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The Origin and Evolution of Mammals - Moodle

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<strong>and</strong> thus remain in their preferred habitat.<br />

However, no such option <strong>of</strong> extending their geographical<br />

range was available to the rain-forest<br />

fauna <strong>of</strong> Central America, which merely experienced<br />

a decline in the area <strong>of</strong> their available habitat.<br />

<strong>The</strong> Great American Biotic Interchange was<br />

unique <strong>and</strong> dramatic because it brought together<br />

two biotas that had been virtually completely isolated<br />

from one another for the whole <strong>of</strong> the previous<br />

60–70 Ma. <strong>The</strong>re were nevertheless significant<br />

biotic changes in other parts <strong>of</strong> the world. <strong>The</strong><br />

severe glaciation 2.4 Ma, shortly after the establishment<br />

<strong>of</strong> the Panamanian l<strong>and</strong> bridge, was the start<br />

<strong>of</strong> the particular cycle <strong>of</strong> ice ages <strong>and</strong> interglacial<br />

periods that is still the dominant feature <strong>of</strong> the<br />

Earth’s climate. In Europe, approximately 17<br />

such ice ages are recorded in the 1.7 Ma <strong>of</strong> the<br />

Pleistocene. Typically, an ice age is associated with<br />

a reduction in the snowline by about 1000 m <strong>and</strong> a<br />

fall in average global temperature <strong>of</strong> about 6 o C.<br />

Sea-levels fall by between 100 <strong>and</strong> 150 m as water is<br />

locked up in the ice caps. <strong>The</strong> effects on the environment<br />

are worldwide, but differ from place to<br />

place. In the higher latitudes <strong>of</strong> the Holarctic continents,<br />

North America <strong>and</strong> Eurasia, there is an oscillation<br />

between temperate forest <strong>and</strong> steppe. In<br />

lower latitudes warm, moist canopied forest alternates<br />

with drier, more open woodl<strong>and</strong> <strong>and</strong> grassl<strong>and</strong><br />

conditions, <strong>and</strong> it was indeed during the Pleistocene<br />

that the fully modern version <strong>of</strong> savannah, prairie,<br />

steppe, <strong>and</strong> pampas developed. Superimposed on<br />

these general changes were particular events affecting<br />

the environment. <strong>The</strong> uplifting <strong>of</strong> the Himalayas<br />

<strong>and</strong> associated formation <strong>of</strong> the Tibetan Plateau<br />

had a pr<strong>of</strong>ound effect both directly by the increase<br />

in the area <strong>of</strong> habitable upl<strong>and</strong>, <strong>and</strong> indirectly by<br />

modifying the pattern <strong>of</strong> the Monsoon winds affecting<br />

the Asian continent. Further north, the Bering<br />

Strait dried up from time to time, allowing free dispersal<br />

<strong>of</strong> tundra <strong>and</strong> cool forest mammals, with the<br />

result that the Holarctic mammalian fauna became<br />

relatively homogenous.<br />

<strong>The</strong> outcome <strong>of</strong> this complex oscillation <strong>of</strong> environmental<br />

factors on the mammalian fauna was a<br />

series <strong>of</strong> waves <strong>of</strong> interchanges <strong>and</strong> extinctions. An<br />

enormous amount <strong>of</strong> detailed information is available<br />

from the Pleistocene palaeoenvironmental <strong>and</strong><br />

fossil record (e.g. Potts <strong>and</strong> Behrensmeyer 1992;<br />

LIVING AND FOSSIL PLACENTALS 287<br />

Graham 1997; Agustí <strong>and</strong> Wenderli 1995; Agustí<br />

<strong>and</strong> Antón 2002), <strong>and</strong> there is space here for no<br />

more than a single taste <strong>of</strong> it. Of the 130 genera <strong>of</strong><br />

Early Pleistocene mammals in North America,<br />

Webb (1985b) found that 40 <strong>of</strong> them were new. Of<br />

these, 20 were immigrants, composed <strong>of</strong> 8 from<br />

South America, <strong>and</strong> 12 from Asia via the Bering<br />

l<strong>and</strong> bridge; the latter notably included mammoths<br />

<strong>and</strong> musk oxen. <strong>The</strong> other new genera, many <strong>of</strong><br />

them cricetid rodents, had presumably evolved in<br />

situ. In similar vein, Maglio (1978) quotes figures<br />

for the Early Pleistocene <strong>of</strong> Africa. Fifty-three new<br />

genera appeared, <strong>of</strong> which about half were immigrants<br />

from the north, including members <strong>of</strong> such<br />

disparate taxa as bovids, primates, bats, pigs, <strong>and</strong><br />

rodents. Simultaneously, there were 33 generic<br />

extinctions.<br />

<strong>The</strong> end-Pleistocene: megafaunal extinction<br />

<strong>The</strong> end <strong>of</strong> the Pleistocene was marked by the<br />

abrupt extinction <strong>of</strong> many mammals, with a strong<br />

bias against species <strong>of</strong> larger body size, the mammalian<br />

megafauna (Martin <strong>and</strong> Klein 1984; McPhee<br />

1999). <strong>The</strong> timing <strong>and</strong> severity <strong>of</strong> this extinction<br />

varies from continent to continent (Fig. 7.28), <strong>and</strong><br />

its unusual features led to a controversy concerning<br />

the cause that has continued for the last century<br />

<strong>and</strong> a half (Grayson 1984). Was it due to the direct<br />

effect <strong>of</strong> human over-hunting (Martin <strong>and</strong><br />

Steadman 1999), or was it due to the kind <strong>of</strong> environmental<br />

change that caused all the mass extinction<br />

events prior to this one (Webb 1985b). Leaving<br />

aside the case <strong>of</strong> Africa <strong>and</strong> southern Asia, all mammals<br />

<strong>of</strong> body size greater than 1000 kg disappeared<br />

<strong>and</strong> about 80% <strong>of</strong> those weighing 100–1000 kg.<br />

Africa is the most glaring exception, with only<br />

about 10% loss <strong>of</strong> its mammalian megafauna; elephants,<br />

hippos, <strong>and</strong> rhinos continue to represent<br />

mammals <strong>of</strong> over 1 tonne body weight. Were this<br />

continent to be considered in isolation, no particularly<br />

significant increase in extinction would even<br />

be noted at the end <strong>of</strong> the Pleistocene. Survival <strong>of</strong><br />

mammals over 1 tonne in weight also occurred in<br />

southern Asia, but the fossil record is presently too<br />

poor to be able to quantify the extinction pattern.<br />

<strong>The</strong> timing <strong>of</strong> the extinction was also variable. In<br />

North America it is well dated to within a narrow

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