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the extinction of the species in the intermediate regions. It cannot be denied that we are as yet very ignorant of the full extent of the various climatal and geographical changes which have affected the earth during modern periods; and such changes will obviously have greatly facilitated migration. As an example, I have attempted to show how potent has been the influence of the Glacial period on the distribution both of the same and of representative species throughout the world. We are as yet profoundly ignorant of the many occasional means of transport. With respect to distinct species of the same genus inhabiting very distant and isolated regions, as the process of modification has necessarily been slow, all the means of migration will have been possible during a very long period ; and consequently the difficulty of the wide diffusion of species of the same genus is in some degree lessened.
As on the theory of natural selection an interminable number of intermediate forms must have existed, linking together all the species in each group by gradations as fine as our present varieties, fit may be asked, Why do we not see these linking forms all around us? Why are not all organic beings biended together in an inextricable chaos? With respect to existing forms, we should remember that we have no right to expect (excepting in rare cases) to discover directly connecting links between them, but only between each and some extinct and supplanted form. Even on a wide area, which has during a long period remained continuous, and of which the climate and other conditions of life change insensibly in going from a district occupied by one species into another district occupied by a closely allied species, we have no just right to expect often to find intermediate varieties in the intermediate zone. For we have reason to believe that only a few species are undergoing change at any one period; and all changes are slowly effected. I have also shown that the intermediate varieties which will at first probably exist in the intermediate zones, will be liable to be
supplanted by the allied forms on either hand ; and the latter, from existing in greater numbers, will generally be modified and improved at a quicker rate than the intermediate varieties, which exist in lesser numbers ; so that the intermediate varieties will, in the long run, be supplanted and exterminated.
On this doctrine of the extermination of an infinitude of connecting links, between the living and extinct inhabitants of the world, and at each successive period between the extinct and still older species, why is not every geological formation charged with such links? Why does not every collection of fossil remains afford plain evidence of the gradation and mutation of the forms of life? We meet with no such evidence, and this is the most obvious and forcible of the many objections which may be urged against my theory. Why, again, do whole groups of allied species appear, though certainly they often falsely appear, to have come in suddenly on the several geological stages? Why do we not find great piles of strata beneath the Silurian system, stored with the remains of the progenitors of the Silurian groups of fossils ? For certainly on my theory such strata must somewhere have been deposited at these ancient and utterly unknown epochs in the world's history.
I can answer these questions and grave objections only on the supposition that the geological record is far more imperfect than most geologists believe. It cannot be objected that there has not beerr time "sufficient for any amount of organic change ; for the lapse of time has been so great as to be utterly inappreciable by the human intellect. The number of specimens in all our museums is absolutely as nothing compared with the countless generations of countless species which certainly have existed. We should not be able to recognise a species as the parent of any one or more species if we were to examine them ever so closely, unless we likewise possessed many of the intermediate links between their past or parent and present states ; and these many links we could hardly ever expect to
discover, owing to the imperfection of the geological record. Numerous existing doubtful forms could be named which are probably varieties ; but who will pre tend that in future ages so many fossil links will be discovered, that naturalists will be able to decide, on the common view, whether or not these doubtful forms are varieties? As long as most of the links between any two species are unknown, if any one link or intermediate variety be discovered, it will simply be classed as another and distinct species. Only a small portion of the world has been geologically explored. Only organic beings of certain classes can be preserved in a fossil condition, at least in any great number. Widely ranging species vary most, and varieties are often at
first local,- both causes rendering the discovery of o intermediate links less likely. Local varieties will not
spread into other and distant regions until they are considerably modified and improved ; and when they do
spread, if discovered in a geological formation, they ble will appear as if suddenly created there, and will be is simply classed as new species. Most formations have \been intermittent in their accumulation ; and their duration, I am inclined to believe, has been shorter than the average duration of specific forms. Successive formations are separated from each other by enormous blank intervals of time; for fossiliferous formations, thick enough to resist future degradation, can be accumulated only where much sediment is deposited on the subsiding bed of the sea. During the alternate periods of elevation and of stationary level the record will be blank. During these latter periods there will probably be more variability in the forms of life ; during periods of subsidence, more extinction.
With respect to the absence of fossiliferous formations beneath the lowest Silurian strata, I can only recur to the hypothesis given in the ninth chapter. That the geological record is imperfect all will admit; but that it is imperfect to the degree which I require, few will be inclined to admit. If we look to long enough intervals of time, geology plainly declares that
all species have changed ; and they have changed in the manner which my theory requires, for they have changed slowly and in a graduated manner. We clearly see this in the fossil remains from consecutive formations invariably being much more closely related to each other, than are the fossils from formations distant from each other in time.
Such is the sum of the several chief objections and difficulties which may justly be urged against my theory; and I have now briefly recapitulated the answers and explanations which can be given to them. I have felt these difficulties far too heavily during many years to doubt their weight. But it deserves especial notice that the more important objections relate to questions on which we are confessedly ignorant ; nor do we know how ignorant we are. We do not know all the possible transitional gradations between the simplest and the most perfect organs; it cannot be pretended that we know all the varied means of Distribution during the long lapse of years, or that we know how imperfect the Geological Record is. Grave as these several difficulties are, in my judgment they do not overthrow the theory of descent from a few created forms with subsequent modification.
Now let us turn to the other side of the argument. Under domestication we see much variability. This seems to be mainly due to the reproductive system being eminently susceptible to changes in the conditions of life, so that this system, when not rendered impotent, fails to reproduce offspring exactly like the parent-form. Variability is governed by many complex laws,-by correlation of growth, by use and disuse, and by the direct action of the physical conditions of life. There is much difficulty in ascertaining how much modification our domestic productions have undergone; but we may safely infer that the amount has been large, and that modifications can be inherited for long periods. As long as the conditions of life remain the same, we have reason to believe that a modification, which has already been inherited for many generations, may continue to be inherited for an almost infinite number of generations. On the other hand we have evidence that variability, when it has once come into play, does not wholly cease ; for new varieties are still occasionally produced by our most anciently domesticated productions.
Man does not actually produce variability ; he only unintentionally exposes organic beings to new conditions of life, and then nature acts on the organisation, and causes variability. But man can and does select the variations given to him by nature, and thus accumulate them in any desired manner. He thus adapts animals and plants for his own benefit or pleasure. He may do this methodically, or he may do it unconsciously by preserving the individuals most useful to him at the time, without any thought of altering the breed. It is certain that he can largely influence the character of a breed by selecting, in each successive generation, individual differences so slight as to be quite inappreciable by an uneducated eye. This process of selection has been the great agency in the production of the most distinct and useful domestic breeds. That many of the breeds produced by man have to a large extent the character of natural species, is shown by the inextricable doubts whether very many of them are varieties or aboriginal species.
There is no obvious reason why the principles which Whave acted so efficiontly under domestication should not have acted under nature. In the preservation of favoured individuals and races, during the constantlyrecurrent Struggle for Existence, we see the most powerful and ever-acting means of selection. The struggle for existence inevitably follows from the high geometrical ratio of increase which is common to all organic beings. This high rate of increase is proved by calculation,-by the rapid increase of many animals and plants during a succession of peculiar seasons, or when naturalised in a new country. More individuals are born than can possibly survive. A grain in the