XVIII. On the Law which has regulated the Introduction of New Species. By ALFRED R. WALLACE, F.R.G.S.

EVERY naturalist who has directed his attention to the subject of the geographical distribution of animals and plants, must have been interested in the singular facts which it presents. Many of these facts are quite different from what would have been anticipated, and have hitherto been considered as highly curious, but quite inexplicable. None of the explanations attempted from the time of Linnaeus are now considered at all satisfactory; none of them have given a cause sufficient to account for the facts known at the time, or comprehensive enough to include all the new facts which have since been, and are daily being added. Of late years, however, a great light has been thrown upon the subject by geological investigations, which have shown that the present state of the earth, and the organisms now inhabiting it, are but the last stage of a long and uninterrupted series of changes which it has undergone, and conse quently, that to endeavour to explain and account for its present condition without any reference to those changes (as has frequently been done) must lead to very imperfect and erroneous conclusions.

The facts proved by geology are briefly these :-That during an immense, but unknown period, the surface of the earth has undergone successive changes; land has sunk beneath the ocean, while fresh land has risen up from it; mountain chains have been elevated; islands have been formed into continents, and continents submerged till they have become islands; and these changes have taken place, not once merely, but perhaps hundreds, perhaps thousands of times :-That all these operations have been more or less continuous, but unequal in their progress, and during the whole series the organic life of the earth has undergone a corresponding alteration. This alteration also has been gradual, but complete; after a certain interval not a single species existing which had lived at the commencement of the period. This complete renewal of the forms of life also appears to have occurred several times :-That from the last of the Geological epochs to the present or Historical epoch, the change of organic life has been gradual: the first appearance of animals now existing can in many cases be traced, their numbers gradually increasing in the more recent formations, while other species. continually die out and disappear, so that the present condition of the organic world is clearly derived by a natural process of gradual extinction and creation of species from that of the latest geological periods. We may therefore safely infer a like gradation and natural sequence from one geological epoch to another.

Now, taking this as a fair statement of the results of geological inquiry, we see that the present geographical distribution of life upon the earth must be the result of all the previous changes, both of the surface of the earth itself and of its inhabitants. Many causes no doubt have operated of which we must ever remain in ignorance, and we may therefore expect to find many details very difficult of explanation, and in attempting to give one, must allow ourselves to call into our service geological changes which it is highly probable may have occurred, though we have no direct evidence of their individual operation.

The great increase of our knowledge within the last twenty years, both of the present and past history of the organic world, has accumulated a body of facts which should afford a sufficient foundation for a comprehensive law embracing and explaining them all, and giving a direction to new researches. It is about ten years since the idea of such a law suggested itself to the writer of this paper, and he has since taken every opportunity of testing it by all the newly ascertained facts with which he has become acquainted, or has been able to observe himself. These have all served to convince him of the correctness of his hypothesis. Fully to enter into such a subject would occupy much space, and it is only in consequence of some views having been lately promulgated, he believes in a wrong direction, that he now ventures to present his ideas to the public, with only such obvious illustrations of the arguments and results as occur to him in a place far removed from all means of reference and exact information.

The following propositions in Organic Geography and Geology give the main facts on which the hypothesis is founded.

Geography.

1. Large groups, such as classes and orders, are generally spread over the whole earth, while smaller ones, such as families and genera, are frequently confined to one portion, often to a very limited district.

2. In' widely distributed families the genera are often limited in range; in widely distributed genera, well-marked groups of species are peculiar to each geographical district.

3. When a group is confined to one district, and is rich in species, it is almost invariably the case that the most closely allied species are found in the same locality or in closely adjoining localities, and that therefore the natural sequence of the species by affinity is also geographical.

4. In countries of a similar climate, but separated by a wide sea or lofty mountains, the families, genera and species of the Ann. & Mag. N. Hist. Ser. 2. Vol. xvi.

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one are often represented by closely allied families, genera and species peculiar to the other.

Geology.

5. The distribution of the organic world in time is very similar to its present distribution in space.

6. Most of the larger and some small groups extend through several geological periods.

7. In each period, however, there are peculiar groups, found nowhere else, and extending through one or several formations. 8. Species of one genus, or genera of one family occurring in the same geological time are more closely allied than those separated in time.

9. As generally in geography no species or genus occurs in two very distant localities without being also found in intermediate places, so in geology the life of a species or genus has not been interrupted. In other words, no group or species has come into existence twice.

10. The following law may be deduced from these facts:-Every species has come into existence coincident both in space and time with a pre-existing closely allied species.

This law agrees with, explains and illustrates all the facts connected with the following branches of the subject:-1st. The system of natural affinities. 2nd. The distribution of animals and plants in space. 3rd. The same in time, including all the phænomena of representative groups, and those which Professor Forbes supposed to manifest polarity. 4th. The phænomena of rudimentary organs. We will briefly endeavour to show its bearing upon each of these.

If the law above enunciated be true, it follows that the natural series of affinities will also represent the order in which the several species came into existence, each one having had for its immediate antitype a closely allied species existing at the time of its origin. It is evidently possible that two or three distinct species may have had a common antitype, and that each of these may again have become the antitypes from which other closely allied species were created. The effect of this would be, that so long as each species has had but one new species formed on its model, the line of affinities will be simple, and may be represented by placing the several species in direct succession in a straight line. But if two or more species have been independently formed on the plan of a common antitype, then the series of affinities will be compound, and can only be represented by a forked or many-branched line. Now, all attempts at a Natural classification and arrangement of organic beings show, that both

these plans have obtained in creation. Sometimes the series of affinities can be well represented for a space by a direct progression from species to species or from group to group, but it is generally found impossible so to continue. There constantly occur two or more modifications of an organ or modifications of two distinct organs, leading us on to two distinct series of species, which at length differ so much from each other as to form 'distinct genera or families. These are the parallel series or representative groups of naturalists, and they often occur in different countries, or are found fossil in different formations. They are said to have an analogy to each other when they are so far removed from their common antitype as to differ in many important points of structure, while they still preserve a family resemblance. We thus see how difficult it is to determine in every case whether a given relation is an analogy or an affinity, for it is evident that as we go back along the parallel or divergent series, towards the common antitype, the analogy which existed between the two groups becomes an affinity. We are also made aware of the difficulty of arriving at a true classification, even in a small and perfect group;-in the actual state of nature it is almost impossible, the species being so numerous and the modifications of form and structure so varied, arising probably from the immense number of species which have served as antitypes for the existing species, and thus produced a complicated branching of the lines of affinity, as intricate as the twigs of a gnarled oak or the vascular system of the human body. Again, if we consider that we have only fragments of this vast system, the stem and main branches being represented by extinct species of which we have no knowledge, while a vast mass of limbs and boughs and minute twigs and scattered leaves is what we have to place in order, and determine the true position each originally occupied with regard to the others, the whole difficulty of the true Natural System of classification becomes apparent to us.

We shall thus find ourselves obliged to reject all those systems of classification which arrange species or groups in circles, as well as those which fix a definite number for the divisions of each group. The latter class have been very generally rejected by naturalists, as contrary to nature, notwithstanding the ability with which they have been advocated; but the circular system of affinities seems to have obtained a deeper hold, many eminent naturalists having to some extent adopted it. We have, however, never been able to find a case in which the circle has been closed by a direct and close affinity. In most cases a palpable analogy has been substituted, in others the affinity is very obscure or altogether doubtful. The complicated branching of the lines of affinities in extensive groups must also afford great

facilities for giving a show of probability to any such purely artificial arrangements. Their death-blow was given by the admirable paper of the lamented Mr. Strickland, published in the Annals of Natural History,' in which he so clearly showed the true synthetical method of discovering the Natural System. If we now consider the geographical distribution of animals and plants upon the earth, we shall find all the facts beautifully in accordance with, and readily explained by, the present hypothesis. A country having species, genera, and whole families peculiar to it, will be the necessary result of its having been isolated for a long period, sufficient for many series of species to have been created on the type of pre-existing ones, which, as well as many of the earlier-formed species, have become extinct, and thus made the groups appear isolated. If in any case the antitype had an extensive range, two or more groups of species might have been formed, each varying from it in a different manner, and thus producing several representative or analogous groups. The Sylviade of Europe and the Sylvicolide of North America, the Heliconide of South America and the Euplaas of the East, the group of Trogons inhabiting Asia, and that peculiar to South America, are examples that may be accounted for in this manner,

Such phænomena as are exhibited by the Galapagos Islands, which contain little groups of plants and animals peculiar to themselves, but most nearly allied to those of South America, have not hitherto received any, even a conjectural explanation. The Galapagos are a volcanic group of high antiquity, and have probably never been more closely connected with the continent than they are at present. They must have been first peopled, like other newly-formed islands, by the action of winds and currents, and at a period sufficiently remote to have had the original species die ont, and the modified prototypes only remain. In the same way we can account for the separate islands having each their peculiar species, either on the supposition that the same original emigration peopled the whole of the islands with the same species from which differently modified prototypes were created, or that the islands were successively peopled from each other, but that new species have been created in each on the plan of the pre-existing ones. St. Helena is a similar case of a very ancient island having obtained an entirely peculiar, though limited, flora. On the other hand, no example is known of an island which can be proved geologically to be of very recent origin (late in the Tertiary, for instance), and yet possesses generic or family groups, or even many species peculiar to itself.

When a range of mountains has attained a great elevation, and has so remained during a long geological period, the species