Cope's rule

Cope's rule, named after American paleontologist Edward Drinker Cope, postulates that population lineages tend to increase in body size over evolutionary time. It was never actually stated by Cope, although he favoured the occurrence of linear evolutionary trends. It is sometimes also known as the Cope-Depéret rule, because Charles Depéret explicitly advocated the idea.Theodor Eimer had also done so earlier. The term "Cope's rule" was apparently coined by Bernhard Rensch, based on the fact that Depéret had "lionized Cope" in his book. While the rule has been demonstrated in many instances, it does not hold true at all taxonomic levels, or in all clades. Larger body size is associated with increased fitness for a number of reasons, although there are also some disadvantages both on an individual and on a clade level: clades comprising larger individuals are more prone to extinction, which may act to limit the maximum size of organisms.

Directional selection appears to act on organisms' size, whereas it exhibits a far smaller effect on other morphological traits, though it is possible that this perception may be a result of sample bias. This selectional pressure can be explained by a number of advantages, both in terms of mating success and survival rate.

For example, larger organisms find it easier to avoid or fight off predators and capture prey, to reproduce, to kill competitors, to survive temporary lean times, and to resist rapid climatic changes. They may also potentially benefit from better thermal efficiency, increased intelligence, and a longer lifespan.

Offsetting these advantages, larger organisms require more food and water, and shift from r to K-selection. Their longer generation time means a longer period of reliance on the mother, and on a macroevolutionary scale restricts the clade's ability to evolve rapidly in response to changing environments.

Left unfettered, the trend of ever-larger size would produce organisms of gargantuan proportions. Therefore, some factors must limit this process. At one level, it is possible that the clade's increased vulnerability to extinction, as its members become larger, means that no taxon survives long enough for individuals to reach huge sizes. There are probably also physically imposed limits to the size of some organisms; for instance, insects must be small enough for oxygen to diffuse to all parts of their bodies, flying birds must be light enough to fly, and the length of giraffes' necks may be limited by the amount of pressure it is possible for their hearts to generate. Finally, there may be a competitive element, in that changes in size are necessarily accompanied by changes in ecological niche. For example, terrestrial carnivores over 21 kg almost always prey on organisms larger, not smaller, than themselves. If such a niche is already occupied, competitive pressure may oppose the directional selection. The three Canidae clades show a trend towards larger size before becoming extinct.

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