On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life

Effects of external conditions — Use and disuse, combined with natural selection ; organs of flight and of vision—Acclimatisa- tion — Correlation of growth — Compensation and economy of growth—False correlations—Multiple, rudimentary, and lowly organised structures variable—Parts developed in an unusual manner are highly variable: specific characters more variable than generic: secondary sexual characters variable—Species of the same genus vary in an analogous manner—Reversions to long lost characters—Summary.

I HAVE hitherto sometimes spoken as if the variations —so common and multiform in organic beings under domestication, and in a lesser degree in those in a state of nature—had been due to chance. This, of course, is a wholly incorrect expression, but it serves to acknow- ledge plainly our ignorance of the cause of each parti- cular variation. Some authors believe it to be as much the function of the reproductive system to produce individual differences, or very slight deviations of struc- ture, as to make the child like its parents. But the much greater variability, as well as the greater fre- quency of monstrosities, under domestication or culti- vation, than under nature, leads me to believe that deviations of structure are in some way due to the nature of the conditions of life, to which the parents and their more remote ancestors have been exposed during several generations. I have remarked in the first chapter—but a long catalogue of facts which can- not be here given would be necessary to show the truth of the remark—that the reproductive system is eminently susceptible to changes in the conditions of life; and to

this system being functionally disturbed in the parents, I chiefly attribute the varying or plastic condition of the offspring. The male and female sexual elements seem to be affected before that union takes place which is to form a new being. In the case of “sporting” plants, the bud, which in its earliest condition does not apparently differ essentially from an ovule, is alone affected. But why, because the reproductive system is disturbed, this or that part should vary more or less, we are profoundly ignorant. Nevertheless, we can here and there dimly catch a faint ray of light, and we may feel sure that there must be some cause for each deviation of structure, however slight.

How much direct effect difference of climate, food, &e., produces on any being is extremely doubtful. My impression is, that the effect is extremely small im the case of animals, but perhaps rather more in that of plants. We may, at least, safely conclude that such influences cannot have produced the many striking and complex co-adaptations of structure between one organic being and another, which we see everywhere throughout nature. Some little influence may be attri- buted to climate, food, &e.: thus, E. Forbes speaks confidently that shells at their southern limit, and when living in shallow water, are more brightly coloured than those of the same species further north or from greater

are more brightly coloured under a clear atmosphere, than when living on islands or near the coast. So with insects, Wollaston is convinced that residence near the sea affects their colours. Moquin-Tandon gives a list of plants which when growing near the sea-shore have their leaves in some degree fleshy, though not elsewhere fleshy. Several other such cases could be given. into the zone of habitation of other species, often acquiring in a very slight degree some of the characters of such species, accords with our view that species of all kinds are only well-marked and permanent varieties. Thus the species of shells which are confined to tropical and shallow seas are generally brighter-coloured than those confined to cold and deeper seas. The birds which are confined to continents are, according to Mr. Gould, brighter-coloured than those of islands. The insect-species confined to sea-coasts, as every collector knows, are often brassy or lurid. Plants which live exclusively on the sea-side are very apt to have fleshy leaves. He who believes in the creation of each spe- cies, will have to say that this shell, for instance, was created with bright colours for a warm sea; but that this other shell became bright-coloured by variation when it ranged into warmer or shallower waters.

When a variation is of the slightest use to a being, we cannot tell how much of it to attribute to the accu- mulative action of natural selection, and how much to the conditions of life. Thus, it is well known to fur- riers that animals of the same species have thicker and better fur the more severe the climate is under which they have lived; but who can tell how much of this difference may be due to the warmest-clad individuals having been favoured and preserved during many generations, and how much to the direct action of the severe climate? for it would appear that climate has some direct action on the hair of our domestic quadrupeds.

Instances could be given of the same variety being produced under conditions of life as different as can well be conceived ; and, on the other hand, of different yarieties being produced from the same species under the same conditions. Such facts show how indirectly the conditions of life must act. Agaim, innumerable instances are known to every naturalist of species keeping true, or not varying at all, although living under the most opposite climates. Such considerations as these incline me to lay very little weight on the direct action of the conditions of life. Indirectly, as already remarked, they seem to play an important part in affecting the reproductive system, and in thus in- ducing variability ; and natural selection will then accu- mulate all profitable variations, however slight, until they become plainly developed and appreciable by us.

Effects of Use and Disuse.-—From the facts alluded to in the first chapter, I think there can be little doubt that use in our domestic animals strengthens and en- larges certain parts, and disuse diminishes them; and that such modifications are inherited. Under free nature, we can have no standard of comparison, by which to judge of the effects of long-continued use or disuse, for we know not the parent-forms; but many animals have structures which can be explained by the effects of disuse. As Professor Owen has remarked, there is no greater anomaly in nature than a bird that cannot fly; yet there are several in this state. The logger- headed duck of South America can only flap along the surface of the water, and has its wings in nearly the same condition as the domestic Aylesbury duck. As the larger ground-feeding birds seldom take flight except to escape danger, I believe that the nearly wingless condi- tion of several birds, which now inhabit or have lately inhabited several oceanic islands, tenanted by no beast of prey, has been caused by disuse. The ostrich indeed inhabits continents and is exposed to danger from which it cannot escape by flight, but by kicking it can defend itself from enemies, as well as any of the smaller

quadrupeds. We may imagine that the early progenitor of the ostrich had habits like those of a bustard, and that as natural selection increased in successive genera- tions the size and weight of its body, its legs were used more, and its wings less, until they became incapable of flight. Kirby has remarked (and I have observed the same fact) that the anterior tarsi, or feet, of many male dung- feeding beetles are very often broken off; he examined seventeen specimens in his own collection, and not one had even a relic left. In the Onites apelles the tarsi are so habitually lost, that the insect has been described as not having them. In some other genera they are pre- sent, but in a rudimentary condition. In the Ateuchus or sacred beetle of the Egyptians, they are totally defi- cient. There is not sufficient evidence to induce us to believe that mutilations are ever inherited; and I should prefer explaining the entire absence of the ante- rior tarsi in Ateuchus, and their rudimentary condition in some other genera, by the long-continued effects of disuse in their progenitors; for as the tarsi are almost always lost in many dung-feeding beetles, they must be lost early in life, and therefore cannot be much used by these insects.

In some cases we might easily put down to disuse modifications of structure which are wholly, or mainly, due to natural selection. Mr. Wollaston has discovered the remarkable fact that 200 beetles, out of the 550 species inhabiting Madeira, are so far deficient in wings that they cannot fly; and that of the twenty-nine endemic genera, no less than twenty-three genera have all their species in this condition! Several facts, namely, that beetles in many parts of the world are very fre- quently blown to sea and perish; that the beetles in Madeira, as observed by Mr. Wollaston, lie much con-

cealed, until the wind lulls and the sun shines; that the proportion of wingless beetles is larger on the ex- posed Dezertas than in Madeira itself; and especially the extraordinary fact, so strongly insisted on by Mr. Wollaston, of the almost entire absence of certain large groups of beetles, elsewhere excessively numerous, and which groups have habits of life almost necessitating frequent flight ;—these several considerations have made me believe that the wingless condition of so many Madeira beetles is mainly due to the action of natural selection, but combined probably with disuse. For during thousands of successive generations each indi- vidual beetle which flew least, either from its wings having been ever so little less perfectly developed or from indolent habit, will have had the best chance of surviving from not being blown out to sea; and, on the other hand, those beetles which most readily took to flight will oftenest have been blown to sea and thus have been destroyed.

The insects in Madeira which are not ground-feeders, and which, as the flower-feeding coleoptera and lepidop- tera, must habitually use their wings to gain their subsist- ence, have, as Mr. Wollaston suspects, their wings not at all reduced, but even enlarged. This is quite com- patible with the action of natural selection. For when a new insect first arrived on the island, the tendency of natural selection to enlarge or to reduce the wings, would depend on whether a greater number of indivi- duals were saved by successfully battling with the winds, or by giving up the attempt and rarely or never flying. As with mariners shipwrecked near a coast, it would have been better for the good swimmers if they had been able to swim still further, whereas it would have been better for the bad swimmers if they had not been able to swim at all and had stuck to the wreck.

The eyes of moles and of some burrowing rodents are rudimentary in size, and in some cases are quite covered up by skin and fur. This state of the eyes is probably due to gradual reduction from disuse, but aided perhaps by natural selection. In South America, a burrowing rodent, the tuco-tuco, or Ctenomys, is even more subter- ranean in its habits than the mole; and I was assured by a Spaniard, who had often caught them, that they were frequently blind; one which I kept alive was cer- tainly in this condition, the cause, as appeared on dis- section, having been inflammation of the nictitating membrane. As frequent inflammation of the eyes must be injurious to any animal, and as eyes are certainly not indispensable to animals with subterranean habits, a reduction in their size with the adhesion of the eye- lids and growth of fur over them, might in such case be an advantage; and if so, natural selection would con- stantly aid the effects of disuse.

It is well known that several animals, belonging to the most different classes, which inhabit the caves of Styria and of Kentucky, are blind. In some of the crabs the foot-stalk for the eye remains, though the eye is gone; the stand for the telescope is there, though the telescope with its glasses has been lost. As it is difficult to ima- gine that eyes, though useless, could be in any way injurious to animals living in darkness, I attribute their loss wholly to disuse. In one of the blind animals, namely, the caye-rat, the eyes are of immense size; and Professor Silliman thought that it regained, after living some days in the light, some slight power of vision. In the same manner as in Madeira the wings of some of the insects have been enlarged, and the wings of others have been reduced by natural selection aided by use and disuse, so in the case of the cave-rat natural selec- tion seems to have struggled with the loss of light and

to have increased the size of the eyes; whereas with all the other inhabitants of the caves, disuse by itself seems to have done its work. It is difficult to imagine conditions of life more similar than deep limestone caverns under a nearly similar climate; so that on the common view of the blind ani- mals having been separately created for the American and European caverns, close similarity in their organisa- tion and affinities might have been expected; but, as Schiddte and others have remarked, this is not the case, and the cave-insects of the two continents are not more closely allied than might have been anticipated from the general resemblance of the other inhabitants of North America and Europe. On my view we must suppose that American animals, having ordinary powers of vision, slowly migrated by successive generations from the outer world into the deeper and deeper recesses of the Kentucky caves, as did European animals into the caves of Europe. We have some evidence of this gra- dation of habit; for, as Schiédte remarks, “animals not far remote from ordinary forms, prepare the transi-

constructed for twilight; and, last of all, those destined for total darkness.” By the time that an animal had reached, after numberless generations, the deepest re- cesses, disuse will on this view have more or less per- fectly obliterated its eyes, and natural selection will often have effected other changes, such as an increase in the length of the antennz or palpi, as a compensa- tion for blindness. Notwithstanding such modifications, we might expect still to see in the cave-animals of America, affinities to the other inhabitants of that con- tinent, and in those of Europe, to the inhabitants of the European continent. And this is the case with some of the American cave-animals, as I hear from

Professor Dana; and some of the European cave- insects are very closely allied to those of the surround- ing country. It would be most difficult to give any rational explanation of the affinities of the blind cave- animals to the other inhabitants of the two continents on the ordinary view of their independent creation. That several of the inhabitants of the caves of the Old and New Worlds should be closely related, we might expect from the well-known relationship of most of their other productions. Far from feeling any sur- prise that some of the cave-animals should be very anomalous, as Agassiz has remarked in regard to the blind fish, the Amblyopsis, and as is the case with the blind Proteus with reference to the reptiles of Europe, Tam only surprised that more wrecks of ancient life have not been preserved, owing to the less severe com- petition to which the inhabitants of these dark abodes will probably have been exposed.

Acclimatisation—Habit is hereditary with plants, as in the period of flowering, in the amount of rain requi- site for seeds to germinate, in the time of sleep, &c., and this leads me to say a few words on acclimatisa- tion. As it is extremely common for species of the same genus to inhabit very hot and very cold countries, and as I believe that all the species of the same genus have descended from a single parent, if this view be correct, acclimatisation must be readily effected during long-continued descent. It is notorious that each species is adapted to the climate of its own home: species from an arctic or even from a temperate region cannot endure a tropical climate, or conversely. So again, many succulent plants cannot endure a damp climate. But the degree of adaptation of species to the climates under which they live is often overrated.

We may infer this from our frequent inability to pre- dict whether or not an imported plant will endure our climate, and from the number of plants and animals brought from warmer countries which here enjoy good health. We have reason to believe that species in a state of nature are limited in their ranges by the com- petition of other organic beings quite as much as, or more than, by adaptation to particular climates. But whether or not the adaptation be generally very close, we have evidence, in the case of some few plants, of their becoming, to a certain extent, naturally habituated to different temperatures, or becoming acclimatised: thus the pines and rhododendrong, raised from seed collected by Dr. Hooker from trees growing at different heights on the Himalaya, were found in this country to possess different constitutional powers of resisting cold. Mr. Thwaites informs me that he has observed similar facts in Ceylon, and analogous observations have been made by Mr. H. C. Watson on European species of plants brought from the Azores to England. In regard to animals, several authentic cases could be given of species within historical times having largely extended their range from warmer to cooler latitudes, and con- versely; but we do not positively know that these ani- mals were strictly adapted to their native climate, but in all ordinary cases we assume such to be the case ; nor do we know that they have subsequently become acclimatised to their new homes.

As I believe that our domestic animals were origin- ally chosen by uncivilised man because they were use- ful and bred readily under confinement, and not because they were subsequently found capable of far-extended transportation, I think the common and extraordinary capacity in our domestic animals of not only withstand- ing the most different climates but of being perfectly fertile (a far severer test) under them, may be used as an argument that a large proportion of other animals, now in a state of nature, could easily be brought to bear widely different climates. We must not, however, push the foregomg argument too far, on account of the pro- bable origin of some of our domestic animals from seve- ral wild stocks: the blood, for instance, of a tropical and arctic wolf or wild dog may perhaps be mingled in our domestic breeds. The rat and mouse cannot be considered as domestic animals, but they have been transported by man to many parts of the world, and now have a far wider range than any other rodent, living free under the cold climate of Faroe in the north and of the Falklands in the south, and on many islands in the torrid zones. Hence I am inclined to look at adaptation to any special climate as a quality readily grafted on an innate wide flexibility of constitution, which is common to most animals. On this view, the capacity of enduring the most different climates by man himself and by his domestic animals, and such facts as that former species of the elephant and rhinoceros were capable of enduring a glacial climate, whereas the liv- ing species are now all tropical or sub-tropical in their habits, ought not to be looked at as anomalies, but merely as examples of a very common flexibility of con- stitution, brought, under peculiar circumstances, into play.

How much of the acclimatisation of species to any peculiar climate is due to mere habit, and how much to the natural selection of varieties having different innate constitutions, and how much to both means combined, is a very obscure question. That habit or custom has some influence I must believe, both from analogy, and from the incessant advice given in agricultural works, even in the ancient Encyclopedias of China, to be very cau- tious in transposing animals from one district to an- other; for it is not likely that man should have suc- ceeded in selecting so many breeds and sub-breeds with constitutions specially fitted for their own districts: the result must, I think, be due to habit. On the other hand, I can see no reason to doubt that natural selection will continually tend to preserve those individuals which are born with constitutions best adapted to their native countries. In treatises on many kinds of cultivated plants, certain varieties are said to withstand certain climates better than others: this is very strikingly shown in works on fruit trees published in the United States, in which certain varieties are habitually recom- mended for the northern, and others for the southern States; and as most of these varieties are of recent origin, they cannot owe their constitutional differences to habit. The case of the Jerusalem artichoke, which is never propagated by seed, and of which consequently new varieties have not been produced, has even been advanced—for it is now as tender as ever it was—as proving that acclimatisation cannot be effected! The case, also, of the kidney-bean has been often cited for a similar purpose, and with much greater weight; but until some one will sow, during a score of generations, his kidney-beans so early that a very large proportion are destroyed by frost, and then collect seed from the few survivors, with care to prevent accidental crosses, and then again get seed from these seedlings, with the same precautions, the experiment cannot be said to have been even tried. Nor let it be supposed that no differences in the constitution of seedling kidney-beans ever appear, for an account has been published how much more hardy some seedlings appeared to be than others.

use, and disuse, have, in some cases, played a consider- able part in the modification of the constitution, and of the structure of various organs; but that the effects of use and disuse have often been largely combined with, and sometimes overmastered by, the natural selec- tion of innate differences. Correlation of Growth—I mean by this expression that the whole organisation is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified. This is a very important subject, most imperfectly understood. The most obvious case is, that modifications accumulated solely for the good of the young or larva, will, it may safely be concluded, affect the structure of the adult; in the same manner as any malconformation affecting the early embryo, seriously affects the whole organisa- tion of the adult. The several parts of the body which are homologous, and which, at an early embryonic period, are alike, seem liable to vary in an allied manner: we see this in the right and left sides of the body varying in the same manner; in the front and hind legs, and eyen in the jaws and limbs, varying together, for the lower jaw is believed to be homologous with the limbs. These tendencies, I do not doubt, may be mastered more or less completely by natural selection: thus a family of stags once existed with an antler only on one side; and if this had been of any great use to the breed it might probably have been rendered permanent by natural selection.

Homologous parts, as has been remarked by some authors, tend to cohere; this is often seen in monstrous plants; and nothing is more common than the union of homologous parts in normal structures, as the union of the petals of the corolla into a tube. Hard parts seem to affect the form of adjoining soft parts; it is believed by some authors that the diversity in the shape of the pelvis in birds causes the remarkable diversity in the shape of their kidneys. Others believe that the shape of the pelvis in the human mother influences by pres- sure the shape of the head of the child. In snakes, according to Schlegel, the shape of the body and the manner of swallowing determine the position of several of the most important viscera.

The nature of the bond of correlation is very fre- quently quite obscure. M. Is. Geoffroy St. Hilaire has forcibly remarked, that certam malconformations very frequently, and that others rarely coexist, without our being able to assign any reason. What can be more singular than the relation between blue eyes and deaf- ness in cats, and the tortoise-shell colour with the female sex; the feathered feet and skin between the outer toes in pigeons, and the presence of more or less down on the young birds when first hatched, with the future colour of their plumage ; or, again, the relation between the hair and teeth in the naked Turkish dog, though here pro- bably homology comes into play? With respect to this latter case of correlation, I think it can hardly be acci- dental, that if we pick out the two orders of mammalia which are most abnormal in their dermal covering, viz. Cetacea (whales) and Edentata (armadilloes, scaly ant- eaters, &c.), that these are likewise the most abnormal in their teeth.

I know of no case better adapted to show the im- portance of the laws of correlation in modifying im- portant structures, independently of utility and, there- fore, of natural selection, than that of the difference between the outer and inner flowers in some Compo- sitous and Umbelliferous plants. Every one knows the difference in the ray and central florets of, for instance, the daisy, and this difference is often accompanied with the abortion of parts of the flower. But, in some Com- positous plants, the seeds also differ in shape and sculp- ture; and even the ovary itself, with its accessory parts, differs, as has been described by Cassini. These differ- ences have been attributed bz some authors to pressure, and the shape of the seeds in the ray-florets in some Composite countenances this idea; but, in the case of the corolla of the Umbelliferze, it is by no means, as Dr. Hooker informs me, in species with the densest heads that the inner and outer flowers most frequently differ. It might have been thought that the development of the ray-petals by drawing nourishment from certain other parts of the flower had caused their abortion ; but in some Composite there is a difference in the seeds of the outer and inner florets without any difference in the corolla. Possibly, these several differences may be con- nected with some difference in the flow of nutriment towards the central and external flowers: we know, at least, that in irregular flowers, those nearest to the axis are oftenest subject to peloria, and become regular. I may add, as an instance of this, and of a striking case of correlation, that I have recently observed in some garden pelargoniums, that the central flower of the truss often loses the patches of darker colour in the two upper petals; and that when this occurs, the adhe- rent nectary is quite aborted; when the colour is absent from only one of the two upper petals, the nectary is only much shortened.

With respect to the difference in the corolla of the central and exterior flowers of a head or umbel, I do not feel at all sure that C. C. Sprengel’s idea that the ray-florets serve to attract insects, whose agency is highly advantageous in the fertilisation of plants of these two orders, is so far-fetched, as it may at first appear: and if it be advantageous, natural selection may have come into play. But in regard to the differ- ences both in the internal and external structure of the seeds, which are not always correlated with any differ- ences in the flowers, it seems impossible that they can be in any way advantageous to the plant: yet in the Umbelliferce these differences are of such apparent im- portance—the seeds being in some cases, according to Tausch, orthospermous in the exterior flowers and cce- lospermous in the central flowers,—that the elder De Candolle founded his main divisions of the order on analogous differences. Hence we see that modifica- tions of structure, viewed by systematists as of high value, may be wholly due to unknown laws of correlated erowth, and without being, as far as we can see, of the slightest service to the species.

We may often falsely attribute to correlation of growth, structures which are common to whole groups of species, and which in truth are simply due to inheritance ; for an ancient progenitor may have acquired through natural selection some one modification in structure, and, after thousands of generations, some other and in- dependent modification; and these two modifications, having been transmitted to a whole group of descendants with diverse habits, would naturally be thought to be correlated in some necessary manner. So, again, I do not doubt that some apparent correlations, occurring throughout whole orders, are entirely due to the manner alone in which natural selection can act. For instance, Alph. De Candolle has remarked that winged seeds are never found in fruits which do not open: I should ex- plain the rule by the fact that seeds could not gradually become winged through natural selection, except in fruits which opened; so that the individual plants producing

seeds which were a little better fitted to be wafted further, might get an advantage over those producing seed less fitted for dispersal; and this process could not possibly go on in fruit which did not open. The elder Geoffroy and Goethe propounded, at about the same period, their law of compensation or balance- ment of growth; or, as Goethe expressed it, “in order to spend on one side, nature is forced to economise on the other side.” I think this holds true to a certain ex- tent with our domestic productions: if nourishment flows to one part or organ in excess, it rarely flows, at least in excess, to another part; thus it is difficult to get a cow to give much milk and to fatten readily. The same va- rieties of the cabbage do not yield abundant and nutri- tious foliage and a copious supply of oil-bearing seeds. When the seeds in our fruits become atrophied, the fruit itself gains largely in size and quality. In our poultry, a large tuft of feathers on the head is generally accom- panied by a diminished comb, and a large beard by diminished wattles. With species in a state of nature it can hardly be maintained that the law is of universal application ; but many good observers, more especially botanists, believe in its truth. I will not, however, here give any instances, for I see hardly any way of distin- guishing between the effects, on the one hand, of a part being largely developed through natural selection and another and adjoining part being reduced by this same process or by disuse, and, on the other hand, the actual withdrawal of nutriment from one part owing to the excess of growth in another and adjoining part.

I suspect, also, that some of the cases of compensation which have been advanced, and likewise some other facts, may be merged under a more general principle, namely, that natural selection is continually trying to economise in every part of the organisation. If under changed conditions of life a structure before useful be- comes less useful, any diminution, however slight, in its development, will be seized on by natural selection, for it will profit the individual not to have its nutriment wasted in building up an useless structure. I can thus only understand a fact with which I was much struck when examining cirripedes, and of which many other instances could be given: namely, that when a cirripede is parasitic within another and is thus protected, it loses more or less completely its own shell or carapace. This is the case with the male Ibla, and in a truly extraordi- nary manner with the Proteolepas: for the carapace in all other cirripedes consists of the three highly-important anterior segments of the head enormously developed, and furnished with great nerves and muscles; but in the parasitic and protected Proteolepas, the whole ante- rior part of the head is reduced to the merest rudiment attached to the bases of the prehensile antenne. Now the saving of a large and complex structure, when ren- dered superfluous by the parasitic habits of the Proteo- lepas, though effected by slow steps, would be a decided advantage to each successive individual of the species ; for in the struggle for life to which every animal is ex- posed, each individual Proteolepas would have a better chance of supporting itself, by less nutriment being wasted in developing a structure now become useless.

Thus, as I believe, natural selection will always suc- ceed in the long run in reducing and saving every part of the organisation, as soon as it is rendered superfluous, without by any means causing some other part to be largely developed in a corresponding degree. And, con- versely, that natural selection may perfectly well suc- ceed in largely developing any organ, without requiring as a necessary compensation the reduction of some ad- joining part.