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Origin of Species
chapter xi. on the geological succession of organic beings   On the state of development of ancient compared with living forms
Charles Darwin
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       We have seen in the fourth chapter that the degree of differentiation and specialisation of the parts in organic beings, when arrived at maturity, is the best standard, as yet suggested, of their degree of perfection or highness. We have also seen that, as the specialisation of parts is an advantage to each being, so natural selection will tend to render the organisation of each being more specialised and perfect, and in this sense higher; not but that it may leave many creatures with simple and unimproved structures fitted for simple conditions of life, and in some cases will even degrade or simplify the organisation, yet leaving such degraded beings better fitted for their new walks of life. In another and more general manner, new species become superior to their predecessors; for they have to beat in the struggle for life all the older forms, with which they come into close competition. We may therefore conclude that if under a nearly similar climate the eocene inhabitants of the world could be put into competition with the existing inhabitants, the former would be beaten and exterminated by the latter, as would the secondary by the eocene, and the palaeozoic by the secondary forms. So that by this fundamental test of victory in the battle for life, as well as by the standard of the specialisation of organs, modern forms ought, on the theory of natural selection, to stand higher than ancient forms. Is this the case? A large majority of palaeontologists would answer in the affirmative; and it seems that this answer must be admitted as true, though difficult of proof.
       It is no valid objection to this conclusion, that certain Brachiopods have been but slightly modified from an extremely remote geological epoch; and that certain land and fresh-water shells have remained nearly the same, from the time when, as far as is known, they first appeared. It is not an insuperable difficulty that Foraminifera have not, as insisted on by Dr. Carpenter, progressed in organisation since even the Laurentian epoch; for some organisms would have to remain fitted for simple conditions of life, and what could be better fitted for this end than these lowly organised Protozoa? Such objections as the above would be fatal to my view, if it included advance in organisation as a necessary contingent. They would likewise be fatal, if the above Foraminifera, for instance, could be proved to have first come into existence during the Laurentian epoch, or the above Brachiopods during the Cambrian formation; for in this case, there would not have been time sufficient for the development of these organisms up to the standard which they had then reached. When advanced up to any given point, there is no necessity, on the theory of natural selection, for their further continued process; though they will, during each successive age, have to be slightly modified, so as to hold their places in relation to slight changes in their conditions. The foregoing objections hinge on the question whether we really know how old the world is, and at what period the various forms of life first appeared; and this may well be disputed.
       The problem whether organisation on the whole has advanced is in many ways excessively intricate. The geological record, at all times imperfect, does not extend far enough back to show with unmistakable clearness that within the known history of the world organisation has largely advanced. Even at the present day, looking to members of the same class, naturalists are not unanimous which forms ought to be ranked as highest: thus, some look at the selaceans or sharks, from their approach in some important points of structure to reptiles, as the highest fish; others look at the teleosteans as the highest. The ganoids stand intermediate between the selaceans and teleosteans; the latter at the present day are largely preponderant in number; but formerly selaceans and ganoids alone existed; and in this case, according to the standard of highness chosen, so will it be said that fishes have advanced or retrograded in organisation. To attempt to compare members of distinct types in the scale of highness seems hopeless; who will decide whether a cuttle-fish be higher than a bee--that insect which the great Von Baer believed to be "in fact more highly organised than a fish, although upon another type?" In the complex struggle for life it is quite credible that crustaceans, not very high in their own class, might beat cephalopods, the highest molluscs; and such crustaceans, though not highly developed, would stand very high in the scale of invertebrate animals, if judged by the most decisive of all trials--the law of battle. Beside these inherent difficulties in deciding which forms are the most advanced in organisation, we ought not solely to compare the highest members of a class at any two periods--though undoubtedly this is one and perhaps the most important element in striking a balance--but we ought to compare all the members, high and low, at two periods. At an ancient epoch the highest and lowest molluscoidal animals, namely, cephalopods and brachiopods, swarmed in numbers; at the present time both groups are greatly reduced, while others, intermediate in organisation, have largely increased; consequently some naturalists maintain that molluscs were formerly more highly developed than at present; but a stronger case can be made out on the opposite side, by considering the vast reduction of brachiopods, and the fact that our existing cephalopods, though few in number, are more highly organised than their ancient representatives. We ought also to compare the relative proportional numbers, at any two periods, of the high and low classes throughout the world: if, for instance, at the present day fifty thousand kinds of vertebrate animals exist, and if we knew that at some former period only ten thousand kinds existed, we ought to look at this increase in number in the highest class, which implies a great displacement of lower forms, as a decided advance in the organisation of the world. We thus see how hopelessly difficult it is to compare with perfect fairness, under such extremely complex relations, the standard of organisation of the imperfectly-known faunas of successive periods.
       We shall appreciate this difficulty more clearly by looking to certain existing faunas and floras. From the extraordinary manner in which European productions have recently spread over New Zealand, and have seized on places which must have been previously occupied by the indigenes, we must believe, that if all the animals and plants of Great Britain were set free in New Zealand, a multitude of British forms would in the course of time become thoroughly naturalized there, and would exterminate many of the natives. On the other hand, from the fact that hardly a single inhabitant of the southern hemisphere has become wild in any part of Europe, we may well doubt whether, if all the productions of New Zealand were set free in Great Britain, any considerable number would be enabled to seize on places now occupied by our native plants and animals. Under this point of view, the productions of Great Britain stand much higher in the scale than those of New Zealand. Yet the most skilful naturalist, from an examination of the species of the two countries, could not have foreseen this result.
       Agassiz and several other highly competent judges insist that ancient animals resemble to a certain extent the embryos of recent animals belonging to the same classes; and that the geological succession of extinct forms is nearly parallel with the embryological development of existing forms. This view accords admirably well with our theory. In a future chapter I shall attempt to show that the adult differs from its embryo, owing to variations having supervened at a not early age, and having been inherited at a corresponding age. This process, whilst it leaves the embryo almost unaltered, continually adds, in the course of successive generations, more and more difference to the adult. Thus the embryo comes to be left as a sort of picture, preserved by nature, of the former and less modified condition of the species. This view may be true, and yet may never be capable of proof. Seeing, for instance, that the oldest known mammals, reptiles, and fishes strictly belong to their proper classes, though some of these old forms are in a slight degree less distinct from each other than are the typical members of the same groups at the present day, it would be vain to look for animals having the common embryological character of the Vertebrata, until beds rich in fossils are discovered far beneath the lowest Cambrian strata--a discovery of which the chance is small.
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Introduction
chapter i. variation under domestication
   Causes of Variability
   Effects of Habit and the use or disuse of Parts; Correlated Variation; Inheritance
   Character of Domestic Varieties; Difficulty of distinguishing between Varieties and Species; Origin of Domestic Varieties from one or more Species
   Breeds of the Domestic Pigeon, Their Differences and Origin
   Principles of Selection, anciently followed, their Effects
   Unconscious Selection
   Circumstances favourable to Man's power of Selection
chapter ii. variation under nature
   Variability
   Individual Differences
   Doubtful species
   Wide ranging, much diffused, and common species, vary most
   Species of the larger genera in each country vary more frequently than the species of the smaller genera
   Many of the species of the larger genera resemble varieties in being very closely, but unequally, related to each other, and in having restricted ranges.
   Summary
chapter iii. struggle for existence
   Its bearing on natural selection
   The term, Struggle for Existence, used in a large sense
   Geometrical ratio of increase
   Nature of the checks to increase
   Complex relations of all animals and plants to each other in the struggle for existence
   Struggle for life most severe between individuals and varieties of the same species
chapter iv. natural selection; or the survival of the fittest
   Natural Selection
   Sexual Selection
   Illustrations of the action of Natural Selection, or the survival of the fittest
   On the Intercrossing of Individuals
   Circumstances favourable for the production of new forms through Natural Selection
   Extinction caused by Natural Selection
   Divergence of Character
   The Probable Effects of the Action of Natural Selection through Divergence of Character and Extinction, on the Descendants of a Common Ancestor
   On the degree to which Organisation tends to advance
   Convergence of character
   Summary
chapter v. laws of variation
   Effects of changed conditions
   Effects of the increased use and disuse of parts, as controlled by Natural Selection
   Acclimatisation
   Correlated variation
   Compensation and economy of growth
   Multiple, rudimentary, and lowly organised structures are variable
   A part developed in any species in an extraordinary degree or manner, in comparison with the same part in allied species, tends to be highly variable
   Specific characters more variable than generic characters
   Secondary sexual characters variable
   Distinct species present analogous variations, so that a variety of one species often assumes a character proper to an allied species, or reverts to some of the characters of an early progenitor
   Summary
chapter vi. difficulties of the theory
   Difficulties of the theory of descent with modification
   Absence or rarity of transitional varieties
   On the origin and transition of organic beings with peculiar habits and structure
   Organs of extreme perfection and complication
   Modes of transition
   Special difficulties of the theory of Natural Selection
   Organs of little apparent importance, as affected by Natural Selection
   Utilitarian doctrine, how far true: Beauty, how acquired
   Summary
chapter vii
   Miscellaneous Objections to the Theory of Natural Selection
chapter viii. instinct
   Instincts comparable with habits, but different in their origin
   Inherited changes of habit or instinct in domesticated animals
   Special instincts; Instincts of the cuckoo
   Slave-making instinct
   Cell-making instinct of the hive-bee
   Objections to the theory of natural selection as applied to instincts: neuter and sterile insects
   Summary
chapter ix. hybridism
   Distinction between the sterility of first crosses and of hybrids
   Degrees of sterility
   Laws governing the sterility of first crosses and of hybrids
   Origin and causes of the sterility of first crosses and of hybrids
   Reciprocal dimorphism and trimorphism
   Fertility of varieties when crossed and of their mongrel offspring not universal
   Hybrids and mongrels compared independently of their fertility
   Summary of Chapter
chapter x. on the imperfection of the geological record
   On the absence of intermediate varieties at the present day
   On the lapse of time, as inferred from the rate of denudation and of deposition
   On the poorness of our palaeontological collections
   On the absence of numerous intermediate varieties in any single formation
   On the sudden appearance of whole groups of allied species
   On the sudden appearance of groups of allied species in the lowest known fossiliferous strata
chapter xi. on the geological succession of organic beings
   On the slow and successive appearance of new species
   On extinction
   On the forms of life changing almost simultaneously throughout the world
   On the affinities of extinct species to each other and to living species
   On the state of development of ancient compared with living forms
   On the succession of the same types within the same areas, during the later Tertiary Periods.
   Summary of preceding and present chapter
chapter xii. geographical distribution
   Present distribution cannot be accounted for by differences in physical conditions
   Single centres of supposed creation
   Means of dispersal
   Dispersal during the Glacial period
   Alternate Glacial periods in the north and south
chapter xiii. geographical distribution -- continued
   Distribution of fresh-water productions
   On the inhabitants of oceanic islands
   Absence of Batrachians and terrestrial Mammals on oceanic islands
   On the relation of the inhabitants of islands to those of the nearest mainland
   Summary of the last and present chapter
chapter xiv. mutual affinities of organic beings: morphology -- embryology -- rudimentary organs
   Classification
   Analogical resemblances
   On the nature of the affinities connecting organic beings
   Morphology
   Development and embryology
   Rudimentary, atrophied, and aborted organs
   Summary
chapter xv
   Recapitulation and Conclusion
Glossary of Scientific Terms