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Punctuated equilibrium (or punctuated equilibria) is a theory in evolutionary biology which states that most sexually reproducing species will show little to no evolutionary change throughout their history. When evolution does occur, it happens sporadically (by splitting) and occurs relatively quickly compared to the species' full duration on earth. Punctuated equilibrium is commonly contrasted against the theory of phyletic gradualism, which hypothesizes that most evolution occurs uniformly and by the steady and gradual transformation of whole lineages (anagenesis).

Discussion

Punctuated equilibrium originated as an extension of Ernst Mayr's concept of genetic revolutions by peripatric and allopatric speciation. Although the workings of the theory were proposed and specifically identified by Mayr in 1954, most historians of science recognize Niles Eldredge and Stephen Jay Gould's 1972 paper as the principal source of its acceptance (by both paleontologists and evolutionists) and as the foundational document of a new and serious paleontological research program (Mayr 1992: 25-26, Shermer 2001: 102-113). Punctuated equilibrium differed from Mayr simply in that Eldredge and Gould had placed considerably greater emphasis on stasis, whereas Mayr was generally concerned with explaining the morphological discontinuity (or punctuational patterns) found in the fossil record.

Tempo and Mode

Ernst Mayr's paper "Change of genetic environment and evolution" (1954) emphasized the homogenizing effects of gene flow and the stabilizing influence of large interbreeding populations. These populations exemplified "ecotypic variation." Peripherally isolated populations, in contrast, possess "typostrophic variation" which "have the characteristic features of incipient species, but what is more important they often are species or incipient species of an entirely new type. That is, they may have morphological or ecological features that deviate quite strikingly and unexpectedly from the parental 'pattern' " (1954:160)

Stephen Jay Gould summarized the theory, and its consequences for punctuated equilibrium, in a 1977 essay for Natural History:

"A new species can arise when a small segment of the ancestral population is isolated at the periphery of the ancestral range. Large, stable central populations exert a strong homogenizing influence. New and favorable mutations are diluted by the sheer bulk of the population through which they must spread. They may build slowly in frequency, but changing environments usually cancel their selective value long before they reach fixation. Thus, phyletic transformation in large populations should be very rare�as the fossil record proclaims. But small, peripherally isolated groups are cut off from their parental stock. They live as tiny populations in geographic corners of the ancestral range. Selective pressures are usually intense because peripheries mark the edge of ecological tolerance for ancestral forms. Favorable variations spread quickly. Small peripheral isolates are a laboratory of evolutionary change.

"What should the fossil record include if most evolution occurs by speciation in peripheral isolates? Species should be static through their range because our fossils are the remains of large central populations. In any local area inhabited by ancestors, a descendant species should appear suddenly by migration from the peripheral region in which it evolved. In the peripheral region itself, we might find direct evidence of speciation, but such good fortune would be rare indeed because the event occurs so rapidly in such a small population. Thus, the fossil record is a faithful rendering of what evolutionary theory predicts, not a pitiful vestige of a once bountiful tale." (1980:184)

Relation to Darwinism

The sudden appearance and lack of substantial gradual change of most species in the geologic record�from their initial appearance until their extinction�has long been noted, including by Charles Darwin (1859:301, 1871:119-120) who appealed to the imperfection of the record as the favored explanation. Nevertheless, with the influence of catastrophism, Darwin needed to forcefully stress the gradual nature of evolution. It is often incorrectly assumed that he insisted that the rate of change must be constant, or nearly so. Though in The Origin of Species Darwin wrote that "the periods during which species have undergone modification, though long as measured in years, have probably been short in comparison with the periods during which they retain the same form." (1872:619) Thus punctuationism in general is consistent with Darwin's conception of evolution, and with the independent proposals of natural selection by William Charles Wells, Patrick Matthew, Charles Darwin and Alfred Russel Wallace.

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