A heated debate has enlivened recent studies of evolution. Darwin' s originalthesis, and the viewpoint supported by evolutionary gradualists, is that species change continuously but slowly and in small increments. Such changes are all but invisible over the short time scale of modern observations, and, it is argued, they are usually obscured by innumerable gaps in the imperfect fossilrecord. Gradualism, with its stress on the slow pace of change, is a comforting position, repeated over and over again in generations of textbooks. By the early twentieth century, the question about the rate of evolution had been answered in favor of gradualism to most biologists' satisfaction.
Sometimes a closed question must be reopened as new evidence or new arguments based on old evidence come to light. In 1972 paleontologist Stephen Jay Gould and Niles Eldredge challenged conventional wisdom with an opposing viewpoint, the punctuated equilibriumhypothesis, which posits that species give rise to new species in relatively sudden bursts, without a lengthytransition period. These episodes of rapid evolution are separated by relatively long static spans during which a species may hardly change at all.
The punctuated equilibriumhypothesis attempts to explain a curious feature of the fossilrecord --- one that has been familiar to paleontologist for more than a century but has usually been ignored. Many speciesappear to remain unchanged in the fossilrecord for millions of years --- a situation that seems to be at odds with Darwin' s model of continuous change. Intermediated fossil forms, predicted by gradualism, are typically lacking. In most localities a givenspecies of clam or coral persists essentially unchanged throughout a thick formation of rock, only to be replaced suddenly by a new and different species.
The evolution of North American horse, which was once presented as a classic textbook example of gradualevolution, is now providing equally compellingevidence for punctuated equilibrium. A convincing 50-million-year sequence of modern horse ancestors --- each slightly larger, with more complex teeth, a longer face, and a more prominent central toe ---seemed to provide strong support for Darwin' s contention that speciesevolvegradually. But close examination of those fossil deposits now reveals a somewhat different story. Horses evolved in discrete steps, each of which persisted almost unchanged for millions of years and was eventually replaced by a distinctive newer model. The four-toed Eohippus preceded the three-toed Miohippus, for example, but North American fossilevidence suggests a jerky, uneventransition between the two. If evolution had been a continuous, gradualprocess, one might expect that almost every fossilspecimen would be slightly different from every year.
If it seems difficult to conceive how major changes could occur rapidly, consider this: an alteration of a single gene in files is enough to turn a normal fly with a single pair of wings into one that has two pairs of wings.
The question about the rate of evolution must now be turned around: does evolution ever proceedgradually, or does it always occur in short bursts? Detailed field studies of thick rock formations containing fossils provide the best potential tests of the competing theories.
Occasionally , a sequence of fossil-rich layers of rock permits a comprehensive look at one type of organism over a long period of time. For example, Peter Sheldon' s studies of trilobites, a now extinctmarine animal with a segmented body, offer a detailed glimpse into three million years of evolution in one marine environment. In that study, each of eight different trilobite species was observed to undergo a gradual change in the number of segments --- typically an increase of one or two segments over the whole time interval. No significant discontinuous were observed, leading Sheldon to conclude that environmental conditions were quite stable during the period he examined.
Similar exhaustive studies are required for many different kinds of organisms from many different periods.
Most researchers expect to find that both modes of transition from one species
to another are at work in evolution. Slow, continuous change may be the norm during periods of environmentalstability, while rapid evolution of new species occurs during periods of environment
stress. But a lot more studies like Sheldon' s are needed before we can say for sure.
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