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09 - On the Imperfection of the Geological Record 09-01 -On the absence of intermediate varieties at the present day 10 IN the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume.

Most of them have now been discussed.

One, namely the distinctness of specific forms, and their not being blended together by innumerable transitional links, is a very obvious difficulty.

I assigned reasons why such links do not commonly occur at the present day, under the circumstances apparently most favourable for their presence, namely on an extensive and continuous area with graduated physical conditions.

I endeavoured to show, that the life of each species depends in a more important manner on the presence of other already defined organic forms, than on climate; and, therefore, that the really governing conditions of life do not graduate away quite insensibly like heat or moisture.

I endeavoured, also, to show that intermediate varieties, from existing in lesser numbers than the forms which they connect, will generally be beaten out and exterminated during the course of further modification and improvement.

The main cause, however, of innumerable intermediate links not now occurring everywhere throughout nature depends on the very process of natural selection, through which new varieties continually take the places of and exterminate their parent-forms.

But just in proportion as this process of extermination has acted on an enormous scale, so must the number of intermediate varieties, which have formerly existed on the earth, be truly enormous.

Why then is not every geological formation and every stratum full of such intermediate links? Geology assuredly does not reveal any such finely graduated organic chain; and this, perhaps, is the most obvious and gravest objection which can be urged against my theory.

The explanation lies, as I believe, in the extreme imperfection of the geological record.

In the first place it should always be borne in mind what sort of intermediate forms must, on my theory, have formerly existed.

I have found it difficult, when looking at any two species, to avoid picturing to myself, forms directly intermediate between them.

But this is a wholly false view; we should always look for forms intermediate between each species and a common but unknown progenitor; and the progenitor will generally have differed in some respects from all its modified descendants.

To give a simple illustration: the fantail and pouter pigeons have both descended from the rock-pigeon; if we possessed all the intermediate varieties which have ever existed, we should have an extremely close series between both and the rock-pigeon; but we should have no varieties directly intermediate between the fantail and pouter; none, for instance, combining a tail somewhat expanded with a crop somewhat enlarged, the characteristic features of these two breeds.

Fantail Pigeon
Fantail Pigeon

Pouter Pigeon
Pouter Pigeon

Rock Pigeon
Rock Pigeon


These two breeds, moreover, have become so much modified, that if we had no historical or indirect evidence regarding their origin, it would not have been possible to have determined from a mere comparison of their structure with that of the rock-pigeon, whether they had descended from this species or from some other allied species, such as C. oenas.

skogsduva (Columba Oenas)
skogsduva (Columba Oenas)
12 - Geographical Distribution -- continued 12-60 - Summary of the last and present chapters 10 In these chapters I have endeavoured to show, that if we make due allowance for our ignorance of the full effects of all the changes of climate and of the level of the land, which have certainly occurred within the recent period, and of other similar changes which may have occurred within the same period; if we remember how profoundly ignorant we are with respect to the many and curious means of occasional transport, a subject which has hardly ever been properly experimentised on; if we bear in mind how often a species may have ranged continuously over a wide area, and then have become extinct in the intermediate tracts, I think the difficulties in believing that all the individuals of the same species, wherever located, have descended from the same parents, are not insuperable.

And we are led to this conclusion, which has been arrived at by many naturalists under the designation of single centres of creation, by some general considerations, more especially from the importance of barriers and from the analogical distribution of sub-genera, genera, and families.

With respect to the distinct species of the same genus, which on my theory must have spread from one parent-source; if we make the same allowances as before for our ignorance, and remember that some forms of life change most slowly, enormous periods of time being thus granted for their migration, I do not think that the difficulties are insuperable; though they often are in this case, and in that of the individuals of the same species, extremely grave.

As exemplifying the effects of climatal changes on distribution, I have attempted to show how important has been the influence of the modern Glacial period, which I am fully convinced simultaneously affected the whole world, or at least great meridional belts.

As showing how diversified are the means of occasional transport, I have discussed at some little length the means of dispersal of fresh-water productions.

If the difficulties be not insuperable in admitting that in the long course of time the individuals of the same species, and likewise of allied species, have proceeded from some one source; then I think all the grand leading facts of geographical distribution are explicable on the theory of migration (generally of the more dominant forms of life), together with subsequent modification and the multiplication of new forms.

We can thus understand the high importance of barriers, whether of land or water, which separate our several zoological and botanical provinces.
island
island
13 - Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Or 13-12 - Summary 10 In this chapter I have attempted to show, that the subordination of group to group in all organisms throughout all time; that the nature of the relationship, by which all living and extinct beings are united by complex, radiating, and circuitous lines of affinities into one grand system; the rules followed and the difficulties encountered by naturalists in their classifications; the value set upon characters, if constant and prevalent, whether of high vital importance, or of the most trifling importance, or, as in rudimentary organs, of no importance; the wide opposition in value between analogical or adaptive characters, and characters of true affinity; and other such rules; all naturally follow on the view of the common parentage of those forms which are considered by naturalists as allied, together with their modification through natural selection, with its contingencies of extinction and divergence of character.

In considering this view of classification, it should be borne in mind that the element of descent has been universally used in ranking together the sexes, ages, and acknowledged varieties of the same species, however different they may be in structure.

If we extend the use of this element of descent, the only certainly known cause of similarity in organic beings, we shall understand what is meant by the natural system: it is genealogical in its attempted arrangement, with the grades of acquired difference marked by the terms varieties, species, genera, families, orders, and classes.

On this same view of descent with modification, all the great facts in Morphology become intelligible, whether we look to the same pattern displayed in the homologous organs, to whatever purpose applied, of the different species of a class; or to the homologous parts constructed on the same pattern in each individual animal and plant.

On the principle of successive slight variations, not necessarily or generally supervening at a very early period of life, and being inherited at a corresponding period, we can understand the great leading facts in Embryology; namely, the resemblance in an individual embryo of the homologous parts, which when matured will become widely different from each other in structure and function; and the resemblance in different species of a class of the homologous parts or organs, though fitted in the adult members for purposes as different as possible.

Larvae are active embryos, which have become specially modified in relation to their habits of life, through the principle of modifications being inherited at corresponding ages.

On this same principle and bearing in mind, that when organs are reduced in size, either from disuse or selection, it will generally be at that period of life when the being has to provide for its own wants, and bearing in mind how strong is the principle of inheritance the occurrence of rudimentary organs and their final abortion, present to us no inexplicable difficulties; on the contrary, their presence might have been even anticipated.

The importance of embryological characters and of rudimentary organs in classification is intelligible, on the view that an arrangement is only so far natural as it is genealogical.

Finally, the several classes of facts which have been considered in this chapter, seem to me to proclaim so plainly, that the innumerable species, genera, and families of organic beings, with which this world is peopled, have all descended, each within its own class or group, from common parents, and have all been modified in the course of descent, that I should without hesitation adopt this view, even if it were unsupported by other facts or arguments.
04 - Natural Selection 04-05 - Sexual Selection 10 Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex, so no doubt it will be under nature.

Thus it is rendered possible for the two sexes to be modified through natural selection in relation to different habits of life, as is sometimes the case; or for one sex to be modified in relation to the other sex, as commonly occurs.

This leads me to say a few words on what I have called Sexual Selection. This form of selection depends, not on a struggle for existence in relation to other organic beings or to external conditions, but on a struggle between the individuals of one sex, generally the males, for the possession of the other sex.

The result is not death to the unsuccessful competitor, but few or no offspring. Sexual selection is, therefore, less rigorous than natural selection.

Generally, the most vigorous males, those which are best fitted for their places in nature, will leave most progeny.

But in many cases, victory depends not so much on general vigor, as on having special weapons, confined to the male sex.

A hornless stag or spurless cock would have a poor chance of leaving numerous offspring.

stag
stag

cock
cock

spurs
spurs


Sexual selection, by always allowing the victor to breed, might surely give indomitable courage, length to the spur, and strength to the wing to strike in the spurred leg, in nearly the same manner as does the brutal cockfighter by the careful selection of his best cocks.

cockfight
cockfight


How low in the scale of nature the law of battle descends, I know not; male alligators have been described as fighting, bellowing, and whirling round, like Indians in a war-dance, for the possession of the females; male salmons have been observed fighting all day long; male stagbeetles sometimes bear wounds from the huge mandibles of other males; the males of certain hymenopterous insects have been frequently seen by that inimitable observer M. Fabre, fighting for a particular female who sits by, an apparently unconcerned beholder of the struggle, and then retires with the conqueror.

alligator
alligator

crocodile
crocodile

Stag Beetle
Stag Beetle

Indian
Indian


The war is, perhaps, severest between the males of polygamous animals, and these seem oftenest provided with special weapons.

The males of carnivorous animals are already well armed; though to them and to others, special means of defence may be given through means of sexual selection, as the mane of the lion, and the hooked jaw to the male salmon; for the shield may be as important for victory, as the sword or spear.

mane
mane

salmon
salmon

sword
sword

spear
spear