcass was found on Maly it died of defects at less than two weeks old.[114] Lyakhovsky Island, one of the islands in the New Siberian Islands archipelago, a female between 50 and 60 years old In 2008, a Japanese team found usable DNA in the at the time of death. The carcass contained well preserved brains of mice that had been frozen for 16 years. They muscular tissue. When it was extracted from the ice, liq- hope to use similar methods to find usable mammoth uid blood spilled from the abdominal cavity. The find- DNA.[115] In 2009, the Pyrenean Ibex (a subspecies of ers interpreted this as indicating woolly mammoth blood the Spanish ibex) was the first extinct animal to be cloned possessed anti-freezing properties.[112] back to life; the clone lived for only seven minutes before dying of lung defects.[116] As the woolly mammoth genome has been mapped, it may be possible to recre- 6.7.1 Recreating the species ate a complete set of woolly mammoth chromosomes in the future by adding mammoth-only sequences to ele- The existence of preserved soft tissue remains and DNA phant chromosomes.[117] If the process is ever successful, of woolly mammoths has led to the idea that the species there are plans to introduce cloned woolly mammoths to could be recreated by scientific means. Two methods Pleistocene Park, a wildlife reserve in Siberia.[118] have been proposed to achieve this. The first is cloning, Mammoth expert Adrian Lister questions the ethics of which would involve removal of the DNA-containing such recreation attempts. In addition to the technical nucleus of the egg cell of a female elephant, and replace- problems, he notes that there is not much habitat left that ment with a nucleus from woolly mammoth tissue. The would be suitable for woolly mammoths. Because the cell would then be stimulated into dividing, and inserted species was social and gregarious, creating a few spec- back into a female elephant. The resulting calf would imens would not be ideal. He also notes that the time have the genes of the woolly mammoth, although its fetal and resources required would be enormous, and that the environment would be different. To date, even the most scientific benefits would be unclear; he suggests these re- intact mammoths have had little usable DNA because of sources should instead be used to preserve extant elephant their conditions of preservation. There is not enough to [113] [113] species which are endangered. A 2014 article about guide the production of an embryo. potential cloning also questioned the ethics of using ele- The second method involves artificially inseminating an phants as surrogate mothers, as most embryos would not elephant egg cell with sperm cells from a frozen woolly survive, and noted that it would be impossible to know mammoth carcass. The resulting offspring would be an the exact needs of a resurrected calf.[119] 62 CHAPTER 6. WOOLLY MAMMOTH

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

Peter III of Russia carved in mammoth ivory

The woolly mammoth has remained culturally significant long after its extinction. Indigenous peoples of Siberia had long found what are now known to be woolly mammoth remains, collecting their tusks for the ivory trade. Native Siberians believed woolly mammoth remains to be those of giant mole-like animals that lived underground Le Mammouth by Paul Jamin, 1885 and died when burrowing to the surface.[6] Woolly mammoth tusks had been articles of trade in Asia long be- Stories abound about frozen woolly mammoth meat that fore Europeans became acquainted with them. Güyük, was consumed once defrosted, especially that of the the 13th-century Khan of the Mongols, is reputed to have “Berezovka mammoth”, but most of these are considered sat on a throne made from mammoth ivory.[95] Siberian dubious. The carcasses were in most cases decayed, and mammoth ivory is reported to have been exported to Rus- the stench so unbearable that only wild scavengers and sia and Europe in the 10th century. The first Siberian the dogs accompanying the finders showed any interest ivory to reach western Europe was brought to London in in the flesh. It appears that such meat was once recom- 1611.[120] mended against illness in China, and Siberian natives have occasionally cooked the meat of frozen carcasses they When Russia occupied Siberia, the ivory trade grew discovered.[123] and became a widely exported commodity, with huge amounts being excavated for this market. From the 19th century and onwards, woolly mammoth ivory became a 6.8.1 Cryptozoology highly prized commodity, used as raw material for many different products. Today it is still in great demand as a There have been occasional claims that the woolly mam- replacement for the now-banned export of elephant ivory, moth is not extinct, and that small isolated herds might and has been referred to as “white gold”. Local dealers survive in the vast and sparsely inhabited tundra of the estimate that there are 10 million mammoths still frozen Northern Hemisphere. In the 19th century, several rein Siberia, and conservationists have suggested that this ports of “large shaggy beasts” were passed on to the Rus- could help save the living species of elephants from ex- sian authorities by Siberian tribesmen, but no scientific tinction. Elephants are hunted by poachers for their ivory, proof ever surfaced. A French chargé d'affaires work- but if this could instead be supplied by the already ex- ing in Vladivostok, M. Gallon, said in 1946 that in 1920 tinct mammoths, the demand could instead be met by he had met a Russian fur-trapper who claimed to have these. Trade in elephant ivory has been forbidden in seen living giant, furry “elephants” deep into the taiga. most places following the 1989 Lausanne Conference, but Gallon added that the fur-trapper had not heard of mam- dealers have been known to label it as mammoth ivory 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- [8] Reich, M.; Gehler, A. (2008). “Giants’ Bones and Uni- vived into more recent times, but all evidence indicates corn Horns Ice Age Elephants Offer 21st Century In- that they went extinct thousands of years ago. It is likely sights”. Collections - Wisdom, Insight, Innovation 8: 44– that these natives had gained their knowledge of woolly 50. mammoths from carcasses they encountered, and that this [125] [9] Brookes, J. (1828). A catalogue of the anatomical & zo- is the source for their legends of the animal. ological museum of Joshua Brookes 1. London: Richard Legends from several Native American tribes have also Taylor. p. 73. been interpreted by some as indicating folk memory of [10] “Mammoth entry in Oxford English Dictionary”. 2000. extinct elephants.[126][127][128] In the late 19th century, there were persistent rumours about surviving mammoths [11] Lister, 2007. p. 49 in Alaska.[124] In October 1899, Henry Tukeman detailed his killing of a mammoth in Alaska and his subsequent [12] Simpson, J. (2009). "Word Stories: Mammoth.” Oxford donation of the specimen to the Smithsonian Institution English Dictionary Online, Oxford University Press. Ac- in Washington, D.C. The museum denied the existence cessed 5 June 2009. [129] of any mammoth corpse; the story was a hoax. Bengt [13] Lister, 2007. pp. 18–21 Sjögren believed that the myth began when the American biologist Charles Haskins Townsend travelled in Alaska, [14] Shoshani, J.; Tassy, P. (2005). “Advances in pro- saw Eskimos trading mammoth tusks, asked if there still boscidean taxonomy & classification, anatomy & phys- were living mammoths in Alaska, and provided them with iology, and ecology & behavior”. Quaternary Inter- a drawing of the animal.[124] national. 126–128: 5. Bibcode:2005QuInt.126....5S. doi:10.1016/j.quaint.2004.04.011.

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Woolly rhinoceros

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

7.1 Evolution

Cast of the 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 2 m (6.6 ft) tall;[1] the body size was thus comparable, or slightly larger than, the extant White rhinoceros.[4] Two horns on the skull were made of keratin, the anterior horn being 61 cm (24 in) in length,[5] with a smaller horn between its eyes.[6] It had thick, long fur, small ears, short, Chauvet cave art depicting a woolly rhino thick legs, and a stocky body. Cave paintings suggest a wide dark band between the front and hind legs, but As the last and most derived member of the Pleistocene the feature is not universal, and identification of pictured rhinoceros lineage, the woolly rhinoceros was well rhinoceroses as woolly rhinoceros is uncertain. adapted to its environment. Stocky limbs and thick woolly pelage made it well suited to the steppe-tundra Its shape was known only from prehistoric cave drawings environment prevalent across the Palearctic ecozone dur- until a completely preserved specimen (missing only the ing the Pleistocene glaciations. Like the vast majority fur and hooves) was discovered in a tar pit in Starunia, of rhinoceroses, the body plan of the woolly rhinoceros Poland. The specimen, an adult female, is now on display adhered to a conservative morphology, like the first in the Polish Academy of Sciences' Museum of Natural rhinoceroses seen in the late Eocene. History in Kraków. Several frozen specimens have also been found in Siberia, the latest in 2007.[7] A study of 40-70.000 year old DNA samples showed its closest extant relative is the Sumatran rhinoceros.[2] 7.3 Behavior and habitat 7.2 Description The woolly rhinoceros used its horns for defensive pur- The external appearance of woolly rhinos is known from poses and to attract mates. During Greenland Stadial 2 mummified individuals from Siberia as well as cave (the Last Glacial Maximum[8]) the North Sea retreated paintings.[3] An adult woolly rhinoceros was typically northward, as sea levels were up to 125 metres (410 ft) around 3 to 3.8 metres (10 to 12.5 feet) in length, with lower than today. The woolly rhinoceros roamed the ex-

68 7.4. EXTINCTION 69

constructed using several lines of evidence. Climatic re- constructions indicate the preferred environment to have been cold and arid steppe-tundra, with large herbivores forming an important part of the feedback cycle. Pollen analysis shows a prevalence of grasses and sedges within a more complicated vegetation mosaic. A strain vector biomechanical investigation of the skull, mandible and teeth of a well-preserved last cold stage individual recovered from Whitemoor Haye, Staﬀordshire, revealed musculature and dental characteristics that support a grazing feeding preference. In particular, the en- largement of the temporalis and neck muscles is consistent with that required to resist the large tugging forces generated when taking large mouthfuls of fodder from Restoration by Charles R. Knight the ground. The presence of a large diastema supports this theory. posed Doggerland and much of Northern Europe and Comparisons with extant perissodactyls conﬁrm that was common in the cold, arid desert that is southern Coelodonta was a hindgut fermentor with a single stom- [9] England and the North Sea today. Its geographi- ach, and as such would have grazed upon cellulose-rich, cal range expanded and contracted with the alternating protein-poor fodder. This method of digestion would cold and warm cycles, forcing populations to migrate as have required a large throughput of food and thus links glaciers receded. The woolly rhinoceros co-existed with the large mouthful size to the low nutritive content of the woolly mammoths and several other extinct larger mam- chosen grasses and sedges.[12] mals of the Pleistocene megafauna. A close relative, Elasmotherium, had a more southern range. Recent evidence suggests that woolly rhinos alive in the Arctic during the Last Glacial Maximum consumed In 2011, the oldest known woolly rhinoceros fossil was approximately equal volumes of forbs, such as Artemisia, discovered from 3.6 million years in the Himalayas on the and graminoids.[13] cold Tibetan Plateau, suggesting it existed there during a period of general climate warmth around the earth. It is believed that they migrated from there to northern Asia and Europe when the Ice Age began.[10] 7.4 Extinction Females gave birth to one or two calves.[11] Main article: Pleistocene megafauna 7.3.1 Diet Many species of Pleistocene megafauna, like the woolly

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

rhinoceros, became extinct around the same time period. Human and Neanderthal hunting is often cited as one cause.[14] Other theories for the cause of the extinctions Frozen head, leg and horn from Siberia, 1849 are climate change associated with the receding Ice age and the hyperdisease hypothesis (q.v. Quaternary extinc- [15] Controversy has long surrounded the precise dietary pref- tion event). erence of Coelodonta as past investigations have found Recent radiocarbon dating indicates that populations sur- both grazing and browsing modes of life to be plausi- vived as recently as 8,000 BC in western Siberia. How- ble. The palaeodiet of the woolly rhinoceros has been re- ever, the accuracy of this date is uncertain, as several ra- 70 CHAPTER 7. WOOLLY RHINOCEROS diocarbon plateaus exist around this time. The extinction [9] Ian Rolfe, W. D. (1966). “Woolly rhinoceros from the does not coincide with the end of the last ice age but does Scottish Pleistocene”. Scottish Journal of Geology 2 (3): coincide with a minor yet severe climatic reversal that 253. doi:10.1144/sjg02030253. lasted for about 1,000–1,250 years, the Younger Dryas [10] “Ice Age giants may have evolved in Tibet”. CNN. 1 (GS1 - Greenland Stadial 1), characterized by glacial September 2011. Retrieved 2 September 2011. readvances and severe cooling globally, a brief interlude in the continuing warming subsequent to the termination [11] Walker, Matt (6 December 2012). “Prehistoric rhino re- of the last major ice age (GS2), thought to have been due veals secrets”. BBC News. to a shutdown of the thermohaline circulation in the ocean [12] SAS Bulletin, Volume 26, number 3/4, Winter 2003 from due to huge influxes of cold fresh water from the preced- the Society for Archaeological Sciences ing sustained glacial melting during the warmer Intersta- dial (GI1 - Greenland Interstadial 1 - ca. 16,000–11,450 [13] Willerslev E, Davison J, Moora M, Zobel M, Coissac E, 14C years B.P.). Edwards ME, Lorenzen ED, Vestergård M, Gussarova G, Haile J, Craine J, Gielly L, Boessenkool S, Epp LS, The Pinhole Cave Man is a late Paleolithic figure of a man Pearman PB, Cheddadi R, Murray D, Bråthen KA, Yoc- engraved on a rib bone of the Woolly rhinoceros, found coz N, Binney H, Cruaud C, Wincker P, Goslar T, Al- at Creswell Crags in England.[16] 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- 7.5 See also panon F, Rioux D, Niderkorn V, Tikhonov A, Savvinov G, Roberts RG, MacPhee RD, Gilbert MT, Kjær KH, Orlando L, Brochmann C, Taberlet P. (2014). “Fifty 7.6 References thousand years of Arctic vegetation and megafaunal diet”. Nature 506 (7486): 47–51. doi:10.1038/nature12921. PMID 24499916. [1] “Woolly Rhino (Coelodonta antiquitatis)". International Rhino Foundation. Retrieved October 30, 2011. [14] Diamond, Jared (1997). Guns, Germs and Steel. New York: Vintage. ISBN 0-09-930278-0. [2] Orlando, L.; Leonard, J. A.; Thenot, A. L.; Laudet, V.; Guerin, C.; Hänni, C. (2003). “Ancient DNA [15] Grayson, D. K.; Meltzer, D. J. (2003). “A requiem analysis reveals woolly rhino evolutionary relation- for North American overkill”. Journal of Archaeo- ships”. Molecular Phylogenetics and Evolution 28 (3): logical Science 30 (5): 585–593. doi:10.1016/S0305- 485–499. doi:10.1016/S1055-7903(03)00023-X. PMID 4403(02)00205-4. 12927133. [16] “engraved bone/antler”. British Museum. [3] “Frozen Fauna of the Mammoth Steppe”. • [4] Krause, Hans (2011). “HKHPE 07 02”. hanskrause.de. Parker, Steve. Dinosaurus: The Complete Guide to Retrieved July 12, 2012. Dinosaurs. Firefly Books Inc, 2003. Pg. 422.

[5] Haines, Tim; Chambers, Paul (2005). “Coelodonta”. The complete guide to prehistoric life (First ed.). Buﬀalo, N.Y.: Fireﬂy Books. p. 203. ISBN 978-1-55407-181-4. 7.7 External links

[6] Fortelius, Mikael (1983). “The morphology and pa- • More pictures of the fully preserved tar pit wholly leobiological signiﬁcance of the horns ofCoelodonta rhinoceros that was found in Poland (text in Polish) antiquitatis(Mammalia: Rhinocerotidae)". Jour- nal of Vertebrate Paleontology 3 (2): 125–135. • Fossil skull of a woolly rhinoceros from Belgium doi:10.1080/02724634.1983.10011964. ISSN 0272- 4634. • Fossil skull of a woolly rhinoceros from Germany

[7] Boeskorov, G. G. (2012). “Some speciﬁc mor- • International Rhino Foundation: Woolly Rhino phological and ecological features of the fossil woolly 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 radiocarbon ages on woolly rhinoceros (Coelodonta antiquitatis) from western central Scotland: signiﬁcance for timing the extinction of woolly rhinoceros in Britain and 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. 7.8. TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES 71

7.8 Text and image sources, contributors, and licenses

7.8.1 Text

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bot 4, Codyag123, Torrr torrr, Pinethicket, I 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. 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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, The way, the truth, and the light, Scarlet23, Mimihitam, Goustien, RingManX, WikiLaurent, Pavel.Riha.CB, ImageRemovalBot, Sfan00 IMG, ClueBot, The Thing That Should Not Be, Niceguyedc, Estirabot, Amaltheus, Darkicebot, Fastily, T.carnifex, Little Mountain 5, Sleptrip, Addbot, Guoguo12, Ronhjones, Ehrenkater, Lightbot, Laikayiu, Legobot, Luckas-bot, 2D, Azcolvin429, Againme, Kingpin13, 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, Peter M. 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. 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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- ,Solrac1993, Legobot ,ماني ,nutter, Ka Faraq Gatri, LaaknorBot, 37ophiuchi, AndersBot, Spike the Dingo, Tyw7, Tide rolls, Lightbot 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, ClueBot NG, Joefromrandb, Vacation9, Stuartsmally, Widr, Helpful Pixie Bot, Gob Lofa, Yendor of yinn, MangoWong, MusikAnimal, Davidiad, Cadiomals, Ornithodiez, Fjasl;d, Pseudofusulina, BattyBot, Isumbard Prince, Markomazzoni, US Jingoist, Ntra00, Hoppeduppeanut, 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. 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Xspike15, Lizia7, Kimboslicee, JaconaFrere, Dinorexcoolio, Monkbot, Supermariolink777, Braden12345, Iluhrs, Owen minns, Flingbong, HMSLavender, 115ash, PrinceSulaiman, Oinksgiant2000, Ninafundisha, Speedytacos, Surajbryan, Jessica Simpson03, , Guillame Heavensburg and Anonymous: 1000

• Woolly mammoth Source: http://en.wikipedia.org/wiki/Woolly%20mammoth?oldid=636651581 Contributors: William Avery, 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

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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

• File:Arrowhead.jpg Source: http://upload.wikimedia.org/wikipedia/commons/5/54/Arrowhead.jpg License: Public domain Contributors: http://www.ornl.gov/info/news/pulse/pulse_v44_99.htm Original artist: • File:Awashrivermap.png Source: http://upload.wikimedia.org/wikipedia/commons/9/94/Awashrivermap.png License: CC-BY-SA-3.0 Contributors: 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 • File:Bhimbetka_rock_paintng1.jpg Source: http://upload.wikimedia.org/wikipedia/commons/b/b6/Bhimbetka_rock_paintng1.jpg Li- cense: CC-BY-SA-3.0 Contributors: ? Original artist: ? • File:Blakey_Pleistmoll.jpg Source: http://upload.wikimedia.org/wikipedia/commons/7/7e/Blakey_Pleistmoll.jpg License: CC-BY-SA- 3.0 Contributors: http://cpgeosystems.com/mollglobe.html Original artist: Ron Blakey, NAU Geology • File:Boltunov_mammoth.jpg Source: http://upload.wikimedia.org/wikipedia/commons/d/dd/Boltunov_mammoth.jpg License: Public domain Contributors: http://johnmckay.blogspot.dk/2009/11/first-great-mammoth.html Original artist: Roman Boltunov • File:Canto_tallado_2-Guelmim-Es_Semara.jpg Source: http://upload.wikimedia.org/wikipedia/commons/8/87/Canto_tallado_ 2-Guelmim-Es_Semara.jpg License: Public domain Contributors: Own work Original artist: José-Manuel Benito Álvarez (España) —> Locutus Borg • File:Carved_19th_century_Eskimo_mammoth_tusk_De_Young_Museum_34430_2.JPG Source: http://upload.wikimedia.org/ wikipedia/commons/8/8d/Carved_19th_century_Eskimo_mammoth_tusk_De_Young_Museum_34430_2.JPG License: CC-BY-SA-3.0 Contributors: Own work Original artist: BrokenSphere • File:Cenozoic_cosmo_1894_beard_1913.png Source: http://upload.wikimedia.org/wikipedia/commons/9/98/Cenozoic_cosmo_1894_ beard_1913.png License: Public domain Contributors: ? Original artist: ? • File:Co2_glacial_cycles_800k.png Source: http://upload.wikimedia.org/wikipedia/commons/c/c9/Co2_glacial_cycles_800k.png Li- cense: CC-BY-SA-3.0 Contributors: Data from [1], and this looks best:3. Composite CO2 record (0-800 kyr BP), marked up with 230ppm transition between glacial and interglacial periods. Original artist: Tomruen • File:Coelodonta_antiquitatis_.jpg Source: http://upload.wikimedia.org/wikipedia/commons/8/87/Coelodonta_antiquitatis_.jpg Li- cense: CC-BY-SA-4.0 Contributors: Own work Original artist: Didier Descouens • File:Combarelles-mammouth.jpg Source: http://upload.wikimedia.org/wikipedia/commons/3/3e/Combarelles-mammouth.jpg Li- cense: Public domain Contributors: personal scan from Manuel d'archéologie préhistorique, celtique et gallo-romaine, fr:Joseph Déchelette (1862-1914) Original artist: A Cro-Magnon caveman • File:Commons-logo.svg Source: http://upload.wikimedia.org/wikipedia/en/4/4a/Commons-logo.svg License: ? Contributors: ? Original artist: ? • File:Cscr-featured.svg Source: http://upload.wikimedia.org/wikipedia/en/e/e7/Cscr-featured.svg License: ? Contributors: ? Original artist: ? • File:Cuvier_elephant_jaw.jpg Source: http://upload.wikimedia.org/wikipedia/commons/7/74/Cuvier_elephant_jaw.jpg License: Public domain Contributors: This plate was originally published in Georges Cuvier’s paper in 1798–99,[1] and can be seen in: Rudwick, Martin; Cuvier, Georges (1998) Georges Cuvier, Fossil Bones, and Geological Catastrophes: New Translations and Interpretations of the Primary Texts, University of Chicago Press, pp. p. 23 Retrieved on 16 June 2009. ISBN: 0226731073. Original artist: Georges Cuvier (1769–1832) • 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 spread during 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: Original uploader 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 was Botaurus 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 on flickr • File:Five_Myr_Climate_Change.svg Source: http://upload.wikimedia.org/wikipedia/commons/f/f7/Five_Myr_Climate_Change.svg License: CC-BY-SA-3.0 Contributors: see below Original artist: Dragons flight, svg by Jo • 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: ? • File:Font-de-Gaume.jpg Source: http://upload.wikimedia.org/wikipedia/commons/2/27/Font-de-Gaume.jpg License: Public domain Contributors: http://birdbookerreport.blogspot.com/2012/01/new-title_30.html Original artist: Charles R. Knight • 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

• File:Hebior_Mammoth_Clean.png Source: http://upload.wikimedia.org/wikipedia/commons/7/7a/Hebior_Mammoth_Clean.png Li- cense: CC-BY-SA-3.0 Contributors: Own work Original artist: MCDinosaurhunter • File:IceAgeEarth.jpg Source: http://upload.wikimedia.org/wikipedia/commons/b/b4/IceAgeEarth.jpg License: CC-BY-SA-3.0 Contrib- utors: Own work Original artist: Ittiz • File:Ice_Age_Temperature.png Source: http://upload.wikimedia.org/wikipedia/commons/f/f8/Ice_Age_Temperature.png License: CC- BY-SA-3.0 Contributors: ? Original artist: ? • File:Ice_age_fauna_of_northern_Spain_-_Mauricio_Antón.jpg Source: http://upload.wikimedia.org/wikipedia/commons/e/e6/Ice_ age_fauna_of_northern_Spain_-_Mauricio_Ant%C3%B3n.jpg License: CC-BY-2.5 Contributors: http://www.plosbiology.org/article/ slideshow.action?uri=info:doi/10.1371/journal.pbio.0060099&imageURI=info:doi/10.1371/journal.pbio.0060099.g001, from C. Sed- wick (1 April 2008). “What Killed the Woolly Mammoth?". PLoS Biology 6 (4): e99. DOI:10.1371/journal.pbio.0060099. Original artist: Mauricio Antón • File:Iceage_north-glacial_hg.png Source: http://upload.wikimedia.org/wikipedia/commons/2/26/Iceage_north-glacial_hg.png License: CC-BY-3.0 Contributors: Own work Original artist: Hannes Grobe/AWI • File:Iceage_north-intergl_glac_hg.png Source: http://upload.wikimedia.org/wikipedia/commons/e/ef/Iceage_north-intergl_glac_hg. png License: CC-BY-3.0 Contributors: Own work Original artist: Hannes Grobe/AWI • File:Iceage_south-intergl_glac_hg.png Source: http://upload.wikimedia.org/wikipedia/commons/4/44/Iceage_south-intergl_glac_hg. png License: CC-BY-3.0 Contributors: Own work Original artist: Hannes Grobe/AWI • File:Lyuba.jpg Source: http://upload.wikimedia.org/wikipedia/commons/9/90/Lyuba.jpg License: CC-BY-2.0 Contributors: IMG_2718 Original artist: Matt Howry from Ardmore, OK, USA • File:MammothVsMastodon.jpg Source: http://upload.wikimedia.org/wikipedia/commons/0/0a/MammothVsMastodon.jpg License: 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 Original artist: Original uploader was Dantheman9758 at en.wikipedia. “I created this image myself with Adobe Photoshop. 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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 Original artist: Paul Hudson from United Kingdom • File:Wooly_rhinoceros.jpg Source: http://upload.wikimedia.org/wikipedia/commons/d/d3/Wooly_rhinoceros.jpg License: Public domain 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: Own work 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: [email protected]. Do not copy this image illegally by ignoring the terms of the СС-BY-SA or GNU FDL licenses, as it is not in the public domain. Other photos see here. • File:Ямальский_мамонтенок.jpg Source: http://upload.wikimedia.org/wikipedia/commons/5/51/%D0%AF%D0%BC%D0%B0% D0%BB%D1%8C%D1%81%D0%BA%D0%B8%D0%B9_%D0%BC%D0%B0%D0%BC%D0%BE%D0%BD%D1%82%D0%B5% D0%BD%D0%BE%D0%BA.jpg License: CC-BY-SA-3.0 Contributors: Own work Original artist: This picture has been taken by Andrew Butko. Contact e-mail: [email protected]. Do not copy this image illegally by ignoring the terms of the СС-BY-SA or GNU FDL licenses, as it is not in the public domain. Other photos see here.

7.8.3 Content license

• Creative Commons Attribution-Share Alike 3.0