On the Origin of Species (sixth edition) 6 – Darwin · Verses Across
Literature · Animals
On the Origin of Species (sixth edition) 6Natural Selection; Or the Survival of the Fittest (part 2)
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Literature · Animals
On the Origin of Species (sixth edition) 6
Natural Selection; Or the Survival of the Fittest (part 2)
DarwinEnglish
In accordance with this view, we can, perhaps, understand some facts which will be again alluded to in our chapter on Geographical Distribution; for instance, the fact of the productions of the smaller continent of Australia now yielding before those of the larger Europæo-Asiatic area. Thus, also, it is that continental productions have everywhere become so largely naturalised on islands. On a small island, the race for life will have been less severe, and there will have been less modification and less extermination. Hence, we can understand how it is that the flora of Madeira, according to Oswald Heer, resembles to a certain extent the extinct tertiary flora of Europe. All fresh water basins, taken together, make a small area compared with that of the sea or of the land. Consequently, the competition between fresh water productions will have been less severe than elsewhere; new forms will have been more slowly produced, and old forms more slowly exterminated. And it is in fresh water basins that we find seven genera of Ganoid fishes, remnants of a once preponderant order: and in fresh water we find some of the most anomalous forms now known in the world, as the Ornithorhynchus and Lepidosiren, which, like fossils, connect to a certain extent orders at present widely separated in the natural scale. These anomalous forms may be called living fossils; they have endured to the present day, from having inhabited a confined area, and from having been exposed to less varied, and therefore less severe, competition.
To sum up, as far as the extreme intricacy of the subject permits, the circumstances favourable and unfavourable for the production of new species through natural selection. I conclude that for terrestrial productions a large continental area, which has undergone many oscillations of level, will have been the most favourable for the production of many new forms of life, fitted to endure for a long time and to spread widely. While the area existed as a continent the inhabitants will have been numerous in individuals and kinds, and will have been subjected to severe competition. When converted by subsidence into large separate islands there will still have existed many individuals of the same species on each island: intercrossing on the confines of the range of each new species will have been checked: after physical changes of any kind immigration will have been prevented, so that new places in the polity of each island will have had to be filled up by the modification of the old inhabitants; and time will have been allowed for the varieties in each to become well modified and perfected. When, by renewed elevation, the islands were reconverted into a continental area, there will again have been very severe competition; the most favoured or improved varieties will have been enabled to spread; there will have been much extinction of the less improved forms, and the relative proportional numbers of the various inhabitants of the reunited continent will again have been changed; and again there will have been a fair field for natural selection to improve still further the inhabitants, and thus to produce new species.
That natural selection generally act with extreme slowness I fully admit. It can act only when there are places in the natural polity of a district which can be better occupied by the modification of some of its existing inhabitants. The occurrence of such places will often depend on physical changes, which generally take place very slowly, and on the immigration of better adapted forms being prevented. As some few of the old inhabitants become modified the mutual relations of others will often be disturbed; and this will create new places, ready to be filled up by better adapted forms; but all this will take place very slowly. Although all the individuals of the same species differ in some slight degree from each other, it would often be long before differences of the right nature in various parts of the organisation might occur. The result would often be greatly retarded by free intercrossing. Many will exclaim that these several causes are amply sufficient to neutralise the power of natural selection. I do not believe so. But I do believe that natural selection will generally act very slowly, only at long intervals of time, and only on a few of the inhabitants of the same region. I further believe that these slow, intermittent results accord well with what geology tells us of the rate and manner at which the inhabitants of the world have changed.
Slow though the process of selection may be, if feeble man can do much by artificial selection, I can see no limit to the amount of change, to the beauty and complexity of the coadaptations between all organic beings, one with another and with their physical conditions of life, which may have been effected in the long course of time through nature’s power of selection, that is by the survival of the fittest.
Extinction caused by Natural Selection.
This subject will be more fully discussed in our chapter on Geology; but it must here be alluded to from being intimately connected with natural selection. Natural selection acts solely through the preservation of variations in some way advantageous, which consequently endure. Owing to the high geometrical rate of increase of all organic beings, each area is already fully stocked with inhabitants, and it follows from this, that as the favoured forms increase in number, so, generally, will the less favoured decrease and become rare. Rarity, as geology tells us, is the precursor to extinction. We can see that any form which is represented by few individuals will run a good chance of utter extinction, during great fluctuations in the nature or the seasons, or from a temporary increase in the number of its enemies. But we may go further than this; for as new forms are produced, unless we admit that specific forms can go on indefinitely increasing in number, many old forms must become extinct. That the number of specific forms has not indefinitely increased, geology plainly tells us; and we shall presently attempt to show why it is that the number of species throughout the world has not become immeasurably great.
We have seen that the species which are most numerous in individuals have the best chance of producing favourable variations within any given period. We have evidence of this, in the facts stated in the second chapter, showing that it is the common and diffused or dominant species which offer the greatest number of recorded varieties. Hence, rare species will be less quickly modified or improved within any given period; they will consequently be beaten in the race for life by the modified and improved descendants of the commoner species.
From these several considerations I think it inevitably follows, that as new species in the course of time are formed through natural selection, others will become rarer and rarer, and finally extinct. The forms which stand in closest competition with those undergoing modification and improvement, will naturally suffer most. And we have seen in the chapter on the Struggle for Existence that it is the most closely-allied forms,—varieties of the same species, and species of the same genus or related genera,—which, from having nearly the same structure, constitution and habits, generally come into the severest competition with each other. Consequently, each new variety or species, during the progress of its formation, will generally press hardest on its nearest kindred, and tend to exterminate them. We see the same process of extermination among our domesticated productions, through the selection of improved forms by man. Many curious instances could be given showing how quickly new breeds of cattle, sheep and other animals, and varieties of flowers, take the place of older and inferior kinds. In Yorkshire, it is historically known that the ancient black cattle were displaced by the long-horns, and that these “were swept away by the short-horns” (I quote the words of an agricultural writer) “as if by some murderous pestilence.”
Divergence of Character.
The principle, which I have designated by this term, is of high importance, and explains, as I believe, several important facts. In the first place, varieties, even strongly-marked ones, though having somewhat of the character of species—as is shown by the hopeless doubts in many cases how to rank them—yet certainly differ far less from each other than do good and distinct species. Nevertheless according to my view, varieties are species in the process of formation, or are, as I have called them, incipient species. How, then, does the lesser difference between varieties become augmented into the greater difference between species? That this does habitually happen, we must infer from most of the innumerable species throughout nature presenting well-marked differences; whereas varieties, the supposed prototypes and parents of future well-marked species, present slight and ill-defined differences. Mere chance, as we may call it, might cause one variety to differ in some character from its parents, and the offspring of this variety again to differ from its parent in the very same character and in a greater degree; but this alone would never account for so habitual and large a degree of difference as that between the species of the same genus.
As has always been my practice, I have sought light on this head from our domestic productions. We shall here find something analogous. It will be admitted that the production of races so different as short-horn and Hereford cattle, race and cart horses, the several breeds of pigeons, &c., could never have been effected by the mere chance accumulation of similar variations during many successive generations. In practice, a fancier is, for instance, struck by a pigeon having a slightly shorter beak; another fancier is struck by a pigeon having a rather longer beak; and on the acknowledged principle that “fanciers do not and will not admire a medium standard, but like extremes,” they both go on (as has actually occurred with the sub-breeds of the tumbler-pigeon) choosing and breeding from birds with longer and longer beaks, or with shorter and shorter beaks. Again, we may suppose that at an early period of history, the men of one nation or district required swifter horses, while those of another required stronger and bulkier horses. The early differences would be very slight; but, in the course of time, from the continued selection of swifter horses in the one case, and of stronger ones in the other, the differences would become greater, and would be noted as forming two sub-breeds. Ultimately after the lapse of centuries, these sub-breeds would become converted into two well-established and distinct breeds. As the differences became greater, the inferior animals with intermediate characters, being neither very swift nor very strong, would not have been used for breeding, and will thus have tended to disappear. Here, then, we see in man’s productions the action of what may be called the principle of divergence, causing differences, at first barely appreciable, steadily to increase, and the breeds to diverge in character, both from each other and from their common parent.
But how, it may be asked, can any analogous principle apply in nature? I believe it can and does apply most efficiently (though it was a long time before I saw how), from the simple circumstance that the more diversified the descendants from any one species become in structure, constitution, and habits, by so much will they be better enabled to seize on many and widely diversified places in the polity of nature, and so be enabled to increase in numbers.
We can clearly discern this in the case of animals with simple habits. Take the case of a carnivorous quadruped, of which the number that can be supported in any country has long ago arrived at its full average. If its natural power of increase be allowed to act, it can succeed in increasing (the country not undergoing any change in conditions) only by its varying descendants seizing on places at present occupied by other animals: some of them, for instance, being enabled to feed on new kinds of prey, either dead or alive; some inhabiting new stations, climbing trees, frequenting water, and some perhaps becoming less carnivorous. The more diversified in habits and structure the descendants of our carnivorous animals become, the more places they will be enabled to occupy. What applies to one animal will apply throughout all time to all animals—that is, if they vary—for otherwise natural selection can effect nothing. So it will be with plants. It has been experimentally proved, that if a plot of ground be sown with one species of grass, and a similar plot be sown with several distinct genera of grasses, a greater number of plants and a greater weight of dry herbage can be raised in the latter than in the former case. The same has been found to hold good when one variety and several mixed varieties of wheat have been sown on equal spaces of ground. Hence, if any one species of grass were to go on varying, and the varieties were continually selected which differed from each other in the same manner, though in a very slight degree, as do the distinct species and genera of grasses, a greater number of individual plants of this species, including its modified descendants, would succeed in living on the same piece of ground. And we know that each species and each variety of grass is annually sowing almost countless seeds; and is thus striving, as it may be said, to the utmost to increase in number. Consequently, in the course of many thousand generations, the most distinct varieties of any one species of grass would have the best chance of succeeding and of increasing in numbers, and thus of supplanting the less distinct varieties; and varieties, when rendered very distinct from each other, take the rank of species.
The truth of the principle that the greatest amount of life can be supported by great diversification of structure, is seen under many natural circumstances. In an extremely small area, especially if freely open to immigration, and where the contest between individual and individual must be very severe, we always find great diversity in its inhabitants. For instance, I found that a piece of turf, three feet by four in size, which had been exposed for many years to exactly the same conditions, supported twenty species of plants, and these belonged to eighteen genera and to eight orders, which shows how much these plants differed from each other. So it is with the plants and insects on small and uniform islets: also in small ponds of fresh water. Farmers find that they can raise more food by a rotation of plants belonging to the most different orders: nature follows what may be called a simultaneous rotation. Most of the animals and plants which live close round any small piece of ground, could live on it (supposing its nature not to be in any way peculiar), and may be said to be striving to the utmost to live there; but, it is seen, that where they come into the closest competition, the advantages of diversification of structure, with the accompanying differences of habit and constitution, determine that the inhabitants, which thus jostle each other most closely, shall, as a general rule, belong to what we call different genera and orders.
The same principle is seen in the naturalisation of plants through man’s agency in foreign lands. It might have been expected that the plants which would succeed in becoming naturalised in any land would generally have been closely allied to the indigenes; for these are commonly looked at as specially created and adapted for their own country. It might also, perhaps, have been expected that naturalised plants would have belonged to a few groups more especially adapted to certain stations in their new homes. But the case is very different; and Alph. de Candolle has well remarked, in his great and admirable work, that floras gain by naturalisation, proportionally with the number of the native genera and species, far more in new genera than in new species. To give a single instance: in the last edition of Dr. Asa Gray’s “Manual of the Flora of the Northern United States,” 260 naturalised plants are enumerated, and these belong to 162 genera. We thus see that these naturalised plants are of a highly diversified nature. They differ, moreover, to a large extent, from the indigenes, for out of the 162 naturalised genera, no less than 100 genera are not there indigenous, and thus a large proportional addition is made to the genera now living in the United States.
By considering the nature of the plants or animals which have in any country struggled successfully with the indigenes, and have there become naturalised, we may gain some crude idea in what manner some of the natives would have had to be modified in order to gain an advantage over their compatriots; and we may at least infer that diversification of structure, amounting to new generic differences, would be profitable to them.
The 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 individual body—a subject so well elucidated by Milne Edwards. No physiologist doubts that a stomach by being adapted to digest vegetable matter alone, or flesh alone, draws most 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. A set of animals, with their organisation but little diversified, could hardly compete with a set more perfectly diversified in structure. It may be doubted, for instance, whether the Australian marsupials, which are divided into groups differing but little from each other, and feebly representing, as Mr. Waterhouse and others have remarked, our carnivorous, ruminant, and rodent mammals, could successfully compete with these well-developed orders. In the Australian mammals, we see the process of diversification in an early and incomplete stage of development.
The Probable Effects of the Action of Natural Selection through Divergence of Character and Extinction, on the Descendants of a Common Ancestor.
After the foregoing discussion, which has been much compressed, we may assume that the modified descendants of any one species will succeed so much the better as they become more diversified in structure, and are thus enabled to encroach on places occupied by other beings. Now let us see how this principle of benefit being derived from divergence of character, combined with the principles of natural selection and of extinction, tends to act.
The accompanying diagram will aid us in understanding this rather perplexing subject. Let A to L represent the species of a genus large in its own country; these species are supposed to resemble each other in unequal degrees, as is so generally the case in nature, and as is represented in the diagram by the letters standing at unequal distances. I have said a large genus, because as we saw in the second chapter, on an average more species vary in large genera than in small genera; and the varying species of the large genera present a greater number of varieties. We have, also, seen that the species, which are the commonest and most widely-diffused, vary more than do the rare and restricted species. Let (A) be a common, widely-diffused, and varying species, belonging to a genus large in its own country. The branching and diverging dotted lines of unequal lengths proceeding from (A), may represent its varying offspring. The variations are supposed to be extremely slight, but of the most diversified nature; they are not supposed all to appear simultaneously, but often after long intervals of time; nor are they all supposed to endure for equal periods. Only those variations which are in some way profitable will be preserved or naturally selected. And here the importance of the principle of benefit derived from divergence of character comes in; for this will generally lead to the most different or divergent variations (represented by the outer dotted lines) being preserved and accumulated by natural selection. When a dotted line reaches one of the horizontal lines, and is there marked by a small numbered letter, a sufficient amount of variation is supposed to have been accumulated to form it into a fairly well-marked variety, such as would be thought worthy of record in a systematic work.
The intervals between the horizontal lines in the diagram, may represent each a thousand or more generations. After a thousand generations, species (A) is supposed to have produced two fairly well-marked varieties, namely a1 and m1. These two varieties will generally still be exposed to the same conditions which made their parents variable, and the tendency to variability is in itself hereditary; consequently they will likewise tend to vary, and commonly in nearly the same manner as did their parents. Moreover, these two varieties, being only slightly modified forms, will tend to inherit those advantages which made their parent (A) more numerous than most of the other inhabitants of the same country; they will also partake of those more general advantages which made the genus to which the parent-species belonged, a large genus in its own country. And all these circumstances are favourable to the production of new varieties.
If, then, these two varieties be variable, the most divergent of their variations will generally be preserved during the next thousand generations. And after this interval, variety a1 is supposed in the diagram to have produced variety a2, which will, owing to the principle of divergence, differ more from (A) than did variety a1. Variety m1 is supposed to have produced two varieties, namely m2 and s2, differing from each other, and more considerably from their common parent (A). We may continue the process by similar steps for any length of time; some of the varieties, after each thousand generations, producing only a single variety, but in a more and more modified condition, some producing two or three varieties, and some failing to produce any. Thus the varieties or modified descendants of the common parent (A), will generally go on increasing in number and diverging in character. In the diagram the process is represented up to the ten-thousandth generation, and under a condensed and simplified form up to the fourteen-thousandth generation.
But I must here remark that I do not suppose that the process ever goes on so regularly as is represented in the diagram, though in itself made somewhat irregular, nor that it goes on continuously; it is far more probable that each form remains for long periods unaltered, and then again undergoes modification. Nor do I suppose that the most divergent varieties are invariably preserved: a medium form may often long endure, and may or may not produce more than one modified descendant; for natural selection will always act according to the nature of the places which are either unoccupied or not perfectly occupied by other beings; and this will depend on infinitely complex relations. But as a general rule, the more diversified in structure the descendants from any one species can be rendered, the more places they will be enabled to seize on, and the more their modified progeny will increase. In our diagram the line of succession is broken at regular intervals by small numbered letters marking the successive forms which have become sufficiently distinct to be recorded as varieties. But these breaks are imaginary, and might have been inserted anywhere, after intervals long enough to allow the accumulation of a considerable amount of divergent variation.
As all the modified descendants from a common and widely-diffused species, belonging to a large genus, will tend to partake of the same advantages which made their parent successful in life, they will generally go on multiplying in number as well as diverging in character: this is represented in the diagram by the several divergent branches proceeding from (A). The modified offspring from the later and more highly improved branches in the lines of descent, will, it is probable, often take the place of, and so destroy, the earlier and less improved branches: this is represented in the diagram by some of the lower branches not reaching to the upper horizontal lines. In some cases no doubt the process of modification will be confined to a single line of descent, and the number of modified descendants will not be increased; although the amount of divergent modification may have been augmented. This case would be represented in the diagram, if all the lines proceeding from (A) were removed, excepting that from a1 to a10. In the same way the English racehorse and English pointer have apparently both gone on slowly diverging in character from their original stocks, without either having given off any fresh branches or races.
After ten thousand generations, species (A) is supposed to have produced three forms, a10, f10, and m10, which, from having diverged in character during the successive generations, will have come to differ largely, but perhaps unequally, from each other and from their common parent. If we suppose the amount of change between each horizontal line in our diagram to be excessively small, these three forms may still be only well-marked varieties; but we have only to suppose the steps in the process of modification to be more numerous or greater in amount, to convert these three forms into doubtful or at least into well-defined species: thus the diagram illustrates the steps by which the small differences distinguishing varieties are increased into the larger differences distinguishing species. By continuing the same process for a greater number of generations (as shown in the diagram in a condensed and simplified manner), we get eight species, marked by the letters between a14 and m14, all descended from (A). Thus, as I believe, species are multiplied and genera are formed.
In a large genus it is probable that more than one species would vary. In the diagram I have assumed that a second species (I) has produced, by analogous steps, after ten thousand generations, either two well-marked varieties (w10 and z10) or two species, according to the amount of change supposed to be represented between the horizontal lines. After fourteen thousand generations, six new species, marked by the letters n14 to z14, are supposed to have been produced. In any genus, the species which are already very different in character from each other, will generally tend to produce the greatest number of modified descendants; for these will have the best chance of seizing on new and widely different places in the polity of nature: hence in the diagram I have chosen the extreme species (A), and the nearly extreme species (I), as those which have largely varied, and have given rise to new varieties and species. The other nine species (marked by capital letters) of our original genus, may for long but unequal periods continue to transmit unaltered descendants; and this is shown in the diagram by the dotted lines unequally prolonged upwards.
But during the process of modification, represented in the diagram, another of our principles, namely that of extinction, will have played an important part. As in each fully stocked country natural selection necessarily acts by the selected form having some advantage in the struggle for life over other forms, there will be a constant tendency in the improved descendants of any one species to supplant and exterminate in each stage of descent their predecessors and their original progenitor. For it should be remembered that the competition will generally be most severe between those forms which are most nearly related to each other in habits, constitution and structure. Hence all the intermediate forms between the earlier and later states, that is between the less and more improved states of a the same species, as well as the original parent-species itself, will generally tend to become extinct. So it probably will be with many whole collateral lines of descent, which will be conquered by later and improved lines. If, however, the modified offspring of a species get into some distinct country, or become quickly adapted to some quite new station, in which offspring and progenitor do not come into competition, both may continue to exist.
If, then, our diagram be assumed to represent a considerable amount of modification, species (A) and all the earlier varieties will have become extinct, being replaced by eight new species (a14 to m14); and species (I) will be replaced by six (n14 to z14) new species.
But we may go further than this. The original species of our genus were supposed to resemble each other in unequal degrees, as is so generally the case in nature; species (A) being more nearly related to B, C, and D than to the other species; and species (I) more to G, H, K, L, than to the others. These two species (A and I), were also supposed to be very common and widely diffused species, so that they must originally have had some advantage over most of the other species of the genus. Their modified descendants, fourteen in number at the fourteen-thousandth generation, will probably have inherited some of the same advantages: they have also been modified and improved in a diversified manner at each stage of descent, so as to have become adapted to many related places in the natural economy of their country. It seems, therefore, extremely probable that they will have taken the places of, and thus exterminated, not only their parents (A) and (I), but likewise some of the original species which were most nearly related to their parents. Hence very few of the original species will have transmitted offspring to the fourteen-thousandth generation. We may suppose that only one (F) of the two species (E and F) which were least closely related to the other nine original species, has transmitted descendants to this late stage of descent.
The new species in our diagram, descended from the original eleven species, will now be fifteen in number. Owing to the divergent tendency of natural selection, the extreme amount of difference in character between species a14 and z14 will be much greater than that between the most distinct of the original eleven species. The new species, moreover, will be allied to each other in a widely different manner. Of the eight descendants from (A) the three marked a14, q14, p14, will be nearly related from having recently branched off from a10; b14 and f14, from having diverged at an earlier period from a5, will be in some degree distinct from the three first-named species; and lastly, o14, e14, and m14, will be nearly related one to the other, but, from having diverged at the first commencement of the process of modification, will be widely different from the other five species, and may constitute a sub-genus or a distinct genus.
The six descendants from (I) will form two sub-genera or genera. But as the original species (I) differed largely from (A), standing nearly at the extreme end of the original genus, the six descendants from (I) will, owing to inheritance alone, differ considerably from the eight descendants from (A); the two groups, moreover, are supposed to have gone on diverging in different directions. The intermediate species, also (and this is a very important consideration), which connected the original species (A) and (I), have all become, except (F), extinct, and have left no descendants. Hence the six new species descended from (I), and the eight descendants from (A), will have to be ranked as very distinct genera, or even as distinct sub-families.
Thus it is, as I believe, that two or more genera are produced by descent with modification, from two or more species of the same genus. And the two or more parent-species are supposed to be descended from some one species of an earlier genus. In our diagram this is indicated by the broken lines beneath the capital letters, converging in sub-branches downwards towards a single point; this point represents a species, the supposed progenitor of our several new sub-genera and genera.
It is worth while to reflect for a moment on the character of the new species F14, which is supposed not to have diverged much in character, but to have retained the form of (F), either unaltered or altered only in a slight degree. In this case its affinities to the other fourteen new species will be of a curious and circuitous nature. Being descended from a form that stood between the parent-species (A) and (I), now supposed to be extinct and unknown, it will be in some degree intermediate in character between the two groups descended from these two species. But as these two groups have gone on diverging in character from the type of their parents, the new species (F14) will not be directly intermediate between them, but rather between types of the two groups; and every naturalist will be able to call such cases before his mind.
In the diagram each horizontal line has hitherto been supposed to represent a thousand generations, but each may represent a million or more generations; it may also represent a section of the successive strata of the earth’s crust including extinct remains. We shall, when we come to our chapter on geology, have to refer again to this subject, and I think we shall then see that the diagram throws light on the affinities of extinct beings, which, though generally belonging to the same orders, families, or genera, with those now living, yet are often, in some degree, intermediate in character between existing groups; and we can understand this fact, for the extinct species lived at various remote epochs when the branching lines of descent had diverged less.
I see no reason to limit the process of modification, as now explained, to the formation of genera alone. If, in the diagram, we suppose the amount of change represented by each successive group of diverging dotted lines to be great, the forms marked a14 to p14, those marked b14 and f14, and those marked o14 to m14, will form three very distinct genera. We shall also have two very distinct genera descended from (I), differing widely from the descendants of (A). These two groups of genera will thus form two distinct families, or orders, according to the amount of divergent modification supposed to be represented in the diagram. And the two new families, or orders, are descended from two species of the original genus; and these are supposed to be descended from some still more ancient and unknown form.
We have seen that in each country it is the species belonging to the larger genera which oftenest present varieties or incipient species. This, indeed, might have been expected; for as natural selection acts through one form having some advantage over other forms in the struggle for existence, it will chiefly act on those which already have some advantage; and the largeness of any group shows that its species have inherited from a common ancestor some advantage in common. Hence, the struggle for the production of new and modified descendants will mainly lie between the larger groups, which are all trying to increase in number. One large group will slowly conquer another large group, reduce its number, and thus lessen its chance of further variation and improvement. Within the same large group, the later and more highly perfected sub-groups, from branching out and seizing on many new places in the polity of nature, will constantly tend to supplant and destroy the earlier and less improved sub-groups. Small and broken groups and sub-groups will finally disappear. Looking to the future, we can predict that the groups of organic beings which are now large and triumphant, and which are least broken up, that is, which have as yet suffered least extinction, will, for a long period, continue to increase. But which groups will ultimately prevail, no man can predict; for we know that many groups, formerly most extensively developed, have now become extinct. Looking still more remotely to the future, we may predict that, owing to the continued and steady increase of the larger groups, a multitude of smaller groups will become utterly extinct, and leave no modified descendants; and consequently that, of the species living at any one period, extremely few will transmit descendants to a remote futurity. I shall have to return to this subject in the chapter on classification, but I may add that as, according to this view, extremely few of the more ancient species have transmitted descendants to the present day, and, as all the descendants of the same species form a class, we can understand how it is that there exist so few classes in each main division of the animal and vegetable kingdoms. Although few of the most ancient species have left modified descendants, yet, at remote geological periods, the earth may have been almost as well peopled with species of many genera, families, orders and classes, as at the present day.
On the Degree to which Organisation tends to advance.
Natural selection acts exclusively by the preservation and accumulation of variations, which are beneficial under the organic and inorganic conditions to which each creature is exposed at all periods of life. The ultimate result is that each creature tends to become more and more improved in relation to its conditions. This improvement inevitably leads to the gradual advancement of the organisation of the greater number of living beings throughout the world. But here we enter on a very intricate subject, for naturalists have not defined to each other’s satisfaction what is meant by an advance in organisation. Among the vertebrata the degree of intellect and an approach in structure to man clearly come into play. It might be thought that the amount of change which the various parts and organs pass through in their development from embryo to maturity would suffice as a standard of comparison; but there are cases, as with certain parasitic crustaceans, in which several parts of the structure become less perfect, so that the mature animal cannot be called higher than its larva. Von Baer’s standard seems the most widely applicable and the best, namely, the amount of differentiation of the parts of the same organic being, in the adult state, as I should be inclined to add, and their specialisation for different functions; or, as Milne Edwards would express it, the completeness of the division of physiological labour. But we shall see how obscure this subject is if we look, for instance, to fishes, among which some naturalists rank those as highest which, like the sharks, approach nearest to amphibians; while other naturalists rank the common bony or teleostean fishes as the highest, inasmuch as they are most strictly fish-like, and differ most from the other vertebrate classes. We see still more plainly the obscurity of the subject by turning to plants, among which the standard of intellect is of course quite excluded; and here some botanists rank those plants as highest which have every organ, as sepals, petals, stamens and pistils, fully developed in each flower; whereas other botanists, probably with more truth, look at the plants which have their several organs much modified and reduced in number as the highest.
If we take as the standard of high organisation, the amount of differentiation and specialisation of the several organs in each being when adult (and this will include the advancement of the brain for intellectual purposes), natural selection clearly leads towards this standard: for all physiologists admit that the specialisation of organs, inasmuch as in this state they perform their functions better, is an advantage to each being; and hence the accumulation of variations tending towards specialisation is within the scope of natural selection. On the other hand, we can see, bearing in mind that all organic beings are striving to increase at a high ratio and to seize on every unoccupied or less well occupied place in the economy of nature, that it is quite possible for natural selection gradually to fit a being to a situation in which several organs would be superfluous or useless: in such cases there would be retrogression in the scale of organisation. Whether organisation on the whole has actually advanced from the remotest geological periods to the present day will be more conveniently discussed in our chapter on Geological Succession.
But it may be objected that if all organic beings thus tend to rise in the scale, how is it that throughout the world a multitude of the lowest forms still exist; and how is it that in each great class some forms are far more highly developed than others? Why have not the more highly developed forms every where supplanted and exterminated the lower? Lamarck, who believed in an innate and inevitable tendency towards perfection in all organic beings, seems to have felt this difficulty so strongly that he was led to suppose that new and simple forms are continually being produced by spontaneous generation. Science has not as yet proved the truth of this belief, whatever the future may reveal. On our theory the continued existence of lowly organisms offers no difficulty; for natural selection, or the survival of the fittest, does not necessarily include progressive development—it only takes advantage of such variations as arise and are beneficial to each creature under its complex relations of life. And it may be asked what advantage, as far as we can see, would it be to an infusorian animalcule—to an intestinal worm—or even to an earth-worm, to be highly organised. If it were no advantage, these forms would be left, by natural selection, unimproved or but little improved, and might remain for indefinite ages in their present lowly condition. And geology tells us that some of the lowest forms, as the infusoria and rhizopods, have remained for an enormous period in nearly their present state. But to suppose that most of the many now existing low forms have not in the least advanced since the first dawn of life would be extremely rash; for every naturalist who has dissected some of the beings now ranked as very low in the scale, must have been struck with their really wondrous and beautiful organisation.
Nearly the same remarks are applicable, if we look to the different grades of organisation within the same great group; for instance, in the vertebrata, to the co-existence of mammals and fish—among mammalia, to the co-existence of man and the ornithorhynchus—among fishes, to the co-existence of the shark and the lancelet (Amphioxus), which latter fish in the extreme simplicity of its structure approaches the invertebrate classes. But mammals and fish hardly come into competition with each other; the advancement of the whole class of mammals, or of certain members in this class, to the highest grade would not lead to their taking the place of fishes. Physiologists believe that the brain must be bathed by warm blood to be highly active, and this requires aërial respiration; so that warm-blooded mammals when inhabiting the water lie under a disadvantage in having to come continually to the surface to breathe. With fishes, members of the shark family would not tend to supplant the lancelet; for the lancelet, as I hear from Fritz Müller, has as sole companion and competitor on the barren sandy shore of South Brazil, an anomalous annelid. The three lowest orders of mammals, namely, marsupials, edentata, and rodents, co-exist in South America in the same region with numerous monkeys, and probably interfere little with each other. Although organisation, on the whole, may have advanced and be still advancing throughout the world, yet the scale will always present many degrees of perfection; for the high advancement of certain whole classes, or of certain members of each class, does not at all necessarily lead to the extinction of those groups with which they do not enter into close competition. In some cases, as we shall hereafter see, lowly organised forms appear to have been preserved to the present day, from inhabiting confined or peculiar stations, where they have been subjected to less severe competition, and where their scanty numbers have retarded the chance of favourable variations arising.
Finally, I believe that many lowly organised forms now exist throughout the world, from various causes. In some cases variations or individual differences of a favourable nature may never have arisen for natural selection to act on and accumulate. In no case, probably, has time sufficed for the utmost possible amount of development. In some few cases there has been what we must call retrogression or organisation. But the main cause lies in the fact that under very simple conditions of life a high organisation would be of no service—possibly would be of actual disservice, as being of a more delicate nature, and more liable to be put out of order and injured.
Looking to the first dawn of life, when all organic beings, as we may believe, presented the simplest structure, how, it has been asked, could the first step in the advancement or differentiation of parts have arisen? Mr. Herbert Spencer would probably answer that, as soon as simple unicellular organisms came by growth or division to be compounded of several cells, or became attached to any supporting surface, his law “that homologous units of any order become differentiated in proportion as their relations to incident forces become different” would come into action. But as we have no facts to guide us, speculation on the subject is almost useless. It is, however, an error to suppose that there would be no struggle for existence, and, consequently, no natural selection, until many forms had been produced: variations in a single species inhabiting an isolated station might be beneficial, and thus the whole mass of individuals might be modified, or two distinct forms might arise. But, as I remarked towards the close of the introduction, no one ought to feel surprise at much remaining as yet unexplained on the origin of species, if we make due allowance for our profound ignorance on the mutual relations of the inhabitants of the world at the present time, and still more so during past ages.
Convergence of Character.
Mr. H.C. Watson thinks that I have overrated the importance of divergence of character (in which, however, he apparently believes), and that convergence, as it may be called, has likewise played a part. If two species belonging to two distinct though allied genera, had both produced a large number of new and divergent forms, it is conceivable that these might approach each other so closely that they would have all to be classed under the same genus; and thus the descendants of two distinct genera would converge into one. But it would in most cases be extremely rash to attribute to convergence a close and general similarity of structure in the modified descendants of widely distinct forms. The shape of a crystal is determined solely by the molecular forces, and it is not surprising that dissimilar substances should sometimes assume the same form; but with organic beings we should bear in mind that the form of each depends on an infinitude of complex relations, namely on the variations which have arisen, these being due to causes far too intricate to be followed out—on the nature of the variations which have been preserved or selected, and this depends on the surrounding physical conditions, and in a still higher degree on the surrounding organisms with which each being has come into competition—and lastly, on inheritance (in itself a fluctuating element) from innumerable progenitors, all of which have had their forms determined through equally complex relations. It is incredible that the descendants of two organisms, which had originally differed in a marked manner, should ever afterwards converge so closely as to lead to a near approach to identity throughout their whole organisation. If this had occurred, we should meet with the same form, independently of genetic connection, recurring in widely separated geological formations; and the balance of evidence is opposed to any such an admission.
Mr. Watson has also objected that the continued action of natural selection, together with divergence of character, would tend to make an indefinite number of specific forms. As far as mere inorganic conditions are concerned, it seems probable that a sufficient number of species would soon become adapted to all considerable diversities of heat, moisture, &c.; but I fully admit that the mutual relations of organic beings are more important; and as the number of species in any country goes on increasing, the organic conditions of life must become more and more complex. Consequently there seems at first no limit to the amount of profitable diversification of structure, and therefore no limit to the number of species which might be produced. We do not know that even the most prolific area is fully stocked with specific forms: at the Cape of Good Hope and in Australia, which support such an astonishing number of species, many European plants have become naturalised. But geology shows us, that from an early part of the tertiary period the number of species of shells, and that from the middle part of this same period, the number of mammals has not greatly or at all increased. What then checks an indefinite increase in the number of species? The amount of life (I do not mean the number of specific forms) supported on an area must have a limit, depending so largely as it does on physical conditions; therefore, if an area be inhabited by very many species, each or nearly each species will be represented by few individuals; and such species will be liable to extermination from accidental fluctuations in the nature of the seasons or in the number of their enemies. The process of extermination in such cases would be rapid, whereas the production of new species must always be slow. Imagine the extreme case of as many species as individuals in England, and the first severe winter or very dry summer would exterminate thousands on thousands of species. Rare species, and each species will become rare if the number of species in any country becomes indefinitely increased, will, on the principal often explained, present within a given period few favourable variations; consequently, the process of giving birth to new specific forms would thus be retarded. When any species becomes very rare, close interbreeding will help to exterminate it; authors have thought that this comes into play in accounting for the deterioration of the aurochs in Lithuania, of red deer in Scotland and of bears in Norway, &c. Lastly, and this I am inclined to think is the most important element, a dominant species, which has already beaten many competitors in its own home, will tend to spread and supplant many others. Alph. de Candolle has shown that those species which spread widely tend generally to spread very widely, consequently they will tend to supplant and exterminate several species in several areas, and thus check the inordinate increase of specific forms throughout the world. Dr. Hooker has recently shown that in the southeast corner of Australia, where, apparently, there are many invaders from different quarters of the globe, the endemic Australian species have been greatly reduced in number. How much weight to attribute to these several considerations I will not pretend to say; but conjointly they must limit in each country the tendency to an indefinite augmentation of specific forms.
Summary of Chapter.
If under changing conditions of life organic beings present individual differences in almost every part of their structure, and this cannot be disputed; if there be, owing to their geometrical rate of increase, a severe struggle for life at some age, season or year, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of life, causing an infinite diversity in structure, constitution, and habits, to be advantageous to them, it would be a most extraordinary fact if no variations had ever occurred useful to each being’s own welfare, in the same manner as so many variations have occurred useful to man. But if variations useful to any organic being ever do occur, assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life; and from the strong principle of inheritance, these will tend to produce offspring similarly characterised. This principle of preservation, or the survival of the fittest, I have called Natural Selection. It leads to the improvement of each creature in relation to its organic and inorganic conditions of life; and consequently, in most cases, to what must be regarded as an advance in organisation. Nevertheless, low and simple forms will long endure if well fitted for their simple conditions of life.
Natural selection, on the principle of qualities being inherited at corresponding ages, can modify the egg, seed, or young as easily as the adult. Among many animals sexual selection will have given its aid to ordinary selection by assuring to the most vigorous and best adapted males the greatest number of offspring. Sexual selection will also give characters useful to the males alone in their struggles or rivalry with other males; and these characters will be transmitted to one sex or to both sexes, according to the form of inheritance which prevails.
Whether natural selection has really thus acted in adapting the various forms of life to their several conditions and stations, must be judged by the general tenour and balance of evidence given in the following chapters. But we have already seen how it entails extinction; and how largely extinction has acted in the world’s history, geology plainly declares. Natural selection, also, leads to divergence of character; for the more organic beings diverge in structure, habits and constitution, by so much the more can a large number be supported on the area, of which we see proof by looking to the inhabitants of any small spot, and to the productions naturalised in foreign lands. Therefore, during the modification of the descendants of any one species, and during the incessant struggle of all species to increase in numbers, the more diversified the descendants become, the better will be their chance of success in the battle for life. Thus the small differences distinguishing varieties of the same species, steadily tend to increase, till they equal the greater differences between species of the same genus, or even of distinct genera.
We have seen that it is the common, the widely diffused, and widely ranging species, belonging to the larger genera within each class, which vary most; and these tend to transmit to their modified offspring that superiority which now makes them dominant in their own countries. Natural selection, as has just been remarked, leads to divergence of character and to much extinction of the less improved and intermediate forms of life. On these principles, the nature of the affinities, and the generally well defined distinctions between the innumerable organic beings in each class throughout the world, may be explained. It is a truly wonderful fact—the wonder of which we are apt to overlook from familiarity—that all animals and all plants throughout all time and space should be related to each other in groups, subordinate to groups, in the manner which we everywhere behold—namely, varieties of the same species most closely related, species of the same genus less closely and unequally related, forming sections and sub-genera, species of distinct genera much less closely related, and genera related in different degrees, forming sub-families, families, orders, sub-classes, and classes. The several subordinate groups in any class cannot be ranked in a single file, but seem clustered round points, and these round other points, and so on in almost endless cycles. If species had been independently created, no explanation would have been possible of this kind of classification; but it is explained through inheritance and the complex action of natural selection, entailing extinction and divergence of character, as we have seen illustrated in the diagram.
The affinities of all the beings of the same class have sometimes been represented by a great tree. I believe this simile largely speaks the truth. The green and budding twigs may represent existing species; and those produced during former years may represent the long succession of extinct species. At each period of growth all the growing twigs have tried to branch out on all sides, and to overtop and kill the surrounding twigs and branches, in the same manner as species and groups of species have at all times overmastered other species in the great battle for life. The limbs divided into great branches, and these into lesser and lesser branches, were themselves once, when the tree was young, budding twigs; and this connexion of the former and present buds by ramifying branches may well represent the classification of all extinct and living species in groups subordinate to groups. Of the many twigs which flourished when the tree was a mere bush, only two or three, now grown into great branches, yet survive and bear the other branches; so with the species which lived during long-past geological periods, very few have left living and modified descendants. From the first growth of the tree, many a limb and branch has decayed and dropped off; and these fallen branches of various sizes may represent those whole orders, families, and genera which have now no living representatives, and which are known to us only in a fossil state. As we here and there see a thin, straggling branch springing from a fork low down in a tree, and which by some chance has been favoured and is still alive on its summit, so we occasionally see an animal like the Ornithorhynchus or Lepidosiren, which in some small degree connects by its affinities two large branches of life, and which has apparently been saved from fatal competition by having inhabited a protected station. As buds give rise by growth to fresh buds, and these, if vigorous, branch out and overtop on all sides many a feebler branch, so by generation I believe it has been with the great Tree of Life, which fills with its dead and broken branches the crust of the earth, and covers the surface with its ever-branching and beautiful ramifications.
