It [variation and natural selection] would explain, says Mr. Darwin, the unity of type which runs through the whole organic world, and why there is sometimes a fundamental agreement in structure in the same class of beings which is quite independent of their habits of life, for such structure, derived by inheritance from a remote progenitor, has been modified, in the course of ages, in different ways, according to the conditions of existence. It would also explain why all living and extinct beings are united, by complex radiating and circuitous lines of affinity with one another, into one grand system; also, there having been a continued extinction of old races and species in progress, and a formation of new ones by variation, why in some genera which are largely represented, or to which a great many species belong, many of these are closely but unequally related; also, why there are distinct geographical provinces of species of animals and plants, for, after long isolation by physical barriers, each fauna and flora, by varying continually, must become distinct from its ancestral type, and from the new forms assumed by other descendants which have diverged from the same stock.
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As breeders of domestic animals, when they choose certain varieties in preference to others to breed from, speak technically of their method as that of 'selecting,' Mr. Darwin calls the combination of natural causes, which may enable certain varieties of wild animals or plants to prevail over others of the same species, 'natural selection.'
In Darwin's time no serious attempt had been made to examine the manifestations of variability. A vast assemblage of miscellaneous facts could formerly be adduced as seemingly comparable illustrations of the phenomenon "Variation." Time has shown this mass of evidence to be capable of analysis. When first promulgated it produced the impression that variability was a phenomenon generally distributed amongst living things in such a way that the specific divisions must be arbitrary. When this variability is sorted out, and is seen to be in part a result of hybridisation, in part a consequence of the persistence of hybrids by parthenogenetic reproduction, a polymorphism due to the continued presence of individuals representing various combinations of Mendelian allelomorphs, partly also the transient effect of alteration in external circumstances, we see how cautious we must be in drawing inferences as to the indefiniteness of specific limits from a bare knowledge that intermediates exist.
Darwin believed that the crossing of two types generally led to a blend, and that consequently bisexual reproduction tended to make a species uniform. He therefore had to postulate some cause constantly at work to keep up the inheritable variation within a species. He very naturally looked to the effects of differences of environment. ...It was shown that Darwin had been wrong in supposing that variations due to environment were inheritable. Selection merely picked out the best available line from a given population, and would not, as Darwin had believed, give rise to an unlimited amount of change.
It cannot be doubted, I think, that Mr. Darwin has satisfactorily proved that what he terms selection, or selective modification, must occur, and does occur, in nature; and he has also proved to superfluity that such selection is competent to produce forms as distinct, structurally, as some genera even are. If the animated world presented us with none but structural differences, I should have no hesitation in saying that Mr. Darwin has demonstrated the existence of a true physical cause, amply competent to account for the origin of living species, and of man among the rest.
As systematic inquiry into the natural facts was begun it was at once found that the accepted ideas of variation were unfounded. Variation was seen very frequently to be a definite and specific phenomenon, affecting different forms of life in different ways, but in all its diversity showing manifold and often obvious indications of regularity. This observation was not in its essence novel. Several examples of definite variation had been well known to Darwin and others, but many, especially Darwin himself in his later years, had nevertheless been disposed to depreciate the significance of such facts. They consequently then lapsed into general disparagement. Upon more careful inquiry the abundance of such phenomena proved to be far greater than was currently supposed, and a discussion of their nature brought into prominence a consideration of greater weight, namely that the differences by which these definite or discontinuous variations are constituted again and again approximate to and are comparable with the class of differences by which species are distinguished from each other.
The theory of the origin of new species by variation will also explain why a species which has once died out never reappears, and why the fossil fauna and flora recede farther and farther from the living type in proportion as we trace it back to remoter ages. It would also account for the fact, that when we have to intercalate a new set of fossiliferous strata between two groups previously known, the newly discovered fossils serve to fill up gaps between specific or generic types previously familiar to us, supplying often the missing links of the chain, which, if transmutation is accepted, must once have been continuous.
Every naturalist admits that there is a general tendency in animals and plants to vary; but it is usually taken for granted, though we have no means of proving the assumption to be true, that there are certain limits beyond which each species cannot pass under any circumstances, or in any number of generations. Mr. Darwin and Mr. Wallace say that the opposite hypothesis, which assumes that every species is capable of varying indefinitely from its original type, is not a whit more arbitrary, and has this manifest claim to be preferred, that it will account for a multitude of phenomena which the ordinary theory is incapable of explaining.
The only secondary cause... which has, as yet, been even conjecturally brought forward, to explain how, in the ordinary course of nature, a new specific form may be generated is, as Lamarck declared, 'variation,' and this has been rendered a far more probable hypothesis by the way in which Natural Selection is shown to give intensity and permanency to certain varieties.
Variation and natural selection would also afford a key to a multitude of geological facts otherwise wholly unaccounted for, as, for example, why there is generally an intimate connection between the living animals and plants of each great division of the globe and the extinct fauna and flora of the post-tertiary or tertiary formations of the same region...
Like Kant before him, Darwin insists that the source of all error is semblance. Analogy, he says again and again, is always a ‘deceitful guide’ (see pp. 61, 66, 473). As against analogy, or as I would say merely metaphorical characterizations of the facts, Darwin wishes to make a case for the existence of real ‘affinities’ genealogically construed. The establishment of these affinities will permit him to postulate the linkage of all living things to all others by the ‘laws’ or ‘principles’ of genealogical descent, variation, and natural selection. These laws and principles are the formal elements in his mechanistic explanation of why creatures are arranged in families in a time series. But this explanation could not be offered as long as the data remained encoded in the linguistic modes of either metaphor or synecdoche, the modes of qualitative connection. As long as creatures are classified in terms of either semblance or essential unity, the realm of organic things must remain either a chaos of arbitrarily affirmed connectedness or a hierarchy of higher and lower forms. Science as Darwin understood it, however, cannot deal in the categories of the ‘higher’ and ‘lower’ any more than it can deal in the categories of the ‘normal’ and ‘monstrous.’ Everything must be entertained as what it manifestly seems to be. Nothing can be regarded as ‘surprising,’ any more than anything can be regarded as ‘miraculous.’
[A]lthough species may be discrete, they have no immutable essence. Variation is the raw material of evolutionary change. It represents the fundamental reality of nature, not an accident about a created norm. Variation is primary; essences are illusory. Species must be defined as ranges of irreducible variation.
I am well aware that some very eminent systematists regard the whole problem as solved. They hold as Darwin did that specific diversity has no physiological foundation or causation apart from adaptation, and that species are impermanent groups, the delimitations of which are ultimately determined by environmental exigency or "fitness." The specific diversity of living things is thus regarded as being something quite different in nature from the specific diversity of inorganic substances. In practice those who share these opinions are, as might be anticipated, to be found among the 'lumpers' rather than among the 'splitters.' In their work, certainly, the Darwinian theory is actually followed as a guiding principle; unanalysed intergradations of all kinds are accepted as impugning the integrity of species; the underlying physiological problem is forgotten, and while the product is almost valueless as a contribution to biological research, I can scarcely suppose that it aids greatly in the advances of other branches of our science.
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