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such wonderful structure, as the eye, of which we hardly as yet fully understand the inimitable perfection? Thirdly, can instincts be acquired and modified through natural selection ? What shall we say to so marvellous an instinct as that which leads the bee to make cells, which has practically anticipated the discoveries of profound mathematicians? Fourthly, how can we account for species, when crossed, being sterile and producing sterile offspring, whereas, when varieties are crossed, their fertility is unimpaired 2 The two first heads shall be here discussed—Instinct and Hybridism in separate chapters.
On the absence or rarity of transitional varieties.— As natural selection acts solely by the preservation of profitable modifications, each new form will tend in a fully-stocked country to take the place of, and finally to exterminate, its own less improved parent or other lessfavoured forms with which it comes into competition. Thus extinction and natural selection will, as we have seen, go hand in hand. Hence, if we look at each species as descended from some other unknown form, both the parent and all the transitional varieties will generally have been exterminated by the very process of formation and perfection of the new form.
But, as by this theory innumerable transitional forms must have existed, why do we not find them embedded in countless numbers in the crust of the earth? It will be much more convenient to discuss this question in the chapter on the Imperfection of the geological record; and I will here only state that I believe the answer mainly lies in the record being incomparably less perfect than is generally supposed; the imperfection of the record being chiefly due to organic beings not inhabiting profound depths of the sea, and to their remains being embedded and preserved to a future age only in masses of sediment sufficiently thick and extensive to withstand an enormous amount of future degradation; and such fossiliferous masses can be accumulated only where much sediment is deposited on the shallow bed of the sea, whilst it slowly subsides. These contingencies will concur only rarely, and after enormously long intervals. Whilst the bed of the sea is stationary or is rising, or when very little sediment is being deposited, there will be blanks in our geological history. The crust of the earth is a vast museum ; but the natural collections have been made only at intervals of time immensely remote. But it may be urged that when several closely-allied species inhabit the same territory we surely ought to find at the present time many transitional forms. Let us take a simple case: in travelling from north to south over a continent, we generally meet at successive intervals with closely allied or representative species, evidently filling nearly the same place in the natural economy of the land. These representative species often meet and interlock; and as the one becomes rarer and rarer, the other becomes more and more frequent, till the one replaces the other. But if we compare these species where they intermingle, they are generally as absolutely distinct from each other in every detail of structure as are specimens taken from the metropolis inhabited by each. By my theory these allied species have descended from a common parent; and during the process of modification, each has become adapted to the conditions of life of its own region, and has supplanted and exterminated its original parent and all the transitional varieties between its past and present states. Hence we ought not to expect at the
present time to meet with numerous transitional varieties in each region, though they must have existed there, and may be embedded there in a fossil condition. But in the intermediate region, having intermediate conditions of life, why do we not now find closely-linking intermediate varieties? This difficulty for a long time quite confounded me. But I think it can be in large part explained. In the first place we should be extremely cautious in inferring, because an area is now continuous, that it has been continuous during a long period. Geology would lead us to believe that almost every continent has been broken up into islands even during the later tertiary periods; and in such islands distinct species might have been separately formed without the possibility of intermediate varieties existing in the intermediate zones. By changes in the form of the land and of climate, marine areas now continuous must often have existed within recent times in a far less continuous and uniform condition than at present. But I will pass over this way of escaping from the difficulty; for I believe that many perfectly defined species have been formed on strictly continuous areas; though I do not doubt that the formerly broken condition of areas now continuous has played an important part in the formation of new species, more especially with freely-crossing and wandering animals. In looking at species as they are now distributed over a wide area, we generally find them tolerably numerous over a large territory, then becoming somewhat abruptly rarer and rarer on the confines, and finally disappearing. Hence the neutral territory between two representative species is generally narrow in
comparison with the territory proper to each. We see the same fact in ascending mountains, and sometimes it is quite remarkable how abruptly, as Alph. de Candolle has observed, a common alpine species disappears. The same fact has been noticed by E. Forbes in sounding the depths of the sea with the dredge. To those who look at climate and the physical conditions of life as the all-important elements of distribution, these facts ought to cause surprise, as climate and height or depth graduate away insensibly. But when we bear in mind that almost every species, even in its metropolis, would increase immensely in numbers, were it not for other competing species; that nearly all either prey on or serve as prey for others; in short, that each organic being is either directly or indirectly related in the most important manner to other organic beings, we must see that the range of the inhabitants of any country by no means exclusively depends on insensibly changing physical conditions, but in large part on the presence of other species, on which it depends, or by which it is destroyed, or with which it comes into competition; and as these species are already defined objects (however they may have become so), not blending one into another by insensible gradations, the range of any one species, depending as it does on the range of others, will tend to be sharply defined. Moreover, each species on the confines of its range, where it exists in lessened numbers, will, during fluctuations in the number of its enemies or of its prey, or in the seasons, be extremely liable to utter extermination; and thus its geographical range will come to be still more sharply defined. If I am right in believing that allied or representative species, when inhabiting a continuous area, are generally so distributed that each has a wide range, with a comparatively narrow neutral territory between them, in which they become rather suddenly rarer and rarer; then, as varieties do not essentially differ from species,
the same rule will probably apply to both; and if we in imagination adapt a varying species to a very large area, we shall have to adapt two varieties to two large areas, and a third variety to a narrow intermediate zone. The intermediate variety, consequently, will exist in lesser numbers from inhabiting a narrow and lesser area; and practically, as far as I can make out, this rule holds good with varieties in a state of nature. I have met with striking instances of the rule in the case of varieties intermediate between well-marked varieties in the genus Balanus. And it would appear from information given me by Mr. Watson, Dr. Asa Gray, and Mr. Wollaston, that generally when varieties intermediate between two other forms occur, they are much rarer numerically than the forms which they connect. Now, if we may trust these facts and inferences, and therefore conclude that varieties linking two other varieties together have generally existed in lesser numbers than the forms which they connect, then, I think, we can understand why intermediate varieties should not endure for very long periods;–why as a general rule they should be exterminated and disappear, sooner than the forms which they originally linked together. For any form existing in lesser numbers would, as already remarked, run a greater chance of being exterminated than one existing in large numbers; and in this particular case the intermediate form would be eminently liable to the inroads of closely allied forms existing on both sides of it. But a far more important consideration, as I believe, is that, during the process of further modification, by which two varieties are supposed on my theory to be converted and perfected into two distinct species, the two which exist in larger numbers from inhabiting larger areas, will have a great advantage over the intermediate variety, which exists