Gould, The Wheel of Fortune and the Wedge of Progress

(or at least to dilute it to insipidity) with his favorite argument about "the imperfection of the geological record." A hopelessly inadequate record can compress millions of years of missing data into an apparent "event." If we could recover the full flow of time, the individual items of a "mass extinction" would spread out on both sides of illusory suddenness, indicating a much longer period of successive disappearances at a rate little, if at all, accelerated beyond the usual tempo of wedging in ordinary times.

A more popular argument (the ghostbuster of accommodation) admits the unusual character of mass extinctions, acknowledging a substantial acceleration in the tempo of death, while arguing that the environmental stresses of these parlous times do not introduce a new regime in the causality of death, but rather, only accentuate the power of the wedge. In these worst of times, the motor of biological competition runs faster as increased stress drives up the intensity of struggle. Mass extinction only "turns up the gain" on business as usual. If the "ancient and beaten" make their exit in ordinary times as superior wedges push themselves into the bustling economy of nature, then the rate of departure can only accelerate in the tough moments of mass extinction when dogs eat dogs and men must be men.

This honing of the wedge provides the traditional context, usually not well explained in general writing on mass extinction, that has made all the recent news about truly catastrophic causes based on extraterrestrial impact so controversial and so threatening. In 1980, the father and son, physicist and geologist team of Luis and Walter Alvarez, along with Frank Asaro and Helen Michel, first published their evidence that a large extraterrestrial object struck the earth some 65 million years ago and triggered the great Cretaceous extinction.

The explosion of fruitful scientific work that this hypothesis has engendered in just nine short years must match the force or the impact itself. The Alvarezes' hypothesis has opened the small and arcane paleontological field of mass extinction into a grand arena of interdisciplinary cooperation. In October 1988, I spent three wonderful days at Snowbird, Utah, listening to several hundred excited scientists, from geochemists to planetary physicists, paleontologists to climatic modelers, ponder the causes of mass extinction. Debate on the Cretaceous event still swirls about two crucial issues: timing (one impact or many, one moment or an extended period of multiple bombardments) and cause (massive volcanic effusions have been proposed as an alternative, but impact now holds the upper hand according to most workers in this field). Moreover, the case for impact has not been established (or precluded) for other mass extinctions, so we do not know whether the Alvarezes' persuasive account or the episode ranks as the explanation for a single event or a general theory of mass extinction.

Nonetheless, I think we now know enough to summarize the new views in a statement that must suggest a radically revised perspective on the evolutionary meaning of mass extinction. These events, with their catastrophic causes, are more frequent, more sudden, more profound in their extent, and more different (from normal times) in their results than we had imagined. Mass extinction does not just turn up the gain on competition, so that wedging can proceed more ruthlessly and more efficiently; mass extinction entrains new causes that impart a distinctive stamp to evolutionary results. And if the history of life owes its shape more to the differential success of groups in surviving mass extinction than to accumulated victories by wedging in normal times, then a major component of Darwin's world view--and the only sensible argument that he could supply for our deepest, culturally bound hope of progress--has been compromised or even overturned.

I can envision two models of causality in mass extinction that challenge wedging and consequent progress as a prominent vector of life. In the random model, species live or die by the roll of the dice and the luck of the draw; success reduces to little more than being in the right place at the right time when the comets hit, the fires roar, the earth darkens, and the oceans are poisoned. In the different rules model, species live or die for definite and specifiable reasons. But the causes of success are quirky and fortuitous with respect to initial reasons for evolving the features that secure survival. The wedge operates in normal times between mass extinctions. Organisms evolve features to enhance success in continuous ecological struggle. The cause of mass extinction then hits in all its sudden fury. Certain features are the passkeys to survival--tolerance of extreme climatic stress, for example. But these features must have evolved during normal times dominated by wedging. And they must, in principle, have arisen for reasons unrelated to their later (and lucky) use in seeing their possessors through the unanticipated debacle of mass extinction. (I say "in principle" because, unless our basic views on causality are seriously awry and the future can control the present, organisms cannot evolve a feature for its potential utility several million years later when the comet hits.)

I see a role for the random model especially in the most severe events. If we accept David M. Raup's estimate of 96 percent species extinction in the Permian debacle, then entire groups may have been lost by something akin to unalloyed bad luck. Yet I remain committed enough to a more conventional view of causality to think that, of my two proposals for radical reform, the different rules model must apply more often. We can specify causes for differential survival in mass extinction, but the features that secure success must have evolved for unrelated reasons, usually in the regime of wedging during normal times.

As Kant told us that concepts without percepts are empty, and as Harry Truman said, "Show me, I'm from Missouri," the different rules model, however interesting or elegant in the abstract, will have no power without empirical documentation from the fossil record. I have written a peculiar essay with a prologue more than twice as long as the main point--the proposed example of the different rules model to follow. Still, what comes next is the heart of this essay, not its epilogue.

If we wish to illustrate the different rules model, we might begin with the most prominent group to die and the most widely cited cause of extinction. Most people think first of dinosaurs when they ponder the Cretaceous extinction. But our clearest and most extensive evidence comes from the opposite end of the discredited chain of being--the single-celled oceanic plankton. Extinctions are so prominent in these creatures that paleontologists speak of a "plankton line" marking the rapid and simultaneous termination of numerous lineages. As for "killing scenarios" in extraterrestrial impact, the most widely cited reason (though causes must have been complex, interacting, numerous, and varied) invokes a device of Moses against Pharaoh: a thick darkness over all the land, even darkness which might be felt. An impacting comet or asteroid, the argument runs, would excavate a massive crater and send aloft a thick cloud of particles that would envelop the earth in sufficient darkness to shut down all photosynthesis for several months.

A tie between the scenario of darkness and the death of plankton seems easy to formulate. A few months of darkness might not faze a tree (especially if the impact occurred in winter); a plant that lives for decades might shut down its factories for a few months. And even if plants die, seeds can often survive to germinate when the dust cloud dissipates. But the

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