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we now find, On an average, most varieties. Secondary sexual characters are highly variable, and such characters differ much in the species of the same group. Variability in the same parts of the organisation has generally been taken advantage of in giving secondary sexual differences to the two sexes of the same species, and specific differences to the several species of the same genus. Any part or organ developed to an extraordinary size or in an extraordinary manner, in comparison with the same part or organ in the allied species, must have gone through an extraordinary amount of modification since the genus arose; and thus we can understand why it should often still be variable in a much higher degree than other parts; for variation is a long-continued and slow process, and natural selection will in such cases not as yet have had time to overcome the tendency to further variability and to reversion to a less modified state. But when a species with any extraordinarilydeveloped organ has become the parent of many modified descendants—which on our view must be a very slow process, requiring a long lapse of time—in this case, natural selection has succeeded in giving a fixed character to the organ, in however extraordinary a manner it may have been developed. Species inheriting nearly the same constitution from a common parent, and, exr^sed_io_fijrjjilar influences, naturally tend to present analogous_variations, or these same species may occasionally revert to some of the characters of their ancient progenitors. Although new and important modifications may not arise from reversion and analogous variation, such modifications will add to the beautiful and harmonious diversity of nature.
Whatever the cause may be of each slight difference between the offspring and their parents—and a cause for each must exist—we have reason to believe that it is the steady accumulation of beneficial differences which has given rise to all the more important modifications of structure in relation to the habits of each species.
Difficulties Of The Theory.
Difficulties of the theory of descent with modification — Absence or rarity of transitional varieties — Transitions in habits of life — Diversified habits in the same species — Species with habits widely different from those of their allies — Organs of extreme perfection — Modes of transition — Cases of difficulty — Natura non facit saltum—Organs of small importance — Organs not in all cases absolutely perfect ■—F The law of Unity of Type and of the Conditions of Existence embraced by the theory of Natural Selection.
Long before the reader has arrived at this part of my work, a crowd of difficulties will have occurred to him. Some of them are so serious that to this day I can hardly reflect on them without being in some degree staggered; but, to the best of my judgment, the number are only apparent, and those that are real are greater not, I think, fatal to the theory.
These difficulties and objections may be classed under the following heads:—First, why, if species have descended from other species by fine gradations, do we not everywhere see innumerable transitional forms? Why is not all nature in confusion, instead of the species being, as we see them, well defined?
Secondly, is it possible that an animal having, for instance, the structure and habits of a bat, could have been formed by the modification of some other animal with widely-different habits and structure? Can we believe that natural selection could produce, on the one hand, an organ of trifling importance, such as the tail of a giraffe, which serves as a fly-flapper, and, on the other hand, an organ so wonderful as the eye?
Thirdly, can instincts be acquired and modified through natural selection? What shall we say to the instinct which leads the bee to make cells, and 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?
The two first heads will here be discussed; some miscellaneous objections in the following chapter; Instinct and Hybridism in the two succeeding 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-form and other less-favoured forms with which it comes into competition.
| Thus extinction and natural selection go hand in hand. Hence, if we look at each species as descended from some unknown form, both the parent and all the transitional varieties will generally have been, exterminated by the very process of the 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 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 crust of the earth is a vast museum; but the natural collections have been imperfectly made, and only at long intervals of time.
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 are 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-form 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 closelylinking 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 most continents have 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 B. 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 allimportant 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 see that the range of the inhabitants of any country by no means exclusively depends on insensibly changing physical conditions, but in a large part on the presence of other species, on which it lives, or by which it is destroyed, or with which it comes into competition; and as these species are already defined objects,,. 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 nature of the seasons, be extremely liable to utter extermination; and thus its geographical range will come to be still more sharply defined.
As allied or representative species, when inhabiting a continuous area, are generally distributed in such a manner that each has a lyide 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 take a varying species inhabiting 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 d, 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 conclude that varieties linking two other varieties together generally have existed in lesser numbers than the forms which they connect, then 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 closelyallied forms existing on both sides of it. But it is a far more important consideration, that during the process of further modification, by which two varieties are supposed to be converted and perfected into twofjflistinct speoies, the two which exist in larger numbers, from inhabiting larger areas, will have a great advantage over the intermediate variety, which exists in smaller numbers
Jin a narrow and intermediate zone. For forms existing in larger numbers will have a better chance, within any given period, of presenting further favourable variations for natural selection to seize on, than will the rarer forms which exist in lesser numbers. Hence, the more common forms, in the race for life, will tend to boat and supplant the less common forms, for these will be more