In the biomorph model we assumed that this kind of multidimensional improvement could not occur. To recapitulate on why that was a reasonable assumption, to make an eye from nothing you need not just one improvement but a large number of improvements. Any one of these improvements is pretty improbable by itself, but not so improbable as to be impossible. The greater the number of simultaneous improvements we consider, the more improbable is their simultaneous occurrence. The coincidence of their simultaneous occurrence is equivalent to leaping a large distance across Biomorph Land, and happening to land on one particular, predesignated spot. If we choose to consider a sufficiently large number of improvements, their joint occurrence becomes so improbable as to be, to all intents and purposes, impossible. The argument has already been sufficiently made, but it may be helpful to draw a distinction between two kinds of hypothetical macromutation, both of which appear to be ruled out by the complexity argument but only one of which, in fact, is ruled out by the complexity argument. I label them, for reasons that will become clear, Boeing 747 macromutations and Stretched DC8 macromutations.
Boeing 747 macromutations are the ones that really are ruled out by the complexity argument just given. They get their name from the astronomer Sir Fred Hoyle’s memorable misunderstanding of the theory of natural selection. He compared natural selection, in its alleged improbability, to a hurricane blowing through a junkyard and chancing to assemble a Boeing 747. As we saw in Chapter 1, this is an entirely false analogy to apply to natural selection, but it is a very good analogy for the idea of certain kinds of macromutation giving rise to evolutionary change. Indeed, Hoyle’s fundamental error was that he, in effect, thought (without realizing it) that the theory of natural selection did depend upon macromutation. The idea of a single macromutation’s giving rise to a fully functioning eye with the properties listed above, where there was only bare skin before, is, indeed, just about as improbable as a hurricane assembling a Boeing 747. This is why I refer to this kind of hypothetical macromutation as a Boeing 747 macromutation.
Stretched DC8 macromutations are mutations that, although they may be large in the magnitude of their effects, turn out not to be large in terms of their complexity. The Stretched DC8 is an airliner that was made by modifying an earlier airliner, the DC8. It is like a DC8, but with an elongated fuselage. It was an improvement at least from one point of view, in that it could carry more passengers than the original DC8. The stretching is a large increase in length, and in that sense is analogous to a macromutation. More interestingly, the increase in length is, at first sight, a complex one. To elongate the fuselage of an airliner, it is not enough just to insert an extra length of cabin tube. You also have to elongate countless ducts, cables, air tubes and electric wires. You have to put in lots more seats, ashtrays, reading lights, 12-channel music selectors and fresh-air nozzles. At first sight there seems to be much more complexity in a Stretched DC8 than there is in an ordinary DC8, but is there really? The answer is no, at least to the extent that the ‘new’ things in the stretched plane are just ‘more of the same’. The biomorphs of Chapter 3 frequently show macromutations of the Stretched DC8 variety.
What has this to do with mutations in real animals? The answer is that some real mutations cause large changes that are very like the change from DC8 to Stretched DC8, and some of these, although in a sense ‘macro’ mutations, have definitely been incorporated in evolution. Snakes, for instance, all have many more vertebrae than their ancestors. We could be sure of this even if we didn’t have any fossils, because snakes have many more vertebrae than their surviving relatives. Moreover, different species of snakes have different numbers of vertebrae, which means that vertebral number must have changed in evolution since their common ancestor, and quite often at that.
Now, to change the number of vertebrae in an animal, you need to do more than just shove in an extra bone. Each vertebra has, associated with it, a set of nerves, a set of blood vessels, a set of muscles etc., just as each row of seats in an airliner has a set of cushions, a set of head rests, a set of headphone sockets, a set of reading-lights with their associated cables etc. The middle part of the body of a snake, like the middle part of the body of an airliner, is composed of a number of segments, many of which are exactly like each other, however complex they all individually may be. Therefore, in order to add new segments, all that has to be done is a simple process of duplication. Since there already exists genetic machinery for making one snake segment — genetic machinery of great complexity, which took many generations of step-by-step, gradual evolution to build up — new identical segments may easily be added by a single mutational step. If we think of genes as ‘instructions to a developing embryo’, a gene for inserting extra segments may read, simply, ‘more of the same here’. I imagine that the instructions for building the first Stretched DC8 were somewhat similar.
We can be sure that, in the evolution of snakes, numbers of vertebrae changed in whole numbers rather than in fractions. We cannot imagine a snake with 26.3 vertebrae. It either had 26 or 27, and it is obvious that there must have been cases when an offspring snake had at least one whole vertebra more than its parents did. This means that it had a whole extra set of nerves, blood vessels, muscle blocks, etc. In a sense, then, this snake was a macro-mutant, but only in the weak ‘Stretched DC8’ sense. It is easy to believe that individual snakes with half a dozen more vertebrae than their parents could have arisen in a single mutational step. The ‘complexity argument’ against saltatory evolution does not apply to Stretched DC8 macromutations because, if we look in detail at the nature of the change involved, they are in a real sense not true macromutations at all. They are only macromutations if we look, naïvely, at the finished product, the adult. If we look at the processes of embryonic development they turn out to be micromutations, in the sense that only a small change in the embryonic instructions had a large apparent effect in the adult. The same goes for antennapaedia in fruitflies and the many other so-called ‘homeotic mutations’.
This concludes my digression on macromutation and saltatory evolution. It was necessary, because the theory of punctuated equilibria is frequently confused with saltatory evolution. But it was a digression, because the theory of punctuated equilibria is the main topic of this chapter, and that theory in truth has no connection with macromutation and true saltation.
The ‘gaps’ that Eldredge and Gould and the other ‘punctuationists’ are talking about, then, have nothing to do with true saltation, and they are much much smaller gaps than the ones that excite creationists. Moreover, Eldredge and Gould originally introduced their theory, not as radically and revolutionarily antipathetic to ordinary, ‘conventional’ Darwinism — which is how it later came to be sold — but as something that followed from long-accepted conventional Darwinism, properly understood. To gain this proper understanding, I’m afraid we need another digression, this time into the question of how new species originate, the process known as ‘speciation’.
Darwin’s answer to the question of the origin of species was, in a general sense, that species were descended from other species. Moreover, the family tree of life is a branching one, which means that more than one modern species can be traced back to one ancestral one. For instance, lions and tigers are now members of different species, but they have both sprung from a single ancestral species, probably not very long ago. This ancestral species may have been the same as one of the two modern species; or it may have been a third modern species; or maybe it is now extinct. Similarly, humans and chimps now clearly belong to different species, but their ancestors of a few million years ago belonged to one single species. Speciation is the process by which a single species becomes two species, one of which may be the same as the original single one.
The reason speciation is thought to be a difficult problem is this. All the members of the single would-be ancestral species are capable of interbreeding with one another: indeed, to many people, this is what is meant by the phrase ‘single species’. Therefore, ev
ery time a new daughter species begins to be ‘budded off’, the budding off is in danger of being frustrated by interbreeding. We can imagine the would-be ancestors of the lions and the would-be ancestors of the tigers failing to split apart because they keep interbreeding with one another and therefore staying similar to one another. Don’t, incidentally, read too much into my use of words like ‘frustrated’, as though the ancestral lions and tigers, in some sense, ‘wanted’ to separate from each other. It is simply that, as a matter of fact, species obviously have diverged from one another in evolution, and at first sight the fact of interbreeding makes it hard for us to see how this divergence came about.
It seems almost certain that the principal correct answer to this problem is the obvious one. There will be no problem of interbreeding if the ancestral lions and the ancestral tigers happen to be in different parts of the world, where they can’t interbreed with each other. Of course, they didn’t go to different continents in order to allow themselves to diverge from one another: they didn’t think of themselves as ancestral lions or ancestral tigers! But, given that the single ancestral species spread to different continents anyway, say Africa and Asia, the ones that happened to be in Africa could no longer interbreed with the ones that happened to be in Asia because they never met them. If there was any tendency for the animals on the two continents to evolve in different directions, either under the influence of natural selection or under the influence of chance, interbreeding no longer constituted a barrier to their diverging and eventually becoming two distinct species.
I have spoken of different continents to make it clear, but the principle of geographical separation as a barrier to interbreeding can apply to animals on different sides of a desert, a mountain range, a river, or even a motorway. It can also apply to animals separated by no barrier other than sheer distance. Shrews in Spain cannot interbreed with shrews in Mongolia, and they can diverge, evolutionarily speaking, from shrews in Mongolia, even if there is an unbroken chain of interbreeding shrews connecting Spain to Mongolia. Nevertheless the idea of geographical separation as the key to speciation is clearer if we think in terms of an actual physical barrier, such as the sea or a mountain range. Chains of islands, indeed, are probably fertile nurseries for new species.
Here, then, is our orthodox neo-Darwinian picture of how a typical species is ‘born’, by divergence from an ancestral species. We start with the ancestral species, a large population of rather uniform, mutually interbreeding animals, spread over a large land mass. They could be any sort of animal, but let’s carry on thinking of shrews. The landmass is cut in two by a mountain range. This is hostile country and the shrews are unlikely to cross it, but it is not quite impossible and very occasionally one or two do end up in the lowlands on the other side. Here they can flourish, and they give rise to an outlying population of the species, effectively cut off from the main population. Now the two populations breed and breed separately, mixing their genes on each side of the mountains but not across the mountains. As time goes by, any changes in the genetic composition of one population are spread by breeding throughout that population but not across to the other population. Some of these changes may be brought about by natural selection, which may be different on the two sides of the mountain range: we should hardly expect weather conditions, and predators and parasites, to be exactly the same on the two sides. Some of the changes may be due to chance alone. Whatever the genetic changes are due to, breeding tends to spread them within each of the two populations, but not between the two populations. So the two populations diverge genetically: they become progressively more unlike each other.
They become so unlike each other that, after a while, naturalists would see them as belonging to different ‘races’. After a longer time, they will have diverged so far that we should classify them as different species. Now imagine that the climate warms up so that travel through the mountain passes becomes easier and some of the new species start trickling back to their ancestral homelands. When they meet the descendants of their long-lost cousins, it turns out that they have diverged so far in their genetic makeup that they can no longer successfully interbreed with them. If they do hybridize with them the resulting offspring are sickly, or sterile like mules. So natural selection penalizes any predilection, on the part of individuals on either side, towards hybridizing with the other species or even race. Natural selection thereby finishes off the process of ‘reproductive isolation’ that began with the chance intervention of a mountain range. ‘Speciation’ is complete. We now have two species where previously there was one, and the two species can coexist in the same area without interbreeding with one another.
Actually, the likelihood is that the two species would not coexist for very long. This is not because they would interbreed but because they would compete. It is a widely accepted principle of ecology that two species with the same way of life will not coexist for long in one place, because they will compete and one or other will be driven extinct. Of course our two populations of shrews might no longer have the same way of life; for instance, the new species, during its period of evolution on the other side of the mountains, might have come to specialize on a different kind of insect prey. But if there is significant competition between the two species, most ecologists would expect one or other species to go extinct in the area of overlap. If it happened to be the original, ancestral species that was driven extinct, we should say that it had been replaced by the new, immigrant species.
The theory of speciation resulting from initial geographical separation has long been a cornerstone of mainstream, orthodox neo-Darwinism, and it is still accepted on all sides as the main process by which new species come into existence (some people think there are others as well). Its incorporation into modern Darwinism was largely due to the influence of the distinguished zoologist Ernst Mayr. What the ‘punctuationists’ did, when they first proposed their theory, was to ask themselves: Given that, like most neo-Darwinians, we accept the orthodox theory that speciation starts with geographical isolation, what should we expect to see in the fossil record?
Recall the hypothetical population of shrews, with a new species diverging on the far side of a mountain range, then eventually returning to the ancestral homelands and, quite possibly, driving the ancestral species extinct. Suppose that these shrews had left fossils; suppose even that the fossil record was perfect, with no gaps due to the unfortunate omission of key stages. What should we expect these fossils to show us? A smooth transition from ancestral species to daughter species? Certainly not, at least if we are digging in the main landmass where the original ancestral shrews lived, and to which the new species returned. Think of the history of what actually happened in the main landmass. There were the ancestral shrews, living and breeding happily away, with no particular reason to change. Admittedly their cousins the other side of the mountains were busy evolving, but their fossils are all on the other side of the mountain so we don’t find them in the main landmass where we are digging. Then, suddenly (suddenly by geological standards, that is), the new species returns, competes with the main species and, perhaps, replaces the main species. Suddenly the fossils that we find as we move up through the strata of the main landmass change. Previously they were all of the ancestral species. Now, abruptly and without visible transitions, fossils of the new species appear, and fossils of the old species disappear.
The ‘gaps’, far from being annoying imperfections or awkward embarrassments, turn out to be exactly what we should positively expect, if we take seriously our orthodox neo-Darwinian theory of speciation. The reason the ‘transition’ from ancestral species to descendant species appears to be abrupt and jerky is simply that, when we look at a series of fossils from any one place, we are probably not looking at an evolutionary event at all: we are looking at a migrational event, the arrival of a new species from another geographical area. Certainly there were evolutionary events, and one species really did evolve, probably gradually, from another. But in order t
o see the evolutionary transition documented in the fossils we should have to dig elsewhere — in this case on the other side of the mountains.
The point that Eldredge and Gould were making, then, could have been modestly presented as a helpful rescuing of Darwin and his successors from what had seemed to them an awkward difficulty. Indeed that is, at least in part, how it was presented — initially. Darwinians had always been bothered by the apparent gappiness of the fossil record, and had seemed forced to resort to special pleading about imperfect evidence. Darwin himself had written:
The geological record is extremely imperfect and this fact will to a large extent explain why we do not find interminable varieties, connecting together all the extinct and existing forms of life by the finest graduated steps. He who rejects these views on the nature of the geological record, will rightly reject my whole theory.
Eldredge and Gould could have made this their main message: Don’t worry Darwin, even if the fossil record were perfect you shouldn’t expect to see a finely graduated progression if you only dig in one place, for the simple reason that most of the evolutionary change took place somewhere else! They could have gone further and said:
Darwin, when you said that the fossil record was imperfect, you were understating it. Not only is it imperfect, there are good reasons for expecting it to be particularly imperfect just when it gets interesting, just when evolutionary change is taking place; this is partly because evolution usually occurred in a different place from where we find most of our fossils; and it is partly because, even if we are fortunate enough to dig in one of the small outlying areas where most evolutionary change went on, that evolutionary change (though still gradual) occupies such a short time that we should need an extra rich fossil record in order to track it!
But no, instead they chose, especially in their later writings in which they were eagerly followed by journalists, to sell their ideas as being radically opposed to Darwin’s and opposed to the neo-Darwinian synthesis. They did this by emphasizing the ‘gradualism’ of the Darwinian view of evolution as opposed to the sudden, jerky, sporadic ‘punctuationism’ of their own. They even, especially Gould, saw analogies between themselves and the old schools of ‘catas-trophism’ and ‘saltationism’. Saltationism we have already discussed. Catastrophism was an eighteenth- and nineteenth-century attempt to reconcile some form of creationism with the uncomfortable facts of the fossil record. Catastrophists believed that the apparent progression of the fossil record really reflected a series of discrete creations, each one terminated by a catastrophic mass extinction. The latest of these catastrophes was Noah’s flood.