With variations of climate, such as we know to occur from year to year, or from decade to decade, there are variations in the productiveness of the soil; and when we remember how closely interwoven are the web and woof of life, we shall see that the increased or diminished productiveness of any area will affect for good or ill all the life which that area supports. The introduction of new forms of life into an area, or their preponderance at certain periods owing to climatic or other conditions particularly favourable to them as opposed to other forms, may alter the whole balance of life in the district. We are often unable to assign any reason for the sudden increase or diminution of the numbers of a species; we can only presume that it is the result of some favourable or unfavourable change of conditions. Thus Mr. Alexander Becker[AY] has recently drawn attention to the fact that whereas for several years various species of grasshoppers appeared in great numbers in South-east Russia, there came then one year of sudden death for most of them. They were sitting motionless on the grasses and dying. He gives similar cases of butterflies for a while numerous, and then rare, and states that a squirrel common near Sarepta suddenly disappeared in the course of one summer, probably, he adds, succumbing to some contagious disease. Such is the nice balance of life, that the partial disappearance of a single form may produce remarkable and little-expected effects. Darwin amusingly showed how the clover crops might be beneficially affected by the introduction of a family of old maids into a parish. The clover is fertilized by humble-bees, the bees are preyed upon by mice; the relations between cats and mice, and between old maids and cats, are well known and familiar: more old maids, more cats; more cats, less mice; less mice, more humble-bees; more humble-bees, better fertilization. A little thing may modify the balance of life, and increase or diminish the struggle for existence, and the rigour of the process of elimination.
But when we take a more extended view of the matter, and include secular changes of climate, the possible range of variation in the struggle for existence is seen to be enormously increased. It is well known to those who have followed the progress of geology, that in early Kainozoic times a mild climate extended to within the Arctic circle, while during the glacial epoch much of the north temperate zone was fast locked in ice, and the climate of the northern hemisphere was profoundly modified. The animals in the north temperate zone were driven southwards.[AZ] Not only was there much elimination from the severe climatic conditions, but the migrants were driven southwards into areas already well stocked with life, and the competition for means of subsistence in these areas must have been rendered extremely severe. Elimination was at a maximum. Then followed the withdrawal of glacial conditions. The increasing geniality of the climate allowed an expansion of life within a given area, and the withdrawal of snow and ice further and further north set free new areas into which this expanding life could migrate and find subsistence. The hard times of the glacial period were succeeded by good times of returning warmth and an expanding area; and if, as some geologists believe, there was an inter-glacial period (or more than one such period) in the midst of the Great Ice Age, then hard times and good times alternated during the glacial epoch.
Expansion and contraction of life-areas have also been effected again and again in the course of geological history by elevations and subsidences of the land. At the beginning of Mesozoic times much of Europe was dry land. In Triassic and Rhætic times there were lakes in England and in Germany, and a warm Mediterranean Sea to the south. Subsidence of the European area brought with it a lessened land-area and an increased sea-area: bad times and increased competition for land animals; good times and a widening life-area for marine forms of life. This continued, with minor variations, till its culmination in the Cretaceous period. Then came the converse process: the land-areas increased, the sea was driven back. A good time had come for terrestrial life; the marine inhabitants of estuaries and inland seas felt the pressure of increased competition in a lessening area. And so there emerged the continental Europe of the beginning of the Kainozoic era. And it is scarcely necessary to remind those who are in any degree conversant with geology that during tertiary times there have been alternate expansions and contractions of life-areas, marine and terrestrial, the former bringing good times, the latter hard times and a heightened struggle for existence.
Now, what would be the result of this alternation of good times and hard times? During good times varieties, which would be otherwise unable to hold their own, might arise and have time to establish themselves. In an expanding area migration would take place, local segregation in the colonial areas would be rendered possible, differential elimination in the different migration-areas would produce divergence. There would be diminished elimination of neutral variations, thus affording opportunities for experimental combinations. In general, good times would favour variation and divergence.
Intermediate between good times and hard times would come, in logical order, the times in which there is neither an expansion nor a contraction of the life-area. One may suppose that these are times of relatively little change. There is neither the divergence rendered possible by the expansion of life-area, nor the heightened elimination enforced by the contraction of life-area.[BA] Elimination is steadily in progress, for the law of increase must still hold good. Divergence is still taking place, for the law of variation still obtains. But neither is at its maximum. These are the good old-fashioned times of slow and steady conservative progress. They are, perhaps, well exemplified by the fauna of the Carboniferous period, and it is not at all improbable that we are ourselves living in such a quiet, conservative period.
On the other hand, hard times would mean increased elimination. During the exhibitions at South Kensington there were good times for rats. But when the show was over, there followed times that were cruelly hard. The keenest competition for the scanty food arose, and the poor animals were forced to prey upon each other. "Their cravings for food," we read in Nature, "culminated in a fierce onslaught on one another, which was evidenced by the piteous cries of those being devoured. The method of seizing their victims was to suddenly make a raid upon one weaker or smaller than themselves, and, after overpowering it by numbers, to tear it in pieces." Elimination by competition, passing in this way into elimination by battle, would, during hard times, be increased. None but the best organized and best adapted could hope to escape. There would be no room for neutral variations, which, in the keenness of the struggle, would be relatively disadvantageous. Slightly divergent varieties, before kept apart through local segregation, would be brought into competition. The weakest would in some cases be eliminated. In other cases, the best-adapted individuals of each variety might survive. If their experiments in intercrossing, should such occur, gave rise to fertile offspring, more vigorous and better adapted than either parent-race, these would survive, and the parent-forms would be eliminated. But if such experiments in intercrossing gave rise to infertile, weakly offspring, these would be eliminated. Thus sterility between species would become fixed. Wherever, during the preceding good times, divergence had taken place in two different directions of adaptation, and some intermediate forms, fairly good in both directions, had been able to escape elimination, the chances are that these intermediates would be in hard times eliminated, and the divergent forms left in possession of the field. Wherever, during good times, a species had acquired or retained a habit of flexibility, that habit would stand it in good stead in the midst of the changes wrought by hard times; but when it had, on the other hand, acquired rigidity (like the proverbially "inflexible goose"), it would be at a disadvantage in the stress of a heightened elimination.
The alternation of good times and hard times may be illustrated by an example taken from human life. The introduction of ostrich-farming in South Africa brought good times to farmers. Whereupon there followed divergence in two directions. Some devoted increased profits to improvements upon their farms, to irrigation works which could not before be afforded, and so forth. For others increased income meant increased expenditure and an easier, if not more luxurious, mode of life. Then came hard times. Others, in Africa and elsewhere, learnt the secret of ostrich-farming. Competition brought down profits, and elimination set in—of which variety need hardly be stated.
I believe that the alternation of good times and hard times, during secular changes of climate and alternate expansions and contractions of life-areas through geological upheavals and depression of the land, has been a factor of the very greatest importance in the evolution of varied and divergent forms of life, and in the elimination of intermediate forms between adaptive variations. It now only remains in this chapter to say a few words concerning convergence, adaptation, and progress.
Convergence, which is the converse of divergence, is brought about through the adaptation of different forms of life to similar conditions of existence. The somewhat similar form of the body and fin-like limbs of fishes, of ancient reptiles (the ichthyosaurus and its allies), of whales, seals, and manatees, is a case in point. Both birds, bats, and pterodactyls have keeled breastbones for the attachment of the large muscles for flight. A whole series of analogous adaptations, as the result of analogous modes of life, are found in the placental mammals of Europe and Asia, on the one hand, and the marsupial forms of Australia on the other hand. The flying squirrel answers to the flying phalanger, the fox to the vulpine phalanger, the bear to the koala, the badger to the pouched badger, the rabbit to the bandicoot, the wolverine to the Tasmanian devil, the weasel to the pouched weasel, the rats and mice to the kangaroo rats and mice, and so on. A familiar example of convergence is to be seen in our swallows and martins, on the one hand, and the swifts on the other. Notwithstanding their superficial similarity in external form and habits, they are now generally regarded as belonging to distinct orders of birds.
These are examples of convergence.[BB] Animals of diverse descent and ancestry have, through similarity of surrounding conditions or of habits of life, become, in certain respects, assimilated. But some zoologists go further than this. They maintain that the same genus or species may, through adaptation to similar circumstances, be derived from dissimilar ancestors. Some palæontologists, for example, believe that the horse has been independently evolved along parallel lines in Europe and in America. Professor Cope considers that in the one continent Protohippus, and in the other Hipparion, was the immediate ancestor of Equus. The probabilities are, however, so strongly against such a view, that it cannot be accepted until substantiated by stronger evidence than is yet forthcoming.