Read On The Origin Of Species Online
Authors: Charles Darwin
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 frequently blown to sea and perish; that the beetles in Madeira, as observed by Mr. Wollaston, lie much concealed, until the wind lulls and the sun shines; that the proportion of wingless beetles is larger on the exposed 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 individual 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 lepidoptera, must habitually use their wings to gain their subsistence, have, as Mr. Wollaston suspects, their wings not at all reduced, but even enlarged. This is quite compatible 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 individuals 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 subterranean 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 certainly in this condition, the cause, as appeared on dissection, 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 eyelids and growth of fur over them, might in such case be an advantage; and if so, natural selection would constantly 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 imagine 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 cave-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 selection 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 animals having been separately created for the American and European caverns, close similarity in their organisation and affinities might have been expected; but, as Schiodte 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 gradation of habit; for, as Schiodte remarks, "animals not far remote from ordinary forms, prepare the transition from light to darkness. Next follow those that are 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 recesses, disuse will on this view have more or less perfectly obliterated its eyes, and natural selection will often have effected other changes, such as an increase in the length of the antennae or palpi, as a compensation for blindness. Notwithstanding such modifications, we might expect still to see in the cave-animals of America, affinities to the other inhabitants of that continent, 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 surrounding 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 surprise 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, I am only surprised that more wrecks of ancient life have not been preserved, owing to the less severe competition 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 requisite for seeds to germinate, in the time of sleep, etc., and this leads me to say a few words on acclimatisation. 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 predict 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 competition 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 rhododendrons, 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 conversely; but we do not positively know that these animals 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 originally chosen by uncivilised man because they were useful 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 withstanding 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 foregoing argument too far, on account of the probable origin of some of our domestic animals from several 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 living 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 constitution, 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 Encyclopaedias of China, to be very cautious in transposing animals from one district to another; for it is not likely that man should have succeeded 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 recommended 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.
On the whole, I think we may conclude that habit, use, and disuse, have, in some cases, played a considerable 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 selection 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 organisation 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 even 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.