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The Origin and Evolution of Mammals - Moodle

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280 THE ORIGIN AND EVOLUTION OF MAMMALS<br />

taxa crossed the temporarily dry Bering Straits to<br />

enter North America from its northwestern corner.<br />

While the evidence is too sparse <strong>and</strong> ambiguous to<br />

support the East <strong>of</strong> Eden hypothesis for the first half<br />

<strong>of</strong> the Palaeocene, there is strong evidence that a<br />

succession <strong>of</strong> at least three, closely spaced waves <strong>of</strong><br />

immigration by this route subsequently took place,<br />

leading to the spread <strong>of</strong> several new placental<br />

groups by earliest Eocene times. Between about 57<br />

<strong>and</strong> 54 Ma, a period spanning the Palaeocene–<br />

Eocene boundary, several taxa known from earlier<br />

in the Palaeocene <strong>of</strong> Asia first occur in the North<br />

American fossil record (Beard 2002b; Bowen et al.<br />

2002). <strong>The</strong>se included the dinoceratans, arctostylopidans,<br />

tillodonts, <strong>and</strong> hyaenodontid creodonts<br />

amongst the archaic groups. It also includes members<br />

<strong>of</strong> the modern orders Perissodactyla, Artiodactyla,<br />

Rodentia, <strong>and</strong> Primates. <strong>The</strong> palaeoclimatic evidence<br />

is also compatible with the East <strong>of</strong> Eden<br />

hypothesis. A brief, intense period <strong>of</strong> global warming<br />

occurred at this time. It is indicated by a shift in<br />

13 C/ 12 C ratio that is possibly associated with the<br />

release by volcanic activity <strong>of</strong> methane <strong>and</strong> CO 2 ,<br />

causing a greenhouse warming effect (Pearson <strong>and</strong><br />

Palmer 2000). According to Kent et al. (2003), the<br />

event may have been triggered by a cometary<br />

impact. <strong>The</strong> average world temperature reached its<br />

maximum Cenozoic level, <strong>and</strong> the associated climate<br />

would be expected to have allowed the expansion<br />

<strong>of</strong> mammalian faunas into higher northern<br />

latitudes, <strong>and</strong> to open up a dispersal route from the<br />

Asian to the North American continents via the temporarily<br />

dry Bering Straits. <strong>The</strong> East <strong>of</strong> Eden<br />

hypothesis also predicts that there would have been<br />

at least some dispersal in the opposite direction,<br />

from North America to Asia. <strong>The</strong>re are, in fact, no<br />

clear examples <strong>of</strong> this except possibly the Carnivora,<br />

<strong>and</strong> nor is obvious why it was in Asia rather than<br />

elsewhere that new groups were originating in the<br />

first place. Beard (1998) suggested that both phenomena<br />

might be related simply to the much larger<br />

area <strong>of</strong> the Asian l<strong>and</strong>mass compared to the North<br />

American.<br />

<strong>The</strong> Palaeocene mammalian fauna <strong>of</strong> Europe is<br />

sufficiently similar to that <strong>of</strong> contemporary North<br />

America that they are presumed to have been interconnected<br />

through a northern Atlantic route. With<br />

the onset <strong>of</strong> the warm conditions <strong>of</strong> the Palaeocene,<br />

dispersal through what is now northern Canada <strong>and</strong><br />

Greenl<strong>and</strong> is perfectly plausible, <strong>and</strong> there is neither<br />

systematic nor palaeobiogeographical evidence that<br />

any Asian taxa arrived directly from the east.<br />

Of the Palaeocene mammals in the southern continents,<br />

vastly less is known. Early <strong>and</strong> Middle<br />

Palaeocene mammals are virtually unknown in<br />

Africa, <strong>and</strong> therefore there is no palaeontological confirmation<br />

<strong>of</strong> the taxonomic inference discussed earlier,<br />

that the Afrotheria were diversifying at that time,<br />

in isolation <strong>of</strong> the rest <strong>of</strong> the world. A little more is<br />

revealed about the early history <strong>of</strong> placentals in<br />

South America. Pantodonts <strong>and</strong> ‘condylarths’ were<br />

present in the Early Palaeocene, <strong>and</strong> presumably had<br />

arrived from the north along with the marsupials, at<br />

or soon after the end <strong>of</strong> the Cretaceous. Xenarthran<br />

fossils are absent prior to the Late Palaeocene, when<br />

they occur alongside the first <strong>of</strong> the South American<br />

ungulates. <strong>The</strong> latter, Meridiungulata, may have<br />

originated monophyletically from an immigrant<br />

North American ‘condylarth’. However, it is also<br />

possible that some <strong>of</strong> them evolved from an immigrant<br />

dinoceratan that also entered during the<br />

Palaeocene.<br />

<strong>The</strong> Indian subcontinent is another Gondwanan<br />

l<strong>and</strong>mass that may, for all that is known, have had<br />

an important role in the story. During most <strong>of</strong> the<br />

Cretaceous it was an isolated plate, drifting northwards<br />

from Africa, <strong>and</strong> eventually colliding with<br />

the Asian continent. <strong>The</strong> earliest time at which<br />

mammals might have been able to disperse from<br />

India to the main Asian l<strong>and</strong>mass is unclear. It may<br />

have been around the Palaeocene–Eocene boundary,<br />

or possibly significantly earlier, even close to<br />

the Cretaceous–Palaeocene boundary. Any placental<br />

groups that are first recorded in the Asian<br />

Palaeocene could conceivably have originated in<br />

isolation in India, <strong>and</strong>, as mentioned above, this<br />

could have included either or both the Laurasiatheria<br />

<strong>and</strong> Euarchontoglires. Unfortunately, there are as<br />

yet no suitable Palaeocene fossils from the Indian<br />

subcontinent to test this idea.<br />

<strong>The</strong> Eocene: flourishing<br />

<strong>The</strong> very warm conditions arising at the end <strong>of</strong><br />

the Palaeocene continued into the Eocene, <strong>and</strong> are<br />

associated with the time <strong>of</strong> maximum placental

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