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Origin of Species
chapter x. on the imperfection of the geological record   On the sudden appearance of whole groups of allied species
Charles Darwin
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       The abrupt manner in which whole groups of species suddenly appear in certain formations, has been urged by several palaeontologists--for instance, by Agassiz, Pictet, and Sedgwick, as a fatal objection to the belief in the transmutation of species. If numerous species, belonging to the same genera or families, have really started into life at once, the fact would be fatal to the theory of evolution through natural selection. For the development by this means of a group of forms, all of which are descended from some one progenitor, must have been an extremely slow process; and the progenitors must have lived long before their modified descendants. But we continually overrate the perfection of the geological record, and falsely infer, because certain genera or families have not been found beneath a certain stage, that they did not exist before that stage. In all cases positive palaeontological evidence may be implicitly trusted; negative evidence is worthless, as experience has so often shown. We continually forget how large the world is, compared with the area over which our geological formations have been carefully examined; we forget that groups of species may elsewhere have long existed, and have slowly multiplied, before they invaded the ancient archipelagoes of Europe and the United States. We do not make due allowance for the enormous intervals of time which have elapsed between our consecutive formations, longer perhaps in many cases than the time required for the accumulation of each formation. These intervals will have given time for the multiplication of species from some one parent-form: and in the succeeding formation, such groups or species will appear as if suddenly created.
       I may here recall a remark formerly made, namely, that it might require a long succession of ages to adapt an organism to some new and peculiar line of life, for instance, to fly through the air; and consequently that the transitional forms would often long remain confined to some one region; but that, when this adaptation had once been effected, and a few species had thus acquired a great advantage over other organisms, a comparatively short time would be necessary to produce many divergent forms, which would spread rapidly and widely throughout the world. Professor Pictet, in his excellent Review of this work, in commenting on early transitional forms, and taking birds as an illustration, cannot see how the successive modifications of the anterior limbs of a supposed prototype could possibly have been of any advantage. But look at the penguins of the Southern Ocean; have not these birds their front limbs in this precise intermediate state of "neither true arms nor true wings?" Yet these birds hold their place victoriously in the battle for life; for they exist in infinite numbers and of many kinds. I do not suppose that we here see the real transitional grades through which the wings of birds have passed; but what special difficulty is there in believing that it might profit the modified descendants of the penguin, first to become enabled to flap along the surface of the sea like the logger-headed duck, and ultimately to rise from its surface and glide through the air?
       I will now give a few examples to illustrate the foregoing remarks, and to show how liable we are to error in supposing that whole groups of species have suddenly been produced. Even in so short an interval as that between the first and second editions of Pictet's great work on Palaeontology, published in 1844-46 and in 1853-57, the conclusions on the first appearance and disappearance of several groups of animals have been considerably modified; and a third edition would require still further changes. I may recall the well-known fact that in geological treatises, published not many years ago, mammals were always spoken of as having abruptly come in at the commencement of the tertiary series. And now one of the richest known accumulations of fossil mammals belongs to the middle of the secondary series; and true mammals have been discovered in the new red sandstone at nearly the commencement of this great series. Cuvier used to urge that no monkey occurred in any tertiary stratum; but now extinct species have been discovered in India, South America and in Europe, as far back as the miocene stage. Had it not been for the rare accident of the preservation of footsteps in the new red sandstone of the United States, who would have ventured to suppose that no less than at least thirty different bird-like animals, some of gigantic size, existed during that period? Not a fragment of bone has been discovered in these beds. Not long ago, palaeontologists maintained that the whole class of birds came suddenly into existence during the eocene period; but now we know, on the authority of Professor Owen, that a bird certainly lived during the deposition of the upper greensand; and still more recently, that strange bird, the Archeopteryx, with a long lizard-like tail, bearing a pair of feathers on each joint, and with its wings furnished with two free claws, has been discovered in the oolitic slates of Solenhofen. Hardly any recent discovery shows more forcibly than this how little we as yet know of the former inhabitants of the world.
       I may give another instance, which, from having passed under my own eyes has much struck me. In a memoir on Fossil Sessile Cirripedes, I stated that, from the large number of existing and extinct tertiary species; from the extraordinary abundance of the individuals of many species all over the world, from the Arctic regions to the equator, inhabiting various zones of depths, from the upper tidal limits to fifty fathoms; from the perfect manner in which specimens are preserved in the oldest tertiary beds; from the ease with which even a fragment of a valve can be recognised; from all these circumstances, I inferred that, had sessile cirripedes existed during the secondary periods, they would certainly have been preserved and discovered; and as not one species had then been discovered in beds of this age, I concluded that this great group had been suddenly developed at the commencement of the tertiary series. This was a sore trouble to me, adding, as I then thought, one more instance of the abrupt appearance of a great group of species. But my work had hardly been published, when a skilful palaeontologist, M. Bosquet, sent me a drawing of a perfect specimen of an unmistakable sessile cirripede, which he had himself extracted from the chalk of Belgium. And, as if to make the case as striking as possible, this cirripede was a Chthamalus, a very common, large, and ubiquitous genus, of which not one species has as yet been found even in any tertiary stratum. Still more recently, a Pyrgoma, a member of a distinct subfamily of sessile cirripedes, has been discovered by Mr. Woodward in the upper chalk; so that we now have abundant evidence of the existence of this group of animals during the secondary period.
       The case most frequently insisted on by palaeontologists of the apparently sudden appearance of a whole group of species, is that of the teleostean fishes, low down, according to Agassiz, in the Chalk period. This group includes the large majority of existing species. But certain Jurassic and Triassic forms are now commonly admitted to be teleostean; and even some palaeozoic forms have thus been classed by one high authority. If the teleosteans had really appeared suddenly in the northern hemisphere at the commencement of the chalk formation, the fact would have been highly remarkable; but it would not have formed an insuperable difficulty, unless it could likewise have been shown that at the same period the species were suddenly and simultaneously developed in other quarters of the world. It is almost superfluous to remark that hardly any fossil-fish are known from south of the equator; and by running through Pictet's Palaeontology it will be seen that very few species are known from several formations in Europe. Some few families of fish now have a confined range; the teleostean fishes might formerly have had a similarly confined range, and after having been largely developed in some one sea, have spread widely. Nor have we any right to suppose that the seas of the world have always been so freely open from south to north as they are at present. Even at this day, if the Malay Archipelago were converted into land, the tropical parts of the Indian Ocean would form a large and perfectly enclosed basin, in which any great group of marine animals might be multiplied; and here they would remain confined, until some of the species became adapted to a cooler climate, and were enabled to double the southern capes of Africa or Australia, and thus reach other and distant seas.
       >From these considerations, from our ignorance of the geology of other countries beyond the confines of Europe and the United States, and from the revolution in our palaeontological knowledge effected by the discoveries of the last dozen years, it seems to me to be about as rash to dogmatize on the succession of organic forms throughout the world, as it would be for a naturalist to land for five minutes on a barren point in Australia, and then to discuss the number and range of its productions.
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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