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The Origin and Evolution of Mammals - Moodle

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agreement about exactly when within that time<br />

frame (Brook <strong>and</strong> Bowman 2002), nor whether it<br />

occurred rapidly or spread over as much as 10,000<br />

years.<br />

<strong>The</strong> arguments in favour <strong>of</strong> human overkill as the<br />

cause <strong>of</strong> the megafaunal extinctions are tw<strong>of</strong>old<br />

(Martin 1984; Alroy 1999a). <strong>The</strong> first is the claim<br />

that the timing <strong>of</strong> the extinction in each region coincides<br />

with the timing <strong>of</strong> the arrival <strong>of</strong> human<br />

populations. This applies particularly well to North<br />

America, where the Clovis people evidently<br />

crossed the Bering l<strong>and</strong>-bridge from Asia, migrated<br />

down an ice-free corridor along the western side <strong>of</strong><br />

Alaska <strong>and</strong> Canada, <strong>and</strong> entered the southern part<br />

<strong>of</strong> the continent around 12–10,000 years ago. Details<br />

<strong>of</strong> human entry into South America are less clear,<br />

but probably occurred as a continuation <strong>of</strong> the<br />

southerly migration through North America. In<br />

northern Eurasia the evidence is more equivocal.<br />

Ne<strong>and</strong>erthals persisted there until about 30,000<br />

years ago, at which time they were replaced by<br />

modern humans (Stuart 1999). This was as much as<br />

20,000 years before many <strong>of</strong> the megafaunal extinctions<br />

actually occurred. <strong>The</strong> time <strong>of</strong> arrival <strong>of</strong><br />

humans into Australia is disputed <strong>and</strong> estimates<br />

vary from as long as 70,000 to as recently as 40,000<br />

years ago (Brook <strong>and</strong> Bowman 2002). Thus, while<br />

the possibility <strong>of</strong> a close overlap between human<br />

arrival <strong>and</strong> the Australian extinction certainly exists<br />

(Miller et al. 1999), it cannot be claimed to be substantiated<br />

by the evidence. More limited support<br />

for the overkill hypothesis is also provided by the<br />

much more accurately measured correlation<br />

between human appearance <strong>and</strong> mammal extinctions<br />

on various isl<strong>and</strong>s, notably Madagascar<br />

where most <strong>of</strong> the extinction <strong>of</strong> giant lemurs, hippopotami,<br />

the mysterious Plesiorycteropus, <strong>and</strong> also<br />

the elephant bird Aepyornis occurred soon after the<br />

arrival <strong>of</strong> humans about 1,500 years ago (Dewar<br />

1984; Burney 1999). <strong>The</strong> reason <strong>of</strong>fered by overkill<br />

enthusiasts for the low level <strong>of</strong> extinction <strong>of</strong> large<br />

mammals in Africa is attributed paradoxically to<br />

the fact that humans had been there for so long that<br />

they <strong>and</strong> the mammals had co-evolved behavioural<br />

strategies, allowing them to coexist.<br />

<strong>The</strong> second kind <strong>of</strong> argument for the overkill<br />

hypothesis is the nature <strong>of</strong> the extinction itself. <strong>The</strong><br />

end-Pleistocene was a time <strong>of</strong> net increase in habit-<br />

LIVING AND FOSSIL PLACENTALS 289<br />

able area, as the ice retreated, <strong>and</strong> so a diversification<br />

rather than an extinction phase would be expected if<br />

climate change alone were affecting the fauna.<br />

Furthermore, none <strong>of</strong> the numerous earlier episodes<br />

<strong>of</strong> mammalian extinction had ever had such a<br />

marked discriminatory effect against large body<br />

size. Human hunting activity, however, would be<br />

expected to focus more on larger mammals because<br />

<strong>of</strong> the greater ease <strong>of</strong> finding <strong>and</strong> killing them, <strong>and</strong><br />

because <strong>of</strong> the greater resource each one represents.<br />

<strong>The</strong> large mammals that did survive also supports<br />

the hypothesis. <strong>The</strong>y occupy habitats that are particularly<br />

hostile to humans, such as the musk ox <strong>and</strong><br />

caribou <strong>of</strong> the tundra, <strong>and</strong> mountain dwelling<br />

bovids, (Alroy 1999a), or else inaccessible such as the<br />

tropical forests <strong>of</strong> southern Asia or arboreal <strong>and</strong> noctumal<br />

species (Johnson 2002).<br />

<strong>The</strong> arguments in favour <strong>of</strong> climatic change as<br />

the cause <strong>of</strong> the extinction are first that a climatic<br />

change <strong>of</strong> the same order <strong>of</strong> magnitude as those<br />

that had caused earlier severe extinctions did<br />

indeed occur (Webb 1985a), <strong>and</strong> second that small<br />

populations <strong>of</strong> humans possessing very limited<br />

hunting technologies could not plausibly have had<br />

such a large effect. A number <strong>of</strong> authors have proposed<br />

more specific theories <strong>of</strong> how the climate<br />

change might have resulted in the end-Pleistocene<br />

extinction, although the picture is so complex that it<br />

is most unlikely a single, climatic factor could ever<br />

be identified as the cause (Graham 1997). Guthrie<br />

(1984) argued that the end-Pleistocene saw a continuation<br />

<strong>of</strong> the increasing seasonality that had<br />

been developing. <strong>The</strong> non-growing cold season in<br />

the temperate latitudes, <strong>and</strong> the non-growing dry<br />

season in the tropics, both increased in length. This,<br />

he proposed, caused reduced plant diversity <strong>and</strong><br />

increased plant zonation, resulting in the change<br />

from highly mosaic habitats to much more homogeneous<br />

ones dominated by large, continuous areas<br />

<strong>of</strong>, respectively, grassl<strong>and</strong>, broad-leafed forest, <strong>and</strong><br />

coniferous forest, as is seen today. Large herbivores<br />

would be the most directly affected by such a<br />

shift, followed inevitably by their large carnivore<br />

predators. Graham <strong>and</strong> Lundelius (1984) pointed<br />

to evidence <strong>of</strong> the breaking up <strong>of</strong> co-adapted biotas<br />

at this time, which they claim would have led<br />

to a variety <strong>of</strong> possible kinds <strong>of</strong> environmental<br />

stresses on individual species, such as the need for

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