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Evolution__3rd_Edition

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640 PART 5 / Macroevolution<br />

Coevolution occurs ...<br />

. . . but its relative contribution to<br />

macroevolution is unknown<br />

Summary<br />

stimulate research, and it likely accounts for some fraction of macroevolution. Just how<br />

large a fraction that is remains to be seen.<br />

22.9 Both biological and physical hypotheses should be<br />

tested on macroevolutionary observations<br />

1 Coevolution occurs when two or more lineages<br />

reciprocally influence each other’s evolution. Coadaptation<br />

between species, such as in any example of mutualism,<br />

is probably but not necessarily the result of<br />

coevolution.<br />

2 Insects and flowering plants have influenced each<br />

other’s evolution. Adaptations concerned with phytophagy<br />

(animals feeding on plants) and pollination<br />

provide examples. The evolutionary relations of flowering<br />

plants and insects are sometimes fully coevolutionary.<br />

In other cases, evolution may be sequential,<br />

Coevolution is one of several general processes that can account for evolution on a large<br />

scale a that is, macroevolution a as well as on a small scale. No one doubts its importance<br />

in microevolution, for instance in the evolution of mutualists or of parasites and<br />

their hosts. However, unanswered microevolutionary questions remain, such as the<br />

question of whether evolution in plants and insects is more often sequential evolution<br />

or fully reciprocal coevolution.<br />

The contribution of coevolution to macroevolution is more controversial. Coevolution<br />

is not the only macroevolutionary force. Many macroevolutionary events are<br />

likely caused by changes in the physical environment a climatic change, or tectonic<br />

change, or asteroid impacts such as we look at in Chapter 23. <strong>Evolution</strong>ary biologists<br />

are interested in the relative contribution of physical and biological factors, and their<br />

interaction, in driving macroevolution.<br />

One way to dramatize the issue is to ask a grand (and unanswered) question: if<br />

change in the physical environment ceased, do you think that evolution would soon<br />

come to a stop? If physical factors dominate evolution, it surely would, perhaps after a<br />

short period during which the species adjusted to the final permanent physical conditions.<br />

In the Red Queen view of evolution, the coevolutionary and biological relations<br />

between species might have a life of their own and evolution would carry on much as<br />

before after the cessation of physical environmental change. The question in its general<br />

form is too difficult for us to answer yet, but it puts under a spotlight many of the ideas<br />

that we need to examine to understand macroevolution.<br />

as plant evolution influences insect evolution but not<br />

vice versa.<br />

3 Taxa of insects and flowering plants that interact<br />

with each other may show cophylogenies, or mirrorimage<br />

phylogenies. Deviations from cophylogenies<br />

can be caused by host shifts. For instance, an insect<br />

species may colonize a new plant species that is chemically<br />

but not phylogenetically similar to the plant<br />

species it currently lives on.<br />

4 Insects and flowering plants may have promoted<br />

each other’s diversification, from the Cretaceous to the<br />

..

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