Origin of Species by Charles Darwin
Chapter XIII. Geographical Distribution -- continued
On the inhabitants of oceanic islands
We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty with respect to distribution, on the view that not only all the individuals of the same species have migrated from some one area, but that allied species, although now inhabiting the most distant points, have proceeded from a single area, the birthplace of their early progenitors. I have already given my reasons for disbelieving in continental extensions within the period of existing species on so enormous a scale that all the many islands of the several oceans were thus stocked with their present terrestrial inhabitants. This view removes many difficulties, but it does not accord with all the facts in regard to the productions of islands. In the following remarks I shall not confine myself to the mere question of dispersal, but shall consider some other cases bearing on the truth of the two theories of independent creation and of descent with modification.
The species of all kinds which inhabit oceanic islands are few in number compared with those on equal continental areas: Alph. de Candolle admits this for plants, and Wollaston for insects. New Zealand, for instance, with its lofty mountains and diversified stations, extending over 780 miles of latitude, together with the outlying islands of Auckland, Campbell and Chatham, contain altogether only 960 kinds of flowering plants; if we compare this moderate number with the species which swarm over equal areas in Southwestern Australia or at the Cape of Good Hope, we must admit that some cause, independently of different physical conditions, has given rise to so great a difference in number. Even the uniform county of Cambridge has 847 plants, and the little island of Anglesea 764, but a few ferns and a few introduced plants are included in these numbers, and the comparison in some other respects is not quite fair. We have evidence that the barren island of Ascension aboriginally possessed less than half-a-dozen flowering plants; yet many species have now become naturalised on it, as they have in New Zealand and on every other oceanic island which can be named. In St. Helena there is reason to believe that the naturalised plants and animals have nearly or quite exterminated many native productions. He who admits the doctrine of the creation of each separate species, will have to admit that a sufficient number of the best adapted plants and animals were not created for oceanic islands; for man has unintentionally stocked them far more fully and perfectly than did nature.
Although in oceanic islands the species are few in number, the proportion of endemic kinds (i.e. those found nowhere else in the world) is often extremely large. If we compare, for instance, the number of endemic land-shells in Madeira, or of endemic birds in the Galapagos Archipelago, with the number found on any continent, and then compare the area of the island with that of the continent, we shall see that this is true. This fact might have been theoretically expected, for, as already explained, species occasionally arriving, after long intervals of time in the new and isolated district, and having to compete with new associates, would be eminently liable to modification, and would often produce groups of modified descendants. But it by no means follows that, because in an island nearly all the species of one class are peculiar, those of another class, or of another section of the same class, are peculiar; and this difference seems to depend partly on the species which are not modified having immigrated in a body, so that their mutual relations have not been much disturbed; and partly on the frequent arrival of unmodified immigrants from the mother-country, with which the insular forms have intercrossed. It should be borne in mind that the offspring of such crosses would certainly gain in vigour; so that even an occasional cross would produce more effect than might have been anticipated. I will give a few illustrations of the foregoing remarks: in the Galapagos Islands there are twenty-six land birds; of these twenty-one (or perhaps twenty-three) are peculiar; whereas of the eleven marine birds only two are peculiar; and it is obvious that marine birds could arrive at these islands much more easily and frequently than land-birds. Bermuda, on the other hand, which lies at about the same distance from North America as the Galapagos Islands do from South America, and which has a very peculiar soil, does not possess a single endemic land bird; and we know from Mr. J.M. Jones's admirable account of Bermuda, that very many North American birds occasionally or even frequently visit this island. Almost every year, as I am informed by Mr. E.V. Harcourt, many European and African birds are blown to Madeira; this island is inhabited by ninety-nine kinds, of which one alone is peculiar, though very closely related to a European form; and three or four other species are confined to this island and to the Canaries. So that the islands of Bermuda and Madeira have been stocked from the neighbouring continents with birds, which for long ages have there struggled together, and have become mutually co-adapted. Hence, when settled in their new homes, each kind will have been kept by the others to its proper place and habits, and will consequently have been but little liable to modification. Any tendency to modification will also have been checked by intercrossing with the unmodified immigrants, often arriving from the mother-country. Madeira again is inhabited by a wonderful number of peculiar land-shells, whereas not one species of sea-shell is peculiar to its shores: now, though we do not know how sea-shells are dispersed, yet we can see that their eggs or larvae, perhaps attached to seaweed or floating timber, or to the feet of wading birds, might be transported across three or four hundred miles of open sea far more easily than land-shells. The different orders of insects inhabiting Madeira present nearly parallel cases.
Oceanic islands are sometimes deficient in animals of certain whole classes, and their places are occupied by other classes; thus in the Galapagos Islands reptiles, and in New Zealand gigantic wingless birds, take, or recently took, the place of mammals. Although New Zealand is here spoken of as an oceanic island, it is in some degree doubtful whether it should be so ranked; it is of large size, and is not separated from Australia by a profoundly deep sea; from its geological character and the direction of its mountain ranges, the Rev. W.B. Clarke has lately maintained that this island, as well as New Caledonia, should be considered as appurtenances of Australia. Turning to plants, Dr. Hooker has shown that in the Galapagos Islands the proportional numbers of the different orders are very different from what they are elsewhere. All such differences in number, and the absence of certain whole groups of animals and plants, are generally accounted for by supposed differences in the physical conditions of the islands; but this explanation is not a little doubtful. Facility of immigration seems to have been fully as important as the nature of the conditions.
Many remarkable little facts could be given with respect to the inhabitants of oceanic islands. For instance, in certain islands not tenanted by a single mammal, some of the endemic plants have beautifully hooked seeds; yet few relations are more manifest than that hooks serve for the transportal of seeds in the wool or fur of quadrupeds. But a hooked seed might be carried to an island by other means; and the plant then becoming modified would form an endemic species, still retaining its hooks, which would form a useless appendage, like the shrivelled wings under the soldered wing-covers of many insular beetles. Again, islands often possess trees or bushes belonging to orders which elsewhere include only herbaceous species; now trees, as Alph. de Candolle has shown, generally have, whatever the cause may be, confined ranges. Hence trees would be little likely to reach distant oceanic islands; and an herbaceous plant, which had no chance of successfully competing with the many fully developed trees growing on a continent, might, when established on an island, gain an advantage over other herbaceous plants by growing taller and taller and overtopping them. In this case, natural selection would tend to add to the stature of the plant, to whatever order it belonged, and thus first convert it into a bush and then into a tree.