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Origin of Species
chapter x. on the imperfection of the geological record   On the poorness of our palaeontological collections
Charles Darwin
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       Now let us turn to our richest museums, and what a paltry display we behold! That our collections are imperfect is admitted by every one. The remark of that admirable palaeontologist, Edward Forbes, should never be forgotten, namely, that very many fossil species are known and named from single and often broken specimens, or from a few specimens collected on some one spot. Only a small portion of the surface of the earth has been geologically explored, and no part with sufficient care, as the important discoveries made every year in Europe prove. No organism wholly soft can be preserved. Shells and bones decay and disappear when left on the bottom of the sea, where sediment is not accumulating. We probably take a quite erroneous view, when we assume that sediment is being deposited over nearly the whole bed of the sea, at a rate sufficiently quick to embed and preserve fossil remains. Throughout an enormously large proportion of the ocean, the bright blue tint of the water bespeaks its purity. The many cases on record of a formation conformably covered, after an immense interval of time, by another and later formation, without the underlying bed having suffered in the interval any wear and tear, seem explicable only on the view of the bottom of the sea not rarely lying for ages in an unaltered condition. The remains which do become embedded, if in sand or gravel, will, when the beds are upraised, generally be dissolved by the percolation of rain water charged with carbonic acid. Some of the many kinds of animals which live on the beach between high and low water mark seem to be rarely preserved. For instance, the several species of the Chthamalinae (a sub-family of sessile cirripedes) coat the rocks all over the world in infinite numbers: they are all strictly littoral, with the exception of a single Mediterranean species, which inhabits deep water and this has been found fossil in Sicily, whereas not one other species has hitherto been found in any tertiary formation: yet it is known that the genus Chthamalus existed during the Chalk period. Lastly, many great deposits, requiring a vast length of time for their accumulation, are entirely destitute of organic remains, without our being able to assign any reason: one of the most striking instances is that of the Flysch formation, which consists of shale and sandstone, several thousand, occasionally even six thousand feet in thickness, and extending for at least 300 miles from Vienna to Switzerland; and although this great mass has been most carefully searched, no fossils, except a few vegetable remains, have been found.
       With respect to the terrestrial productions which lived during the Secondary and Palaeozoic periods, it is superfluous to state that our evidence is fragmentary in an extreme degree. For instance, until recently not a land-shell was known belonging to either of these vast periods, with the exception of one species discovered by Sir C. Lyell and Dr. Dawson in the carboniferous strata of North America; but now land-shells have been found in the lias. In regard to mammiferous remains, a glance at the historical table published in Lyell's Manual, will bring home the truth, how accidental and rare is their preservation, far better than pages of detail. Nor is their rarity surprising, when we remember how large a proportion of the bones of tertiary mammals have been discovered either in caves or in lacustrine deposits; and that not a cave or true lacustrine bed is known belonging to the age of our secondary or palaeozoic formations.
       But the imperfection in the geological record largely results from another and more important cause than any of the foregoing; namely, from the several formations being separated from each other by wide intervals of time. This doctrine has been emphatically admitted by many geologists and palaeontologists, who, like E. Forbes, entirely disbelieve in the change of species. When we see the formations tabulated in written works, or when we follow them in nature, it is difficult to avoid believing that they are closely consecutive. But we know, for instance, from Sir R. Murchison's great work on Russia, what wide gaps there are in that country between the superimposed formations; so it is in North America, and in many other parts of the world. The most skilful geologist, if his attention had been confined exclusively to these large territories, would never have suspected that during the periods which were blank and barren in his own country, great piles of sediment, charged with new and peculiar forms of life, had elsewhere been accumulated. And if, in every separate territory, hardly any idea can be formed of the length of time which has elapsed between the consecutive formations, we may infer that this could nowhere be ascertained. The frequent and great changes in the mineralogical composition of consecutive formations, generally implying great changes in the geography of the surrounding lands, whence the sediment was derived, accord with the belief of vast intervals of time having elapsed between each formation.
       We can, I think, see why the geological formations of each region are almost invariably intermittent; that is, have not followed each other in close sequence. Scarcely any fact struck me more when examining many hundred miles of the South American coasts, which have been upraised several hundred feet within the recent period, than the absence of any recent deposits sufficiently extensive to last for even a short geological period. Along the whole west coast, which is inhabited by a peculiar marine fauna, tertiary beds are so poorly developed that no record of several successive and peculiar marine faunas will probably be preserved to a distant age. A little reflection will explain why, along the rising coast of the western side of South America, no extensive formations with recent or tertiary remains can anywhere be found, though the supply of sediment must for ages have been great, from the enormous degradation of the coast rocks and from the muddy streams entering the sea. The explanation, no doubt, is that the littoral and sub-littoral deposits are continually worn away, as soon as they are brought up by the slow and gradual rising of the land within the grinding action of the coast-waves.
       We may, I think, conclude that sediment must be accumulated in extremely thick, solid, or extensive masses, in order to withstand the incessant action of the waves, when first upraised and during subsequent oscillations of level, as well as the subsequent subaerial degradation. Such thick and extensive accumulations of sediment may be formed in two ways; either in profound depths of the sea, in which case the bottom will not be inhabited by so many and such varied forms of life as the more shallow seas; and the mass when upraised will give an imperfect record of the organisms which existed in the neighbourhood during the period of its accumulation. Or sediment may be deposited to any thickness and extent over a shallow bottom, if it continue slowly to subside. In this latter case, as long as the rate of subsidence and supply of sediment nearly balance each other, the sea will remain shallow and favourable for many and varied forms, and thus a rich fossiliferous formation, thick enough, when upraised, to resist a large amount of denudation, may be formed.
       I am convinced that nearly all our ancient formations, which are throughout the greater part of their thickness rich in fossils, have thus been formed during subsidence. Since publishing my views on this subject in 1845, I have watched the progress of geology, and have been surprised to note how author after author, in treating of this or that great formation, has come to the conclusion that it was accumulated during subsidence. I may add, that the only ancient tertiary formation on the west coast of South America, which has been bulky enough to resist such degradation as it has as yet suffered, but which will hardly last to a distant geological age, was deposited during a downward oscillation of level, and thus gained considerable thickness.
       All geological facts tell us plainly that each area has undergone numerous slow oscillations of level, and apparently these oscillations have affected wide spaces. Consequently, formations rich in fossils and sufficiently thick and extensive to resist subsequent degradation, will have been formed over wide spaces during periods of subsidence, but only where the supply of sediment was sufficient to keep the sea shallow and to embed and preserve the remains before they had time to decay. On the other hand, as long as the bed of the sea remained stationary, thick deposits cannot have been accumulated in the shallow parts, which are the most favourable to life. Still less can this have happened during the alternate periods of elevation; or, to speak more accurately, the beds which were then accumulated will generally have been destroyed by being upraised and brought within the limits of the coast-action.
       These remarks apply chiefly to littoral and sublittoral deposits. In the case of an extensive and shallow sea, such as that within a large part of the Malay Archipelago, where the depth varies from thirty or forty to sixty fathoms, a widely extended formation might be formed during a period of elevation, and yet not suffer excessively from denudation during its slow upheaval; but the thickness of the formation could not be great, for owing to the elevatory movement it would be less than the depth in which it was formed; nor would the deposit be much consolidated, nor be capped by overlying formations, so that it would run a good chance of being worn away by atmospheric degradation and by the action of the sea during subsequent oscillations of level. It has, however, been suggested by Mr. Hopkins, that if one part of the area, after rising and before being denuded, subsided, the deposit formed during the rising movement, though not thick, might afterwards become protected by fresh accumulations, and thus be preserved for a long period.
       Mr. Hopkins also expresses his belief that sedimentary beds of considerable horizontal extent have rarely been completely destroyed. But all geologists, excepting the few who believe that our present metamorphic schists and plutonic rocks once formed the primordial nucleus of the globe, will admit that these latter rocks have been stripped of their covering to an enormous extent. For it is scarcely possible that such rocks could have been solidified and crystallised while uncovered; but if the metamorphic action occurred at profound depths of the ocean, the former protecting mantle of rock may not have been very thick. Admitting then that gneiss, mica-schist, granite, diorite, etc., were once necessarily covered up, how can we account for the naked and extensive areas of such rocks in many parts of the world, except on the belief that they have subsequently been completely denuded of all overlying strata? That such extensive areas do exist cannot be doubted: the granitic region of Parime is described by Humboldt as being at least nineteen times as large as Switzerland. South of the Amazon, Boue colours an area composed of rocks of this nature as equal to that of Spain, France, Italy, part of Germany, and the British Islands, all conjoined. This region has not been carefully explored, but from the concurrent testimony of travellers, the granitic area is very large: thus Von Eschwege gives a detailed section of these rocks, stretching from Rio de Janeiro for 260 geographical miles inland in a straight line; and I travelled for 150 miles in another direction, and saw nothing but granitic rocks. Numerous specimens, collected along the whole coast, from near Rio de Janeiro to the mouth of the Plata, a distance of 1,100 geographical miles, were examined by me, and they all belonged to this class. Inland, along the whole northern bank of the Plata, I saw, besides modern tertiary beds, only one small patch of slightly metamorphosed rock, which alone could have formed a part of the original capping of the granitic series. Turning to a well-known region, namely, to the United States and Canada, as shown in Professor H.D. Rogers' beautiful map, I have estimated the areas by cutting out and weighing the paper, and I find that the metamorphic (excluding the "semi-metamorphic") and granite rocks exceed, in the proportion of 19 to 12.5, the whole of the newer Palaeozoic formations. In many regions the metamorphic and granite rocks would be found much more widely extended than they appear to be, if all the sedimentary beds were removed which rest unconformably on them, and which could not have formed part of the original mantle under which they were crystallised. Hence, it is probable that in some parts of the world whole formations have been completely denuded, with not a wreck left behind.
       One remark is here worth a passing notice. During periods of elevation the area of the land and of the adjoining shoal parts of the sea will be increased and new stations will often be formed--all circumstances favourable, as previously explained, for the formation of new varieties and species; but during such periods there will generally be a blank in the geological record. On the other hand, during subsidence, the inhabited area and number of inhabitants will decrease (excepting on the shores of a continent when first broken up into an archipelago), and consequently during subsidence, though there will be much extinction, few new varieties or species will be formed; and it is during these very periods of subsidence that the deposits which are richest in fossils have been accumulated.
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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