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Evolution__3rd_Edition

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..<br />

Figure 23.1<br />

(a) The number of families of<br />

mammals in North America<br />

increased abruptly in the<br />

Paleocene and Eocene, after<br />

which it has remained constant.<br />

(Alroy 1999 shows a similar<br />

pattern for an updated,<br />

taxonomically finer, dataset.)<br />

(b) The number of families of<br />

bivalves has increased steadily<br />

through time. Redrawn, by<br />

permission of the publisher,<br />

from Stanley (1979). © 1979<br />

WH Freeman & Company.<br />

The Cambrian explosion may have<br />

followed an adaptive breakthrough<br />

Number of families<br />

(a) Mammalia (b) Bivalvia<br />

120<br />

120<br />

100<br />

80<br />

60<br />

40<br />

20<br />

0<br />

L. Cret. Pal. Eoc. Olig. Mio.<br />

Pli. P.<br />

100<br />

80<br />

60<br />

40<br />

20<br />

0<br />

CHAPTER 23 / Extinction and Radiation 645<br />

Ord. Sil. Dev. Miss. Pen. Perm. Tri. Jur. Cret. Pal. Neog.<br />

100 80 60 40 20 0 500 400 300 200 100 0<br />

Time (Myr BP)<br />

Time (Myr BP)<br />

3. Replacement of competitors. One taxon may radiate if it is adaptively superior to<br />

its competitors. The superior taxon will take over from the inferior one, which<br />

will be driven extinct. The superior taxon may have become superior because of<br />

environmental change, or because it evolved a new, superior adaptation. We look<br />

at replacements in Section 23.7.<br />

4. Adaptive breakthroughs. A taxon may evolve a new adaptation that allows it to outcompete<br />

another taxon (see point 3 above), or may allow it to exploit a previously<br />

unexploited resource. For instance, we saw in Sections 18.5 and 18.6 (pp. 538–42)<br />

how plants and animals colonized the land as they evolved a set of appropriate<br />

adaptations.<br />

The colonization of land was somewhat analogous to the colonization of a new<br />

island chain (point 1 above). However, we can distinguish between colonization of<br />

an existing area, made possible by the evolution of a new adaptation, and colonization<br />

of a new area that requires no new adaptation. The colonization of land required new<br />

adaptations for support, respiration, and water retention. By contrast, the finch that<br />

first colonized the Galápagos probably did not have to evolve a new adaptation before<br />

it could colonize the island. After the colonization of land, plants and animals both<br />

underwent adaptive radiations. These radiations were made possible by an adaptive<br />

breakthrough.<br />

The proliferation of animals with hard skeletons in the “Cambrian explosion”<br />

(Section 18.4, p. 535) is one of the most important adaptive radiations in the history<br />

of life. But the reason why it occurred remains uncertain. One hypothesis proposes<br />

that predators evolved escalated skills around this time, making hard skeletons advantageous.<br />

If so, the radiation may be an example of a replacement following changed<br />

conditions (factor 3 in the list above). The radiation of animals with hard parts may<br />

have occurred as they replaced soft-bodied predecessors.<br />

Adaptive radiations can be understood in terms of Darwin’s principle of divergence<br />

(Section 16.8, p. 487). Darwin was interested in why evolution usually shows a diverging,<br />

tree-like pattern. He explained the pattern by competition. More similar forms<br />

will compete more strongly than less similar forms, which tends to “push” species<br />

apart during evolution. Species diverge to escape competition. Darwin’s principle of<br />

divergence likely needs to be slightly modified to incorporate the modern theory of

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