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.

In Darwin's theory, you just have to substitute 'mutations' for his 'slight accidental variations' (just as quantum theory substitutes 'quantum jump' for 'continuous transfer of energy'). In all other respects little change was necessary in Darwin's theory...

There can be no question that Darwin had nothing like sufficient evidence to establish his theory of evolution. . . . His general theory, that all life on earth had originated and evolved by a gradual successive accumulation of fortuitous mutations, is still, as it was in Darwin’s time, a highly speculative hypothesis entirely without direct factual support and very far from that self-evident axiom some of its more aggressive advocates would have us believe. . . . One might have expected that a theory of such cardinal importance, a theory that literally changed the world, would have been something more than metaphysics, something more than a myth.

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.

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.

Trained in a less severe school than that of geometry and physics, his reasonings are almost always loose and inconclusive. His generalizations seem to have been reached before he had obtained the materials upon which he rests them: His facts, though frequently new and interesting, are often little more than conjectures; and the grand phenomena of the world of life, and instinct, and reason, which other minds have woven into noble and elevating truths, have thus become in Mr. Darwin's hands the basis of a dangerous and degrading speculation.

Matthew's response: To me the conception of this law of Nature came intuitively as a self-evident fact, almost without an effort of concentrated thought. Mr Darwin here seems to have more merit in the discovery than I have had; to me it did not appear a discovery. He seems to have worked it out by inductive reason, slowly and with due caution to have made his way synthetically from fact to fact onwards; while with me it was by a general glance at the scheme of Nature that I estimated this select production of species as an a priori recognisable fact – an axiom requiring only to be pointed out to be admitted by unprejudiced minds of sufficient grasp.

Although Darwin was an innovator, his ideas rested on a foundation of fact and analysis that naturalists and explorers around the world had built up over the course of a century or more.

A seeker after Truth cannot afford to indulge in generalisation. Darwin for the greater part of his book Origin of the Species [sic] has simply massed fact upon fact without any theorising, and only towards the end has formulated his conclusion which, because of the sheer weight of testimony behind it, becomes almost irresistible. Yes I have criticised even Darwin's generalisation as being unwarranted. Science tells us that a proposition may hold good in nine hundred ninety-nine cases and yet fail in the thousandth case and thus be rendered untenable as a universal statement.

The facts of variability, of the struggle for existence, of adaptation to conditions, were notorious enough; but none of us had suspected that the road to the heart of the species problem lay through them, until Darwin and Wallace dispelled the darkness.

Taken together, these new studies suggest that Darwin did not know the strength of his own theory. He vastly underestimated the power of natural selection. Its action is neither rare nor slow. It leads to evolution daily and hourly, all around us, and we can watch.

Without much preliminary exposition and without feeling himself to be out of touch with his contemporaries, Darwin set to work to explain species in terms of the process out of which they have arisen, rather than out of the prime cause to which the distinction between them may he due.

One of the most important presuppositions of Darwin's entire thesis is gradualism, the idea that mutations to the genome can cause minor variations in the organism's properties, which can be accumulated piecemeal, bit by bit, over the eons to create the complex order found in the organisms we observe.

While the geneticists were disproving many of Darwin's ideas, the palaeontologists were determining the actual historical facts of evolution. ...they were able to verify the law of succession, first explicitly given by Darwin's colleague Wallace. "Every species has come into existence coincident, both in time and space, with a pre-existing closely allied species."