Darwinism (Wallace)/Chapter V
CHAPTER V
NATURAL SELECTION BY VARIATION AND SURVIVAL OF THE FITTEST
Effect of struggle for existence under unchanged conditions—The effect under change of conditions—Divergence of character—In insects—In birds—In mammalia—Divergence leads to a maximum of life in each area—Closely allied species inhabit distinct areas—Adaptation to conditions at various periods of life—The continued existence of low forms of life—Extinction of low types among the higher animals—Circumstances favourable to the origin of new species—Probable origin of the dippers—The importance of isolation—On the advance of organisation by natural selection—Summary of the first five chapters.
In the preceding chapters we have accumulated a body of facts and arguments which will enable us now to deal with the very core of our subject—the formation of species by means of natural selection. We have seen how tremendous is the struggle for existence always going on in nature owing to the great powers of increase of all organisms; we have ascertained the fact of variability extending to every part and organ, each of which varies simultaneously and for the most part independently; and we have seen that this variability is both large in its amount in proportion to the size of each part, and usually affects a considerable proportion of the individuals in the large and dominant species. And, lastly, we have seen how similar variations, occurring in cultivated plants and domestic animals, are capable of being perpetuated and accumulated by artificial selection, till they have resulted in all the wonderful varieties of our fruits, flowers, and vegetables, our domestic animals and household pets, many of which differ from each other far more in external characters, habits, and instincts than do species in a state of nature. We have now to inquire whether there is any analogous process in nature, by which wild animals and plants can be permanently modified and new races or new species produced.
Effect of Struggle for Existence under Unchanged Conditions.
Let us first consider what will be the effect of the struggle for existence upon the animals and plants which we see around us, under conditions which do not perceptibly vary from year to year or from century to century. We have seen that every species is exposed to numerous and varied dangers throughout its entire existence, and that it is only by means of the exact adaptation of its organisation—including its instincts and habits—to its surroundings that it is enabled to live till it produces offspring which may take its place when it ceases to exist. We have seen also that, of the whole annual increase only a very small fraction survives; and though the survival in individual cases may sometimes be due rather to accident than to any real superiority, yet we cannot doubt that, in the long run, those survive which are best fitted by their perfect organisation to escape the dangers that surround them. This "survival of the fittest" is what Darwin termed "natural selection," because it leads to the same results in nature as are produced by man's selection among domestic animals and cultivated plants. Its primary effect will, clearly, be to keep each species in the most perfect health and vigour, with every part of its organisation in full harmony with the conditions of its existence. It prevents any possible deterioration in the organic world, and produces that appearance of exuberant life and enjoyment, of health and beauty, that affords us so much pleasure, and which might lead a superficial observer to suppose that peace and quietude reigned throughout nature.
The Effect under changed Conditions.
But the very same process which, so long as conditions remain substantially the same, secures the continuance of each species of animal or plant in its full perfection, will usually, under changed conditions, bring about whatever change of structure or habits may be necessitated by them. The changed conditions to which we refer are such as we know have occurred throughout all geological time and in every part of the world. Land and water have been continually shifting their positions; some regions are undergoing subsidence with diminution of area, others elevation with extension of area; dry land has been converted into marshes, while marshes have been drained or have even been elevated into plateaux. Climate too has changed again and again, either through the elevation of mountains in high latitudes leading to the accumulation of snow and ice, or by a change in the direction of winds and ocean currents produced by the subsidence or elevation of lands which connected continents and divided oceans. Again, along with all these changes have come not less important changes in the distribution of species. Vegetation has been greatly modified by changes of climate and of altitude; while every union of lands before separated has led to extensive migrations of animals into new countries, disturbing the balance that before existed among its forms of life, leading to the extermination of some species and the increase of others.
When such physical changes as these have taken place, it is evident that many species must either become modified or cease to exist. When the vegetation has changed in character the herbivorous animals must become able to live on new and perhaps less nutritious food; while the change from a damp to a dry climate may necessitate migration at certain periods to escape destruction by drought. This will expose the species to new dangers, and require special modifications of structure to meet them. Greater swiftness, increased cunning, nocturnal habits, change of colour, or the power of climbing trees and living for a time on their foliage or fruit, may be the means adopted by different species to bring themselves into harmony with the new conditions; and by the continued survival of those individuals, only, which varied sufficiently in the right direction, the necessary modifications of structure or of function would be brought about, just as surely as man has been able to breed the greyhound to hunt by sight and the foxhound by scent, or has produced from the same wild plant such distinct forms as the cauliflower and the brussels sprouts.
We will now consider the special characteristics of the changes in species that are likely to be effected, and how far they agree with what we observe in nature.
Divergence of Character.
In species which have a wide range the struggle for existence will often cause some individuals or groups of individuals to adopt new habits in order to seize upon vacant places in nature where the struggle is less severe. Some, living among extensive marshes, may adopt a more aquatic mode of life; others, living where forests abound, may become more arboreal. In either case we cannot doubt that the changes of structure needed to adapt them to their new habits would soon be brought about, because we know that variations in all the external organs and all their separate parts are very abundant and are also considerable in amount. That such divergence of character has actually occurred we have some direct evidence. Mr. Darwin informs us that in the Catskill Mountains in the United States there are two varieties of wolves, one with a light greyhound-like form which pursues deer, the other more bulky with shorter legs, which more frequently attacks sheep.[1] Another good example is that of the insects in the island of Madeira, many of which have either lost their wings or have had them so much reduced as to be useless for flight, while the very same species on the continent of Europe possess fully developed wings. In other cases the wingless Madeira species are distinct from, but closely allied to, winged species of Europe. The explanation of this change is, that Madeira, like many oceanic islands in the temperate zone, is much exposed to sudden gales of wind, and as most of the fertile land is on the coast, insects which flew much would be very liable to be blown out to sea and lost. Year after year, therefore, those individuals which had shorter wings, or which used them least, were preserved; and thus, in time, terrestrial, wingless, or imperfectly winged races or species have been produced. That this is the true explanation of this singular fact is proved by much corroborative evidence. There are some few flower-frequenting insects in Madeira to whom wings are essential, and in these the wings are somewhat larger than in the same species on the mainland. We thus see that there is no general tendency to the abortion of wings in Madeira, but that it is simply a case of adaptation to new conditions. Those insects to whom wings were not absolutely essential escaped a serious danger by not using them, and the wings therefore became reduced or were completely lost. But when they were essential they were enlarged and strengthened, so that the insect could battle against the winds and save itself from destruction at sea. Many flying insects, not varying fast enough, would be destroyed before they could establish themselves, and thus we may explain the total absence from Madeira of several whole families of winged insects which must have had many opportunities of reaching the islands. Such are the large groups of the tiger-beetles (Cicindelidæ), the chafers (Melolonthidæ), the click-beetles (Elateridæ), and many others.
But the most curious and striking confirmation of this portion of Mr. Darwin's theory is afforded by the case of Kerguelen Island. This island was visited by the Transit of Venus expedition. It is one of the stormiest places on the globe, being subject to almost perpetual gales, while, there being no wood, it is almost entirely without shelter. The Rev. A. E. Eaton, an experienced entomologist, was naturalist to the expedition, and he assiduously collected the few insects that were to be found. All were incapable of flight, and most of them entirely without wings. They included a moth, several flies, and numerous beetles. As these insects could hardly have reached the islands in a wingless state, even if there were any other known land inhabited by them—which there is not—we must assume that, like the Madeiran insects, they were originally winged, and lost their power of flight because its possession was injurious to them.
It is no doubt due to the same cause that some butterflies on small and exposed islands have their wings reduced in size, as is strikingly the case with the small tortoise-shell butterfly (Vanessa urtica) inhabiting the Isle of Man, which is only about half the size of the same species in England or Ireland; and Mr. Wollaston notes that Vanessa callirhoe—a closely allied South European form of our red-admiral butterfly—is permanently smaller in the small and bare island of Porto Santo than in the larger and more wooded adjacent island of Madeira.
A very good example of comparatively recent divergence of character, in accordance with new conditions of life, is afforded by our red grouse. This bird, the Lagopus scoticus of naturalists, is entirely confined to the British Isles. It is, however, very closely allied to the willow grouse (Lagopus albus), a bird which ranges all over Europe, Northern Asia, and North America, but which, unlike our species, changes to white in winter. No difference in form or structure can be detected between the two birds, but as they differ so decidedly in colour—our species being usually rather darker in winter than in summer, while there are also slight differences in the call-note and in habits,—the two species are generally considered to be distinct. The differences, however, are so clearly adaptations to changed conditions that we can hardly doubt that, during the early part of the glacial period, when our islands were united to the continent, our grouse was identical with that of the rest of Europe. But when the cold passed away and our islands became permanently separated from the mainland, with a mild and equable climate and very little snow in winter, the change to white at that season became hurtful, rendering the birds more conspicuous instead of serving as a means of concealment. The colour was, therefore, gradually changed by the process of variation and natural selection; and as the birds obtained ample shelter among the heather which clothes so many of our moorlands, it became useful for them to assimilate with its brown and dusky stems and withered flowers rather than with the snow of the higher mountains. An interesting confirmation of this change having really occurred is afforded by the occasional occurrence in Scotland of birds with a considerable amount of white in the winter plumage. This is considered to be a case of reversion to the ancestral type, just as the slaty colours and banded wings of the wild rock-pigeon sometimes reappear in our fancy breeds of domestic pigeons.[2]
The principle of "divergence of character" pervades all nature from the lowest groups to the highest, as may be well seen in the class of birds. Among our native species we see it well marked in the different species of titmice, pipits, and chats. The great titmouse (Parus major) by its larger size and stronger bill is adapted to feed on larger insects, and is even said sometimes to kill small and weak birds. The smaller and weaker coal titmouse (Parus ater) has adopted a more vegetarian diet, eating seeds as well as insects, and feeding on the ground as well as among trees. The delicate little blue titmouse (Parus cœruleus), with its very small bill, feeds on the minutest insects and grubs which it extracts from crevices of bark and from the buds of fruit-trees. The marsh titmouse, again (Parus palustris), has received its name from the low and marshy localities it frequents; while the crested titmouse (Parus cristatus) is a northern bird frequenting especially pine forests, on the seeds of which trees it partially feeds. Then, again, our three common pipits—the tree-pipit (Anthus arboreus), the meadow-pipit (Anthus pratensis), and the rock-pipit or sea-lark (Anthus obscurus) have each occupied a distinct place in nature to which they have become specially adapted, as indicated by the different form and size of the hind toe and claw in each species. So, the stone-chat (Saxicola rubicola), the whin-chat (S. rubetra), and the wheat-ear (S. œnanthe) are more or less divergent forms of one type, with modifications in the shape of the wing, feet, and bill adapting them to slightly different modes of life. The whin-chat is the smallest, and frequents furzy commons, fields, and lowlands, feeding on worms, insects, small molluscs, and berries; the stone-chat is next in size, and is especially active and lively, frequenting heaths and uplands, and is a permanent resident with us, the two other species being migrants; while the larger and more conspicuous wheat-ear, besides feeding on grubs, beetles, etc., is able to capture flying insects on the wing, something after the manner of true flycatchers.
These examples sufficiently indicate how divergence of character has acted, and has led to the adaptation of numerous allied species, each to a more or less special mode of life, with the variety of food, of habits, and of enemies which must necessarily accompany such diversity. And when we extend our inquiries to higher groups we find the same indications of divergence and special adaptation, often to a still more marked extent. Thus we have the larger falcons, which prey upon birds, while some of the smaller species, like the hobby (Falco subbuteo), live largely on insects. The true falcons capture their prey in the air, while the hawks usually seize it on or near the ground, feeding on hares, rabbits, squirrels, grouse, pigeons, and poultry. Kites and buzzards, on the other hand, seize their prey upon the ground, and the former feed largely on reptiles and offal as well as on birds and quadrupeds. Others have adopted fish as their chief food, and the osprey snatches its prey from the water with as much facility as a gull or a petrel; while the South American caracaras (Polyborus) have adopted the habits of vultures and live altogether on carrion. In every great group there is the same divergence of habits. There are ground-pigeons, rock-pigeons, and wood-pigeons,—seed-eating pigeons and fruit-eating pigeons; there are carrion-eating, insect-eating, and fruit-eating crows. Even kingfishers are, some aquatic, some terrestrial in their habits; some live on fish, some on insects, some on reptiles. Lastly, among the primary divisions of birds we find a purely terrestrial group—the Ratitæ, including the ostriches, cassowaries, etc.; other great groups, including the ducks, cormorants, gulls, penguins, etc., are aquatic; while the bulk of the Passerine birds are aerial and arboreal. The same general facts can be detected in all other classes of animals. In the mammalia, for example, we have in the common rat a fish-eater and flesh-eater as well as a grain-eater, which has no doubt helped to give it the power of spreading over the world and driving away the native rats of other countries. Throughout the Rodent tribe we find everywhere aquatic, terrestrial, and arboreal forms. In the weasel and cat tribes some live more in trees, others on the ground; squirrels have diverged into terrestrial, arboreal, and flying species; and finally, in the bats we have a truly aerial, and in the whales a truly aquatic order of mammals. We thus see that, beginning with different varieties of the same species, we have allied species, genera, families, and orders, with similarly divergent habits, and adaptations to different modes of life, indicating some general principle in nature which has been operative in the development of the organic world. But in order to be thus operative it must be a generally useful principle, and Mr. Darwin has very clearly shown us in what this utility consists.
Divergence leads to a Maximum of Organic Forms in each Area.
Divergence of character has a double purpose and use. In the first place it enables a species which is being overcome by rivals, or is in process of extinction by enemies, to save itself by adopting new habits or by occupying vacant places in nature. This is the immediate and obvious effect of all the numerous examples of divergence of character which we have pointed out. But there is another and less obvious result, which is, that the greater the diversity in the organisms inhabiting a country or district the greater will be the total amount of life that can be supported there. Hence the continued action of the struggle for existence will tend to bring about more and more diversity in each area, which may be shown to be the case by several kinds of evidence. As an example, a piece of turf, three feet by four in size, was found by Mr. Darwin to contain twenty species of plants, and these twenty species belonged to eighteen genera and to eight orders, showing how greatly they differed from each other. Farmers find that a greater quantity of hay is obtained from ground sown with a variety of genera of grasses, clover, etc., than from similar land sown with one or two species only; and the same principle applies to rotation of crops, plants differing very widely from each other giving the best results. So, in small and uniform islands, and in small ponds of fresh water, the plants and insects, though few in number, are found to be wonderfully varied in character.
The same principle is seen in the naturalisation of plants and animals by man's agency in distant lands, for the species that thrive best and establish themselves permanently are not only very varied among themselves but differ greatly from the native inhabitants. Thus, in the Northern United States there are, according to Dr. Asa Gray, 260 naturalised flowering plants which belong to no less than 162 genera; and of these, 100 genera are not natives of the United States. So, in Australia, the rabbit, though totally unlike any native animal, has increased so much that it probably outnumbers in individuals all the native mammals of the country; and in New Zealand the rabbit and the pig have equally multiplied. Darwin remarks that this "advantage of diversification of structure in the inhabitants of the same region is, in fact, the same as that of the physiological division of labour in the organs of the same body. No physiologist doubts that a stomach adapted to digest vegetable matter alone, or flesh alone, draws more nutriment from these substances. So, in the general economy of any land, the more widely and perfectly the animals and plants are diversified for different habits of life, so will a greater number of individuals be capable of there supporting themselves."[3]
The most closely allied Species inhabit distinct Areas.
One of the curious results of the general action of this principle in nature is, that the most closely allied species—those whose differences though often real and important are hardly perceptible to any one but a naturalist—are usually not found in the same but in widely separated countries. Thus, the nearest allies to our European golden plover are found in North America and East Asia; the nearest ally of our European jay is found in Japan, although there are several other species of jays in Western Asia and North Africa; and though we have several species of titmice in England they are not very closely allied to each other. The form most akin to our blue tit is the azure tit of Central Asia (Parus azureus); the Parus ledouci of Algeria is very near our coal tit, and the Parus lugubris of South-Eastern Europe and Asia Minor is nearest to our marsh tit. So, our four species of wild pigeons—the ring-dove, stock-dove, rock-pigeon, and turtle-dove—are not closely allied to each other, but each of them belongs, according to some ornithologists, to a separate genus or subgenus, and has its nearest relatives in distant parts of Asia and Africa. In mammalia the same thing occurs. Each mountain region of Europe and Asia has usually its own species of wild sheep and goat, and sometimes of antelope and deer; so that in each region there is found the greatest diversity in this class of animals, while the closest allies inhabit quite distinct and often distant areas. In plants we find the same phenomenon prevalent. Distinct species of columbine are found in Central Europe (Aguilegia vulgaris), in Eastern Europe, and Siberia (A. glandulosa), in the Alps (A. Alpina), in the Pyrenees (A. pyrenaiea), in the Greek mountains (A. ottonis), and in Corsica (A. Bernardi), but rarely are two species found in the same area. So, each part of the world has its own peculiar forms of pines, firs, and cedars, but the closely allied species or varieties are in almost every case inhabitants of distinct areas. Examples are the deodar of the Himalayas, the cedar of Lebanon, and that of North Africa, all very closely allied but confined to distinct areas; and the numerous closely allied species of true pine (genus Pinus), which almost always inhabit different countries or occupy different stations. We will now consider some other modes in which natural selection will act, to adapt organisms to changed conditions.
Adaptation to Conditions at Various Periods of Life.
It is found, that, in domestic animals and cultivated plants, variations occurring at any one period of life reappear in the offspring at the same period, and can be perpetuated and increased by selection without modifying other parts of the organisation. Thus, variations in the caterpillar or the cocoon of the silkworm, in the eggs of poultry, and in the seeds or young shoots of many culinary vegetables, have been accumulated till those parts have become greatly modified and, for man's purposes, improved. Owing to this fact it is easy for organisms to become so modified as to avoid dangers that occur at any one period of life. Thus it is that so many seeds have become adapted to various modes of dissemination or protection. Some are winged, or have down or hairs attached to them, so as to enable them to be carried long distances in the air; others have curious hooks and prickles, which cause them to be attached firmly to the fur of mammals or the feathers of birds; while others are buried within sweet or juicy and brightly coloured fruits, which are seen and devoured by birds, the hard smooth seeds passing through their bodies in a fit state for germination. In the struggle for existence it must benefit a plant to have increased means of dispersing its seeds, and of thus having young plants produced in a greater variety of soils, aspects, and surroundings, with a greater chance of some of them escaping their numerous enemies and arriving at maturity. The various differences referred to would, therefore, be brought about by variation and survival of the fittest, just as surely as the length and quality of cotton on the seed of the cotton-plant have been increased by man's selection.
The larvæ of insects have thus been wonderfully modified in order to escape the numerous enemies to whose attacks they are exposed at this period of their existence. Their colours and markings have become marvellously adapted to conceal them among the foliage of the plant they live upon, and this colour often changes completely after the last moult, when the creature has to descend to the ground for its change to the pupa state, during which period a brown instead of a green colour is protective. Others have acquired curious attitudes and large ocelli, which cause them to resemble the head of some reptile, or they have curious horns or coloured ejectile processes which frighten away enemies; while a great number have acquired secretions which render them offensive to the taste of their enemies, and these are always adorned with very conspicuous markings or brilliant colours, which serve as a sign of inedibility and prevent their being needlessly attacked. This, however, is a portion of the very large subject of organic colour and marking, which will be fully discussed and illustrated in a separate chapter.
In this way every possible modification of an animal or plant, whether in colour, form, structure, or habits, which would be serviceable to it or to its progeny at any period of its existence, may be readily brought about. There are some curious organs which are used only once in a creature's life, but which are yet essential to its existence, and thus have very much the appearance of design by an intelligent designer. Such are, the great jaws possessed by some insects, used exclusively for opening the cocoon, and the hard tip to the beak of unhatched birds used for breaking the eggshell. The increase in thickness or hardness of the cocoons or the eggs being useful for protection against enemies or to avoid accidents, it is probable that the change has been very gradual, because it would be constantly checked by the necessity for a corresponding change in the young insects or birds enabling them to overcome the additional obstacle of a tougher cocoon or a harder eggshell. As we have seen, however, that every part of the organism appears to be varying independently, at the same time, though to different amounts, there seems no reason to believe that the necessity for two or more coincident variations would prevent the required change from taking place.
The Continued Existence of Low Forms of Life.
Since species are continually undergoing modifications giving them some superiority over other species or enabling them to occupy fresh places in nature, it may be asked—Why do any low forms continue to exist? Why have they not long since been improved and developed into higher forms? The answer, probably, is, that these low forms occupy places in nature which cannot be filled by higher forms, and that they have few or no competitors; they therefore continue to exist. Thus, earthworms are adapted to their mode of life better than they would be if more highly organised. So, in the ocean, the minute foraminifera and infusoria, and the larger sponges and corals, occupy places which more highly developed creatures could not fill. They form, as it were, the base of the great structure of animal life, on which the next higher forms rest; and though in the course of ages they may undergo some changes, and diversification of form and structure, in accordance with changed conditions, their essential nature has probably remained the same from the very dawn of life on the earth. The low aquatic diatomaceæ and confervæ, together with the lowest fungi and lichens, occupy a similar position in the vegetable kingdom, filling places in nature which would be left vacant if only highly organised plants existed. There is, therefore, no motive power to destroy or seriously to modify them; and they have thus probably persisted, under slightly varying forms, through all geological time.
Extinction of Lower Types among the Higher Animals.
So soon; however, as we approach the higher and more fully developed groups, we see indications of the often repeated extinction of lower by higher forms. This is shown by the great gaps that separate the mammalia, birds, reptiles, and fishes from each other; while the lowest forms of each are always few in number and confined to limited areas. Such are the lowest mammals—the echidna and ornithorhynchus of Australia; the lowest birds—the apteryx of New Zealand and the cassowaries of the New Guinea region; while the lowest fish—the amphioxus or lancelet, is completely isolated, and has apparently survived only by its habit of burrowing in the sand. The great distinctness of the carnivora, ruminants, rodents, whales, bats, and other orders of mammalia; of the accipitres, pigeons, and parrots, among birds; and of the beetles, bees, flies, and moths, among insects, all indicate an enormous amount of extinction among the comparatively low forms by which, on any theory of evolution, these higher and more specialised groups must have been preceded.
Circumstances favourable to the Origin of New Species by Natural Selection.
We have already seen that, when there is no change in the physical or organic conditions of a country, the effect of natural selection is to keep all the species inhabiting it in a state of perfect health and full development, and to preserve the balance that already exists between the different groups of organisms. But, whenever the physical or organic conditions change, to however small an extent, some corresponding change will be produced in the flora and fauna, since, considering the severe struggle for existence and the complex relations of the various organisms, it is hardly possible that the change should not be beneficial to some species and hurtful to others. The most common effect, therefore, will be that some species will increase and others will diminish; and in cases where a species was already small in numbers a further diminution might lead to extinction. This would afford room for the increase of other species, and thus a considerable readjustment of the proportions of the several species might take place. When, however, the change was of a more important character, directly affecting the existence of many species so as to render it difficult for them to maintain themselves without some considerable change in structure or habits, that change would, in some cases, be brought about by variation and natural selection, and thus new varieties or new species might be formed. We have to consider, then, which are the species that would be most likely to be so modified, while others, not becoming modified, would succumb to the changed conditions and become extinct.
The most important condition of all is, undoubtedly, that variations should occur of sufficient amount, of a sufficiently diverse character, and in a large number of individuals, so as to afford ample materials for natural selection to act upon; and this, we have seen, does occur in most, if not in all, large, wide-ranging, and dominant species. From some of these, therefore, the new species adapted to the changed conditions would usually be derived; and this would especially be the case when the change of conditions was rather rapid, and when a correspondingly rapid modification could alone save some species from extinction. But when the change was very gradual, then even less abundant and less widely distributed species might become modified into new forms, more especially if the extinction of many of the rarer species left vacant places in the economy of nature.
Probable Origin of the Dippers.
An excellent example of how a limited group of species has been able to maintain itself by adaptation to one of these "vacant places" in nature, is afforded by the curious little birds called dippers or water-ouzels, forming the genus Cinclus and the family Cinclidæ of naturalists. These birds are something like small thrushes, with very short wings and tail, and very dense plumage. They frequent, exclusively, mountain torrents in the northern hemisphere, and obtain their food entirely in the water, consisting, as it does, of water-beetles, caddis-worms and other insect-larvæ, as well as numerous small freshwater shells. These birds, although not far removed in structure from thrushes and wrens, have the extraordinary power of flying under water; for such, according to the best observers, is their process of diving in search of their prey, their dense and somewhat fibrous plumage retaining so much air that the water is prevented from touching their bodies or even from wetting their feathers to any great extent. Their powerful feet and long curved claws enable them to hold on to stones at the bottom, and thus to retain their position while picking up insects, shells, etc. As they frequent chiefly the most rapid and boisterous torrents, among rocks, waterfalls, and huge boulders, the water is never frozen over, and they are thus able to live during the severest winters. Only a very few species of dipper are known, all those of the old world being so closely allied to our British bird that some ornithologists consider them to be merely local races of one species; while in North America and the northern Andes there are two other species.
Here then we have a bird, which, in its whole structure, shows a close affinity to the smaller typical perching birds, but which has departed from all its allies in its habits and mode of life, and has secured for itself a place in nature where it has few competitors and few enemies. We may well suppose, that, at some remote period, a bird which was perhaps the common and more generalised ancestor of most of our thrushes, warblers, wrens, etc., had spread widely over the great northern continent, and had given rise to numerous varieties adapted to special conditions of life. Among these some took to feeding on the borders of clear streams, picking out such larvae and molluscs as they could reach in shallow water. When food became scarce they would attempt to pick them out of deeper and deeper water, and while doing this in cold weather many would become frozen and starved. But any which possessed denser and more hairy plumage than usual, which was able to keep out the water, would survive; and thus a race would be formed which would depend more and more on this kind of food. Then, following up the frozen streams into the mountains, they would be able to live there during the winter; and as such places afforded them much protection from enemies and ample shelter for their nests and young, further adaptations would occur, till the wonderful power of diving and flying under water was acquired by a true land-bird.
That such habits might be acquired under stress of need is rendered highly probable by the facts stated by the well-known American naturalist, Dr. Abbott. He says that "the water-thrushes (Seiurus sp.) all wade in water, and often, seeing minute mollusca on the bottom of the stream, plunge both head and neck beneath the surface, so that often, for several seconds, a large part of the body is submerged. Now these birds still have the plumage pervious to water, and so are liable to be drenched and sodden; but they have also the faculty of giving these drenched feathers such a good shaking that flight is practicable a moment after leaving the water. Certainly the water-thrushes (Seiurus ludovicianus, S. auricapillus, and S. noveboracensis) have taken many preliminary steps to becoming as aquatic as the dipper; and the winter-wren, and even the Maryland yellow-throat are not far behind."[4]
Another curious example of the way in which species have been modified to occupy new places in nature, is afforded by the various animals which inhabit the water-vessels formed by the leaves of many epiphytal species of Bromelia. Fritz Müller has described a caddis-fly larva which lives among these leaves, and which has been modified in the pupa state in accordance with its surroundings. The pupæ of caddis-flies inhabiting streams have fringes of hair on the tarsi to enable them to reach the surface on leaving their cases. But in the species inhabiting bromelia leaves there is no need for swimming, and accordingly we find the tarsi entirely bare. In the same plants are found curious little Entomostraca, very abundant there but found nowhere else. These form a new genus, but are most nearly allied to Cythere, a marine type. It is believed that the transmission of this species from one tree to another must be effected by the young crustacea, which are very minute, clinging to beetles, many of which, both terrestrial and aquatic, also inhabit the bromelia leaves; and as some water-beetles are known to frequent the sea, it is perhaps by these means that the first emigrants established themselves in this strange new abode. Bromeliæ are often very abundant on trees growing on the water's edge, and this would facilitate the transition from a marine to an arboreal habitat. Fritz Müller has also found, among the bromelia leaves, a small frog bearing its eggs on its back, and having some other peculiarities of structure. Several beautiful little aquatic plants of the genus Utricularia or bladder-wort also inhabit bromelia leaves; and these send runners out to neighbouring plants and thus spread themselves with great rapidity.
The Importance of Isolation.
Isolation is no doubt an important aid to natural selection, as shown by the fact that islands so often present a number of peculiar species; and the same thing is seen on the two sides of a great mountain range or on opposite coasts of a continent. The importance of isolation is twofold. In the first place, it leads to a body of individuals of each species being limited in their range and thus subjected to uniform conditions for long spaces of time. Both the direct action of the environment and the natural selection of such varieties only as are suited to the conditions, will, therefore, be able to produce their full effect. In the second place, the process of change will not be interfered with by intercrossing with other individuals which are becoming adapted to somewhat different conditions in an adjacent area. But this question of the swamping effects of intercrossing will be considered in another chapter.
Mr. Darwin was of opinion that, on the whole, the largeness of the area occupied by a species was of more importance than isolation, as a factor in the production of new species, and in this I quite agree with him. It must, too, be remembered, that isolation will often be produced in a continuous area whenever a species becomes modified in accordance with varied conditions or diverging habits. For example, a wide-ranging species may in the northern and colder part of its area become modified in one direction, and in the southern part in another direction; and though for a long time an intermediate form may continue to exist in the intervening area, this will be likely soon to die out, both because its numbers will be small, and it will be more or less pressed upon in varying seasons by the modified varieties, each better able to endure extremes of climate. So, when one portion of a terrestrial species takes to a more arboreal or to a more aquatic mode of life, the change of habit itself leads to the isolation of each portion. Again, as will be more fully explained in a future chapter, any difference of habits or of haunts usually leads to some modification of colour or marking, as a means of concealment from enemies; and there is reason to believe that this difference will be intensified by natural selection as a means of identification and recognition by members of the same variety or incipient species. It has also been observed that each differently coloured variety of wild animals, or of domesticated animals which have run wild, keep together, and refuse to pair with individuals of the other colours; and this must of itself act to keep the races separate as completely as physical isolation.
On the Advance of Organisation by Natural Selection.
As natural selection acts solely by the preservation of useful variations, or those which are beneficial to the organism under the conditions to which it is exposed, the result must necessarily be that each species or group tends to become more and more improved in relation to its conditions. Hence we should expect that the larger groups in each class of animals and plants—those which have persisted and have been abundant throughout geological ages—would, almost necessarily, have arrived at a high degree of organisation, both physical and mental. Illustrations of this are to be seen everywhere. Among mammalia we have the carnivora, which from Eocene times have been becoming more and more specialised, till they have culminated in the cat and dog tribes, which have reached a degree of perfection both in structure and intelligence fully equal to that of any other animals. In another line of development, the herbivora have been specialised for living solely on vegetable food till they have culminated in the sheep, the cattle, the deer, and the antelopes. The horse tribe, commencing with an early four-toed ancestor in the Eocene age, has increased in size and in perfect adaptation of feet and teeth to a life on open plains, and has reached its highest perfection in the horse, the ass, and the zebra. In birds, also, we see an advance from the imperfect tooth-billed and reptile-tailed birds of the secondary epoch, to the wonderfully developed falcons, crows, and swallows of our time. So, the ferns, lycopods, conifers, and monocotyledons of the palæozoic and mesozoic rocks, have developed into the marvellous wealth of forms of the higher dicotyledons that now adorn the earth.
But this remarkable advance in the higher and larger groups does not imply any universal law of progress in organisation, because we have at the same time numerous examples (as has been already pointed out) of the persistence of lowly organised forms, and also of absolute degradation or degeneration. Serpents, for example, have been developed from some lizard-like type which has lost its limbs; and though this loss has enabled them to occupy fresh places in nature and to increase and flourish to a marvellous extent, yet it must be considered to be a retrogression rather than an advance in organisation. The same remark will apply to the whale tribe among mammals; to the blind amphibia and insects of the great caverns; and among plants to the numerous cases in which flowers, once specially adapted to be fertilised by insects, have lost their gay corollas and their special adaptations, and have become degraded into wind-fertilised forms. Such are our plantains, our meadow burnet, and even, as some botanists maintain, our rushes, sedges, and grasses. The causes which have led to this degeneration will be discussed in a future chapter; but the facts are undisputed, and they show us that although variation and the struggle for existence may lead, on the whole, to a continued advance of organisation; yet they also lead in many cases to a retrogression, when such retrogression may aid in the preservation of any form under new conditions. They also lead to the persistence, with slight modifications, of numerous lowly organised forms which are suited to places which higher forms could not fully occupy, or to conditions under which they could not exist. Such are the ocean depths, the soil of the earth, the mud of rivers, deep caverns, subterranean waters, etc.; and it is in such places as these, as well as in some oceanic islands which competing higher forms have not been able to reach, that we find many curious relics of an earlier world, which, in the free air and sunlight and in the great continents, have long since been driven out or exterminated by higher types.
Summary of the first Five Chapters.
We have now passed in review, in more or less detail, the main facts on which the theory of "the origin of species by means of natural selection" is founded. In future chapters we shall have to deal mainly with the application of the theory to explain the varied and complex phenomena presented by the organic world; and, also, to discuss some of the theories put forth by modern writers, either as being more fundamental than that of Darwin or as supplementary to it. Before doing this, however, it will be well briefly to summarise the facts and arguments already set forth, because it is only by a clear comprehension of these that the full importance of the theory can be appreciated and its further applications understood.
The theory itself is exceedingly simple, and the facts on which it rests—though excessively numerous individually, and coextensive with the entire organic world—yet come under a few simple and easily understood classes. These facts are,—first, the enormous powers of increase in geometrical progression possessed by all organisms, and the inevitable struggle for existence among them; and, in the second place, the occurrence of much individual variation combined with the hereditary transmission of such variations. From these two great classes of facts, which are universal and indisputable, there necessarily arises, as Darwin termed it, the "preservation of favoured races in the struggle for life," the continuous action of which, under the ever-changing conditions both of the inorganic and organic universe, necessarily leads to the formation or development of new species.
But, although this general statement is complete and indisputable, yet to see its applications under all the complex conditions that actually occur in nature, it is necessary always to bear in mind the tremendous power and universality of the agencies at work. We must never for an instant lose sight of the fact of the enormously rapid increase of all organisms, which has been illustrated by actual cases, given in our second chapter, no less than by calculations of the results of unchecked increase for a few years. Then, never forgetting that the animal and plant population of any country is, on the whole, stationary, we must be always trying to realise the ever-recurring destruction of the enormous annual increase, and asking ourselves what determines, in each individual case, the death of the many, the survival of the few. We must think over all the causes of destruction to each organism,—to the seed, the young shoot, the growing plant, the full-grown tree, or shrub, or herb, and again the fruit and seed; and among animals, to the egg or new-born young, to the youthful, and to the adults. Then, we must always bear in mind that what goes on in the case of the individual or family group we may observe or think of, goes on also among the millions and scores of millions of individuals which are comprised in almost every species; and must get rid of the idea that chance determines which shall live and which die. For, although in many individual cases death may be due to chance rather than to any inferiority in those which die first, yet we cannot possibly believe that this can be the case on the large scale on which nature works. A plant, for instance, cannot be increased unless there are suitable vacant places its seeds can grow in, or stations where it can overcome other less vigorous and healthy plants. The seeds of all plants, by their varied modes of dispersal, may be said to be seeking out such places in which to grow; and we cannot doubt that, in the long run, those individuals whose seeds are the most numerous, have the greatest powers of dispersal, and the greatest vigour of growth, will leave more descendants than the individuals of the same species which are inferior in all these respects, although now and then some seed of an inferior individual may chance to be carried to a spot where it can grow and survive. The same rule will apply to every period of life and to every danger to which plants or animals are exposed. The best organised, or the most healthy, or the most active, or the best protected, or the most intelligent, will inevitably, in the long run, gain an advantage over those which are inferior in these qualities; that is, the fittest will survive, the fittest being, in each particular case, those which are superior in the special qualities on which safety depends. At one period of life, or to escape one kind of danger, concealment may be necessary; at another time, to escape another danger, swiftness; at another, intelligence or cunning; at another, the power to endure rain or cold or hunger; and those which possess all these faculties in the fullest perfection will generally survive.
Having fully grasped these facts in all their fulness and in their endless and complex results, we have next to consider the phenomena of variation, discussed in the third and fourth chapters; and it is here that perhaps the greatest difficulty will be felt in appreciating the full importance of the evidence as set forth. It has been so generally the practice to speak of variation as something exceptional and comparatively rare—as an abnormal deviation from the uniformity and stability of the characters of a species—and so few even among naturalists have ever compared, accurately, considerable numbers of individuals, that the conception of variability as a general characteristic of all dominant and widespread species, large in its amount and affecting, not a few, but considerable masses of the individuals which make up the species, will be to many entirely new. Equally important is the fact that the variability extends to every organ and every structure, external and internal; while perhaps most important of all is the independent variability of these several parts, each one varying without any constant or even usual dependence on, or correlation with, other parts. No doubt there is some such correlation in the differences that exist between species and species—more developed wings usually accompanying smaller feet and vice versâ—but this is, generally, a useful adaptation which has been brought about by natural selection, and does not apply to the individual variability which occurs within the species.
It is because these facts of variation are so important and so little understood, that they have been discussed in what will seem to some readers wearisome and unnecessary detail. Many naturalists, however, will hold that even more evidence is required; and more, to almost any amount, could easily have been given. The character and variety of that already adduced will, however, I trust, convince most readers that the facts are as stated; while they have been drawn from a sufficiently wide area to indicate a general principle throughout nature.
If, now, we fully realise these facts of variation, along with those of rapid multiplication and the struggle for existence, most of the difficulties in the way of comprehending how species have originated through natural selection will disappear. For whenever, through changes of climate, or of altitude, or of the nature of the soil, or of the area of the country, any species are exposed to new dangers, and have to maintain themselves and provide for the safety of their offspring under new and more arduous conditions, then, in the variability of all parts, organs, and structures, no less than of habits and intelligence, we have the means of producing modifications which will certainly bring the species into harmony with its new conditions. And if we remember that all such physical changes are slow and gradual in their operation, we shall see that the amount of variation which we know occurs in every new generation will be quite sufficient to enable modification and adaptation to go on at the same rate. Mr. Darwin was rather inclined to exaggerate the necessary slowness of the action of natural selection; but with the knowledge we now possess of the great amount and range of individual variation, there seems no difficulty in an amount of change, quite equivalent to that which usually distinguishes allied species, sometimes taking place in less than a century, should any rapid change of conditions necessitate an equally rapid adaptation. This may often have occurred, either to immigrants into a new land, or to residents whose country has been cut off by subsidence from a larger and more varied area over which they had formerly roamed. When no change of conditions occurs, species may remain unchanged for very long periods, and thus produce that appearance of stability of species which is even now often adduced as an argument against evolution by natural selection, but which is really quite in harmony with it.
On the principles, and by the light of the facts, now briefly summarised, we have been able, in the present chapter, to indicate how natural selection acts, how divergence of character is set up, how adaptation to conditions at various periods of life has been effected, how it is that low forms of life continue to exist, what kind of circumstances are most favourable to the formation of new species, and, lastly, to what extent the advance of organisation to higher types is produced by natural selection. We will now pass on to consider some of the more important objections and difficulties which have been advanced by eminent naturalists.