Causes of Variability. Effects of Habit. Correlation of Growth. Inheritance. Character of Domestic Varieties. Difficulty of distinguishing between Varieties and Species. Origin of Domestic Varieties from one or more Species. Domestic Pigeons, their Differences and Origin. Principle of Selection anciently followed, its Effects. Methodical and Unconscious Selection. Unknown Origin of our Domestic Productions. Circumstances favourable to Man’s power of Selection.

When we look to the individuals of ...

On The Breeds Of The Domestic Pigeon.

Believing that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every breed which I could purchase or obtain, and have been most kindly favoured with skins from several quarters of the world, more especially by the Honourable W. Elliot from India, and by the Honourable C. Murray from Persia. Many treatises in different languages have been published on pigeons, and some of them are very important, as being of considerable antiquity....

Selection.

Let us now briefly consider the steps by which domestic races have been produced, either from one or from several allied species. Some little effect may, perhaps, be attributed to the direct action of the external conditions of life, and some little to habit; but he would be a bold man who would account by such agencies for the differences of a dray and race horse, a greyhound and bloodhound, a carrier and tumbler pigeon. One of the most remarkable features in our domesticated races is that we s...

Natural Selection: its power compared with man’s selection, its power on characters of trifling importance, its power at all ages and on both sexes. Sexual Selection. On the generality of intercrosses between individuals of the same species. Circumstances favourable and unfavourable to Natural Selection, namely, intercrossing, isolation, number of individuals. Slow action. Extinction caused by Natural Selection. Divergence of Character, related to the diversity of inhabitants of any small area, ...

Sexual Selection.

Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex, the same fact probably occurs under nature, and if so, natural selection will be able to modify one sex in its functional relations to the other sex, or in relation to wholly different habits of life in the two sexes, as is sometimes the case with insects. And this leads me to say a few words on what I call Sexual Selection. This depends, not on a struggle for existence, but on a s...

Illustrations Of The Action Of Natural Selection.

In order to make it clear how, as I believe, natural selection acts, I must beg permission to give one or two imaginary illustrations. Let us take the case of a wolf, which preys on various animals, securing some by craft, some by strength, and some by fleetness; and let us suppose that the fleetest prey, a deer for instance, had from any change in the country increased in numbers, or that other prey had decreased in numbers, during that season of the year when the wolf is hardest pressed for fo...

On The Intercrossing Of Individuals.

I must here introduce a short digression. In the case of animals and plants with separated sexes, it is of course obvious that two individuals must always unite for each birth; but in the case of hermaphrodites this is far from obvious. Nevertheless I am strongly inclined to believe that with all hermaphrodites two individuals, either occasionally or habitually, concur for the reproduction of their kind. This view, I may add, was first suggested by Andrew Knight. We shall presently see its impor...

Circumstances Favourable To Natural Selection.

This is an extremely intricate subject. A large amount of inheritable and diversified variability is favourable, but I believe mere individual differences suffice for the work. A large number of individuals, by giving a better chance for the appearance within any given period of profitable variations, will compensate for a lesser amount of variability in each individual, and is, I believe, an extremely important element of success. Though nature grants vast periods of time for the work of natura...

Extinction.

This subject will be more fully discussed in our chapter on Geology; but it must be here alluded to from being intimately connected with natural selection. Natural selection acts solely through the preservation of variations in some way advantageous, which consequently endure. But as from the high geometrical powers of increase of all organic beings, each area is already fully stocked with inhabitants, it follows that as each selected and favoured form increases in number, so will the less favou...

Divergence Of Character.

The principle, which I have designated by this term, is of high importance on my theory, and explains, as I believe, several important facts. In the first place, varieties, even strongly-marked ones, though having somewhat of the character of species–as is shown by the hopeless doubts in many cases how to rank them–yet certainly differ from each other far less than do good and distinct species. Nevertheless, according to my view, varieties are species in the process of formation, or are, as I ha...

Summary Of Chapter.

If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an ...

Effects of external conditions. Use and disuse, combined with natural selection; organs of flight and of vision. Acclimatisation. Correlation of growth. Compensation and economy of growth. False correlations. Multiple, rudimentary, and lowly organised structures variable. Parts developed in an unusual manner are highly variable: specific characters more variable than generic: secondary sexual characters variable. Species of the same genus vary in an analogous manner. Reversions to long lost char...

Effects Of Use And Disuse.

From the facts alluded to in the first chapter, I think there can be little doubt that use in our domestic animals strengthens and enlarges certain parts, and disuse diminishes them; and that such modifications are inherited. Under free nature, we can have no standard of comparison, by which to judge of the effects of long-continued use or disuse, for we know not the parent-forms; but many animals have structures which can be explained by the effects of disuse. As Professor Owen has remarked, th...

Acclimatisation.

Habit is hereditary with plants, as in the period of flowering, in the amount of rain requisite for seeds to germinate, in the time of sleep, etc., and this leads me to say a few words on acclimatisation. As it is extremely common for species of the same genus to inhabit very hot and very cold countries, and as I believe that all the species of the same genus have descended from a single parent, if this view be correct, acclimatisation must be readily effected during long-continued descent. It i...

Correlation Of Growth.

I mean by this expression that the whole organisation is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified. This is a very important subject, most imperfectly understood. The most obvious case is, that modifications accumulated solely for the good of the young or larva, will, it may safely be concluded, affect the structure of the adult; in the same manner as any malcon...

Summary.

Our ignorance of the laws of variation is profound. Not in one case out of a hundred can we pretend to assign any reason why this or that part differs, more or less, from the same part in the parents. But whenever we have the means of instituting a comparison, the same laws appear to have acted in producing the lesser differences between varieties of the same species, and the greater differences between species of the same genus. The external conditions of life, as climate and food, etc., seem t...

Difficulties on the theory of descent with modification. Transitions. Absence or rarity of transitional varieties. Transitions in habits of life. Diversified habits in the same species. Species with habits widely different from those of their allies. Organs of extreme perfection. Means of transition. Cases of difficulty. Natura non facit saltum. Organs of small importance. Organs not in all cases absolutely perfect. The law of Unity of Type and of the Conditions of Existence embraced by the theo...

On The Absence Or Rarity Of Transitional Varieties.

As natural selection acts solely by the preservation of profitable modifications, each new form will tend in a fully-stocked country to take the place of, and finally to exterminate, its own less improved parent or other less-favoured forms with which it comes into competition. Thus extinction and natural selection will, as we have seen, go hand in hand. Hence, if we look at each species as descended from some other unknown form, both the parent and all the transitional varieties will generally ...

On The Origin And Transitions Of Organic Beings With Peculiar Habits And Structure.

It has been asked by the opponents of such views as I hold, how, for instance, a land carnivorous animal could have been converted into one with aquatic habits; for how could the animal in its transitional state have subsisted? It would be easy to show that within the same group carnivorous animals exist having every intermediate grade between truly aquatic and strictly terrestrial habits; and as each exists by a struggle for life, it is clear that each is well adapted in its habits to its place...

Organs Of Extreme Perfection And Complication.

To suppose that the eye, with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest possible degree. Yet reason tells me, that if numerous gradations from a perfect and complex eye to one very imperfect and simple, each grade being useful to its possessor, can be shown to exist...

Organs Of Little Apparent Importance.

As natural selection acts by life and death,–by the preservation of individuals with any favourable variation, and by the destruction of those with any unfavourable deviation of structure,–I have sometimes felt much difficulty in understanding the origin of simple parts, of which the importance does not seem sufficient to cause the preservation of successively varying individuals. I have sometimes felt as much difficulty, though of a very different kind, on this head, as in the case of an organ ...

Summary Of Chapter.

We have in this chapter discussed some of the difficulties and objections which may be urged against my theory. Many of them are very grave; but I think that in the discussion light has been thrown on several facts, which on the theory of independent acts of creation are utterly obscure. We have seen that species at any one period are not indefinitely variable, and are not linked together by a multitude of intermediate gradations, partly because the process of natural selection will always be ve...

Instincts comparable with habits, but different in their origin. Instincts graduated. Aphides and ants. Instincts variable. Domestic instincts, their origin. Natural instincts of the cuckoo, ostrich, and parasitic bees. Slave-making ants. Hive-bee, its cell-making instinct. Difficulties on the theory of the Natural Selection of instincts. Neuter or sterile insects. Summary.

The subject of instinct might have been worked into the previous chapters; but I have thought that it would be more conveni...

Slave-Making Instinct.

This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber, a better observer even than his celebrated father. This ant is absolutely dependent on its slaves; without their aid, the species would certainly become extinct in a single year. The males and fertile females do no work. The workers or sterile females, though most energetic and courageous in capturing slaves, do no other work. They are incapable of making their own nests, or of feeding their own l...

Cell-Making Instinct Of The Hive-Bee.

I will not here enter on minute details on this subject, but will merely give an outline of the conclusions at which I have arrived. He must be a dull man who can examine the exquisite structure of a comb, so beautifully adapted to its end, without enthusiastic admiration. We hear from mathematicians that bees have practically solved a recondite problem, and have made their cells of the proper shape to hold the greatest possible amount of honey, with the least possible consumption of precious wa...

Summary.

I have endeavoured briefly in this chapter to show that the mental qualities of our domestic animals vary, and that the variations are inherited. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. No one will dispute that instincts are of the highest importance to each animal. Therefore I can see no difficulty, under changing conditions of life, in natural selection accumulating slight modifications of instinct to any extent, in any useful direction. I...

Distinction between the sterility of first crosses and of hybrids. Sterility various in degree, not universal, affected by close interbreeding, removed by domestication. Laws governing the sterility of hybrids. Sterility not a special endowment, but incidental on other differences. Causes of the sterility of first crosses and of hybrids. Parallelism between the effects of changed conditions of life and crossing. Fertility of varieties when crossed and of their mongrel offspring not universal. Hy...

Laws Governing The Sterility Of First Crosses And Of Hybrids.

We will now consider a little more in detail the circumstances and rules governing the sterility of first crosses and of hybrids. Our chief object will be to see whether or not the rules indicate that species have specially been endowed with this quality, in order to prevent their crossing and blending together in utter confusion. The following rules and conclusions are chiefly drawn up from Gartner’s admirable work on the hybridisation of plants. I have taken much pains to ascertain how far the...

Causes Of The Sterility Of First Crosses And Of Hybrids.

We may now look a little closer at the probable causes of the sterility of first crosses and of hybrids. These two cases are fundamentally different, for, as just remarked, in the union of two pure species the male and female sexual elements are perfect, whereas in hybrids they are imperfect. Even in first crosses, the greater or lesser difficulty in effecting a union apparently depends on several distinct causes. There must sometimes be a physical impossibility in the male element reaching the ...

Fertility Of Varieties When Crossed, And Of Their Mongrel Offspring.

It may be urged, as a most forcible argument, that there must be some essential distinction between species and varieties, and that there must be some error in all the foregoing remarks, inasmuch as varieties, however much they may differ from each other in external appearance, cross with perfect facility, and yield perfectly fertile offspring. I fully admit that this is almost invariably the case. But if we look to varieties produced under nature, we are immediately involved in hopeless difficu...

Hybrids And Mongrels Compared, Independently Of Their Fertility.

Independently of the question of fertility, the offspring of species when crossed and of varieties when crossed may be compared in several other respects. Gartner, whose strong wish was to draw a marked line of distinction between species and varieties, could find very few and, as it seems to me, quite unimportant differences between the so-called hybrid offspring of species, and the so-called mongrel offspring of varieties. And, on the other hand, they agree most closely in very many important ...

Summary Of Chapter.

First crosses between forms sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally, sterile. The sterility is of all degrees, and is often so slight that the two most careful experimentalists who have ever lived, have come to diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible of favourable and unfavourable conditions. The degree of ...

On the absence of intermediate varieties at the present day. On the nature of extinct intermediate varieties; on their number. On the vast lapse of time, as inferred from the rate of deposition and of denudation. On the poorness of our palaeontological collections. On the intermittence of geological formations. On the absence of intermediate varieties in any one formation. On the sudden appearance of groups of species. On their sudden appearance in the lowest known fossiliferous strata.

In the s...

On The Lapse Of Time.

Independently of our not finding fossil remains of such infinitely numerous connecting links, it may be objected, that time will not have sufficed for so great an amount of organic change, all changes having been effected very slowly through natural selection. It is hardly possible for me even to recall to the reader, who may not be a practical geologist, the facts leading the mind feebly to comprehend the lapse of time. He who can read Sir Charles Lyell’s grand work on the Principles of Geology...

On The Poorness Of Our Palaeontological Collections.

That our palaeontological collections are very imperfect, is admitted by every one. The remark of that admirable palaeontologist, the late Edward Forbes, should not be forgotten, namely, that numbers of our 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 Eur...

On The Sudden Appearance Of Whole Groups Of Allied Species.

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 by none more forcibly than by Professor 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 all at once, the fact would be fatal to the theory of descent with slow modification through natural selection. For t...

On The Sudden Appearance Of Groups Of Allied Species In The Lowest Known Fossiliferous Strata.

There is another and allied difficulty, which is much graver. I allude to the manner in which numbers of species of the same group, suddenly appear in the lowest known fossiliferous rocks. Most of the arguments which have convinced me that all the existing species of the same group have descended from one progenitor, apply with nearly equal force to the earliest known species. For instance, I cannot doubt that all the Silurian trilobites have descended from some one crustacean, which must have l...

On the slow and successive appearance of new species. On their different rates of change. Species once lost do not reappear. Groups of species follow the same general rules in their appearance and disappearance as do single species. On Extinction. On simultaneous changes in the forms of life throughout the world. On the affinities of extinct species to each other and to living species. On the state of development of ancient forms. On the succession of the same types within the same areas. Summar...

On Extinction.

We have as yet spoken only incidentally of the disappearance of species and of groups of species. On the theory of natural selection the extinction of old forms and the production of new and improved forms are intimately connected together. The old notion of all the inhabitants of the earth having been swept away at successive periods by catastrophes, is very generally given up, even by those geologists, as Elie de Beaumont, Murchison, Barrande, etc., whose general views would naturally lead the...

On The Forms Of Life Changing Almost Simultaneously Throughout The World.

Scarcely any palaeontological discovery is more striking than the fact, that the forms of life change almost simultaneously throughout the world. Thus our European Chalk formation can be recognised in many distant parts of the world, under the most different climates, where not a fragment of the mineral chalk itself can be found; namely, in North America, in equatorial South America, in Tierra del Fuego, at the Cape of Good Hope, and in the peninsula of India. For at these distant points, the or...

On The Affinities Of Extinct Species To Each Other, And To Living Forms.

Let us now look to the mutual affinities of extinct and living species. They all fall into one grand natural system; and this fact is at once explained on the principle of descent. The more ancient any form is, the more, as a general rule, it differs from living forms. But, as Buckland long ago remarked, all fossils can be classed either in still existing groups, or between them. That the extinct forms of life help to fill up the wide intervals between existing genera, families, and orders, cann...

On The State Of Development Of Ancient Forms.

There has been much discussion whether recent forms are more highly developed than ancient. I will not here enter on this subject, for naturalists have not as yet defined to each other’s satisfaction what is meant by high and low forms. But in one particular sense the more recent forms must, on my theory, be higher than the more ancient; for each new species is formed by having had some advantage in the struggle for life over other and preceding forms. If under a nearly similar climate, the eoce...

On The Succession Of The Same Types Within The Same Areas, During The Later Tertiary Periods.

Mr. Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that continent. In South America, a similar relationship is manifest, even to an uneducated eye, in the gigantic pieces of armour like those of the armadillo, found in several parts of La Plata; and Professor Owen has shown in the most striking manner that most of the fossil mammals, buried there in such numbers, are related to South American types. This relationship ...

Summary Of The Preceding And Present Chapters.

I have attempted to show that the geological record is extremely imperfect; that only a small portion of the globe has been geologically explored with care; that only certain classes of organic beings have been largely preserved in a fossil state; that the number both of specimens and of species, preserved in our museums, is absolutely as nothing compared with the incalculable number of generations which must have passed away even during a single formation; that, owing to subsidence being necess...

Present distribution cannot be accounted for by differences in physical conditions. Importance of barriers. Affinity of the productions of the same continent. Centres of creation. Means of dispersal, by changes of climate and of the level of the land, and by occasional means. Dispersal during the Glacial period co-extensive with the world.

In considering the distribution of organic beings over the face of the globe, the first great fact which strikes us is, that neither the similarity nor the di...

Means Of Dispersal.

Sir C. Lyell and other authors have ably treated this subject. I can give here only the briefest abstract of the more important facts. Change of climate must have had a powerful influence on migration: a region when its climate was different may have been a high road for migration, but now be impassable; I shall, however, presently have to discuss this branch of the subject in some detail. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two mar...

Dispersal During The Glacial Period.

The identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where the Alpine species could not possibly exist, is one of the most striking cases known of the same species living at distant points, without the apparent possibility of their having migrated from one to the other. It is indeed a remarkable fact to see so many of the same plants living on the snowy regions of the Alps or Pyrenees, and in the extreme northern parts of Europ...

On The Inhabitants Of Oceanic Islands.

We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty, on the view that all the individuals both of the same and of allied species have descended from a single parent; and therefore have all proceeded from a common birthplace, notwithstanding that in the course of time they have come to inhabit distant points of the globe. I have already stated that I cannot honestly admit Forbes’s view on continental extensions, which, if le...

Summary Of Last And Present Chapters.

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 ...

CLASSIFICATION, groups subordinate to groups. Natural system. Rules and difficulties in classification, explained on the theory of descent with modification. Classification of varieties. Descent always used in classification. Analogical or adaptive characters. Affinities, general, complex and radiating. Extinction separates and defines groups. MORPHOLOGY, between members of the same class, between parts of the same individual. EMBRYOLOGY, laws of, explained by variations not supervening at an ea...

Morphology.

We have seen that the members of the same class, independently of their habits of life, resemble each other in the general plan of their organisation. This resemblance is often expressed by the term “unity of type;” or by saying that the several parts and organs in the different species of the class are homologous. The whole subject is included under the general name of Morphology. This is the most interesting department of natural history, and may be said to be its very soul. What can be more c...

Embryology.

It has already been casually remarked that certain organs in the individual, which when mature become widely different and serve for different purposes, are in the embryo exactly alike. The embryos, also, of distinct animals within the same class are often strikingly similar: a better proof of this cannot be given, than a circumstance mentioned by Agassiz, namely, that having forgotten to ticket the embryo of some vertebrate animal, he cannot now tell whether it be that of a mammal, bird, or rep...

Rudimentary, Atrophied, Or Aborted Organs.

Organs or parts in this strange condition, bearing the stamp of inutility, are extremely common throughout nature. For instance, rudimentary mammae are very general in the males of mammals: I presume that the “bastard-wing” in birds may be safely considered as a digit in a rudimentary state: in very many snakes one lobe of the lungs is rudimentary; in other snakes there are rudiments of the pelvis and hind limbs. Some of the cases of rudimentary organs are extremely curious; for instance, the pr...

Summary.

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 im...

Causes of Variability. Effects of Habit. Correlation of Growth. Inheritance. Character of Domestic Varieties. Difficulty of distinguishing between Varieties and Species. Origin of Domestic Varieties from one or more Species. Domestic Pigeons, their Differences and Origin. Principle of Selection anciently followed, its Effects. Methodical and Unconscious Selection. Unknown Origin of our Domestic Productions. Circumstances favourable to Man’s power of Selection.

When we look to the individuals of ...

On The Breeds Of The Domestic Pigeon.

Believing that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every breed which I could purchase or obtain, and have been most kindly favoured with skins from several quarters of the world, more especially by the Honourable W. Elliot from India, and by the Honourable C. Murray from Persia. Many treatises in different languages have been published on pigeons, and some of them are very important, as being of considerable antiquity....

Selection.

Let us now briefly consider the steps by which domestic races have been produced, either from one or from several allied species. Some little effect may, perhaps, be attributed to the direct action of the external conditions of life, and some little to habit; but he would be a bold man who would account by such agencies for the differences of a dray and race horse, a greyhound and bloodhound, a carrier and tumbler pigeon. One of the most remarkable features in our domesticated races is that we s...

Natural Selection: its power compared with man’s selection, its power on characters of trifling importance, its power at all ages and on both sexes. Sexual Selection. On the generality of intercrosses between individuals of the same species. Circumstances favourable and unfavourable to Natural Selection, namely, intercrossing, isolation, number of individuals. Slow action. Extinction caused by Natural Selection. Divergence of Character, related to the diversity of inhabitants of any small area, ...

Sexual Selection.

Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex, the same fact probably occurs under nature, and if so, natural selection will be able to modify one sex in its functional relations to the other sex, or in relation to wholly different habits of life in the two sexes, as is sometimes the case with insects. And this leads me to say a few words on what I call Sexual Selection. This depends, not on a struggle for existence, but on a s...

Illustrations Of The Action Of Natural Selection.

In order to make it clear how, as I believe, natural selection acts, I must beg permission to give one or two imaginary illustrations. Let us take the case of a wolf, which preys on various animals, securing some by craft, some by strength, and some by fleetness; and let us suppose that the fleetest prey, a deer for instance, had from any change in the country increased in numbers, or that other prey had decreased in numbers, during that season of the year when the wolf is hardest pressed for fo...

On The Intercrossing Of Individuals.

I must here introduce a short digression. In the case of animals and plants with separated sexes, it is of course obvious that two individuals must always unite for each birth; but in the case of hermaphrodites this is far from obvious. Nevertheless I am strongly inclined to believe that with all hermaphrodites two individuals, either occasionally or habitually, concur for the reproduction of their kind. This view, I may add, was first suggested by Andrew Knight. We shall presently see its impor...

Circumstances Favourable To Natural Selection.

This is an extremely intricate subject. A large amount of inheritable and diversified variability is favourable, but I believe mere individual differences suffice for the work. A large number of individuals, by giving a better chance for the appearance within any given period of profitable variations, will compensate for a lesser amount of variability in each individual, and is, I believe, an extremely important element of success. Though nature grants vast periods of time for the work of natura...

Extinction.

This subject will be more fully discussed in our chapter on Geology; but it must be here alluded to from being intimately connected with natural selection. Natural selection acts solely through the preservation of variations in some way advantageous, which consequently endure. But as from the high geometrical powers of increase of all organic beings, each area is already fully stocked with inhabitants, it follows that as each selected and favoured form increases in number, so will the less favou...

Divergence Of Character.

The principle, which I have designated by this term, is of high importance on my theory, and explains, as I believe, several important facts. In the first place, varieties, even strongly-marked ones, though having somewhat of the character of species–as is shown by the hopeless doubts in many cases how to rank them–yet certainly differ from each other far less than do good and distinct species. Nevertheless, according to my view, varieties are species in the process of formation, or are, as I ha...

Summary Of Chapter.

If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an ...

Effects of external conditions. Use and disuse, combined with natural selection; organs of flight and of vision. Acclimatisation. Correlation of growth. Compensation and economy of growth. False correlations. Multiple, rudimentary, and lowly organised structures variable. Parts developed in an unusual manner are highly variable: specific characters more variable than generic: secondary sexual characters variable. Species of the same genus vary in an analogous manner. Reversions to long lost char...

Effects Of Use And Disuse.

From the facts alluded to in the first chapter, I think there can be little doubt that use in our domestic animals strengthens and enlarges certain parts, and disuse diminishes them; and that such modifications are inherited. Under free nature, we can have no standard of comparison, by which to judge of the effects of long-continued use or disuse, for we know not the parent-forms; but many animals have structures which can be explained by the effects of disuse. As Professor Owen has remarked, th...

Acclimatisation.

Habit is hereditary with plants, as in the period of flowering, in the amount of rain requisite for seeds to germinate, in the time of sleep, etc., and this leads me to say a few words on acclimatisation. As it is extremely common for species of the same genus to inhabit very hot and very cold countries, and as I believe that all the species of the same genus have descended from a single parent, if this view be correct, acclimatisation must be readily effected during long-continued descent. It i...

Correlation Of Growth.

I mean by this expression that the whole organisation is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified. This is a very important subject, most imperfectly understood. The most obvious case is, that modifications accumulated solely for the good of the young or larva, will, it may safely be concluded, affect the structure of the adult; in the same manner as any malcon...

Summary.

Our ignorance of the laws of variation is profound. Not in one case out of a hundred can we pretend to assign any reason why this or that part differs, more or less, from the same part in the parents. But whenever we have the means of instituting a comparison, the same laws appear to have acted in producing the lesser differences between varieties of the same species, and the greater differences between species of the same genus. The external conditions of life, as climate and food, etc., seem t...

Difficulties on the theory of descent with modification. Transitions. Absence or rarity of transitional varieties. Transitions in habits of life. Diversified habits in the same species. Species with habits widely different from those of their allies. Organs of extreme perfection. Means of transition. Cases of difficulty. Natura non facit saltum. Organs of small importance. Organs not in all cases absolutely perfect. The law of Unity of Type and of the Conditions of Existence embraced by the theo...

On The Absence Or Rarity Of Transitional Varieties.

As natural selection acts solely by the preservation of profitable modifications, each new form will tend in a fully-stocked country to take the place of, and finally to exterminate, its own less improved parent or other less-favoured forms with which it comes into competition. Thus extinction and natural selection will, as we have seen, go hand in hand. Hence, if we look at each species as descended from some other unknown form, both the parent and all the transitional varieties will generally ...

On The Origin And Transitions Of Organic Beings With Peculiar Habits And Structure.

It has been asked by the opponents of such views as I hold, how, for instance, a land carnivorous animal could have been converted into one with aquatic habits; for how could the animal in its transitional state have subsisted? It would be easy to show that within the same group carnivorous animals exist having every intermediate grade between truly aquatic and strictly terrestrial habits; and as each exists by a struggle for life, it is clear that each is well adapted in its habits to its place...

Organs Of Extreme Perfection And Complication.

To suppose that the eye, with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest possible degree. Yet reason tells me, that if numerous gradations from a perfect and complex eye to one very imperfect and simple, each grade being useful to its possessor, can be shown to exist...

Organs Of Little Apparent Importance.

As natural selection acts by life and death,–by the preservation of individuals with any favourable variation, and by the destruction of those with any unfavourable deviation of structure,–I have sometimes felt much difficulty in understanding the origin of simple parts, of which the importance does not seem sufficient to cause the preservation of successively varying individuals. I have sometimes felt as much difficulty, though of a very different kind, on this head, as in the case of an organ ...

Summary Of Chapter.

We have in this chapter discussed some of the difficulties and objections which may be urged against my theory. Many of them are very grave; but I think that in the discussion light has been thrown on several facts, which on the theory of independent acts of creation are utterly obscure. We have seen that species at any one period are not indefinitely variable, and are not linked together by a multitude of intermediate gradations, partly because the process of natural selection will always be ve...

Instincts comparable with habits, but different in their origin. Instincts graduated. Aphides and ants. Instincts variable. Domestic instincts, their origin. Natural instincts of the cuckoo, ostrich, and parasitic bees. Slave-making ants. Hive-bee, its cell-making instinct. Difficulties on the theory of the Natural Selection of instincts. Neuter or sterile insects. Summary.

The subject of instinct might have been worked into the previous chapters; but I have thought that it would be more conveni...

Slave-Making Instinct.

This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber, a better observer even than his celebrated father. This ant is absolutely dependent on its slaves; without their aid, the species would certainly become extinct in a single year. The males and fertile females do no work. The workers or sterile females, though most energetic and courageous in capturing slaves, do no other work. They are incapable of making their own nests, or of feeding their own l...

Cell-Making Instinct Of The Hive-Bee.

I will not here enter on minute details on this subject, but will merely give an outline of the conclusions at which I have arrived. He must be a dull man who can examine the exquisite structure of a comb, so beautifully adapted to its end, without enthusiastic admiration. We hear from mathematicians that bees have practically solved a recondite problem, and have made their cells of the proper shape to hold the greatest possible amount of honey, with the least possible consumption of precious wa...

Summary.

I have endeavoured briefly in this chapter to show that the mental qualities of our domestic animals vary, and that the variations are inherited. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. No one will dispute that instincts are of the highest importance to each animal. Therefore I can see no difficulty, under changing conditions of life, in natural selection accumulating slight modifications of instinct to any extent, in any useful direction. I...

Distinction between the sterility of first crosses and of hybrids. Sterility various in degree, not universal, affected by close interbreeding, removed by domestication. Laws governing the sterility of hybrids. Sterility not a special endowment, but incidental on other differences. Causes of the sterility of first crosses and of hybrids. Parallelism between the effects of changed conditions of life and crossing. Fertility of varieties when crossed and of their mongrel offspring not universal. Hy...

Laws Governing The Sterility Of First Crosses And Of Hybrids.

We will now consider a little more in detail the circumstances and rules governing the sterility of first crosses and of hybrids. Our chief object will be to see whether or not the rules indicate that species have specially been endowed with this quality, in order to prevent their crossing and blending together in utter confusion. The following rules and conclusions are chiefly drawn up from Gartner’s admirable work on the hybridisation of plants. I have taken much pains to ascertain how far the...

Causes Of The Sterility Of First Crosses And Of Hybrids.

We may now look a little closer at the probable causes of the sterility of first crosses and of hybrids. These two cases are fundamentally different, for, as just remarked, in the union of two pure species the male and female sexual elements are perfect, whereas in hybrids they are imperfect. Even in first crosses, the greater or lesser difficulty in effecting a union apparently depends on several distinct causes. There must sometimes be a physical impossibility in the male element reaching the ...

Fertility Of Varieties When Crossed, And Of Their Mongrel Offspring.

It may be urged, as a most forcible argument, that there must be some essential distinction between species and varieties, and that there must be some error in all the foregoing remarks, inasmuch as varieties, however much they may differ from each other in external appearance, cross with perfect facility, and yield perfectly fertile offspring. I fully admit that this is almost invariably the case. But if we look to varieties produced under nature, we are immediately involved in hopeless difficu...

Hybrids And Mongrels Compared, Independently Of Their Fertility.

Independently of the question of fertility, the offspring of species when crossed and of varieties when crossed may be compared in several other respects. Gartner, whose strong wish was to draw a marked line of distinction between species and varieties, could find very few and, as it seems to me, quite unimportant differences between the so-called hybrid offspring of species, and the so-called mongrel offspring of varieties. And, on the other hand, they agree most closely in very many important ...

Summary Of Chapter.

First crosses between forms sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally, sterile. The sterility is of all degrees, and is often so slight that the two most careful experimentalists who have ever lived, have come to diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible of favourable and unfavourable conditions. The degree of ...

On the absence of intermediate varieties at the present day. On the nature of extinct intermediate varieties; on their number. On the vast lapse of time, as inferred from the rate of deposition and of denudation. On the poorness of our palaeontological collections. On the intermittence of geological formations. On the absence of intermediate varieties in any one formation. On the sudden appearance of groups of species. On their sudden appearance in the lowest known fossiliferous strata.

In the s...

On The Lapse Of Time.

Independently of our not finding fossil remains of such infinitely numerous connecting links, it may be objected, that time will not have sufficed for so great an amount of organic change, all changes having been effected very slowly through natural selection. It is hardly possible for me even to recall to the reader, who may not be a practical geologist, the facts leading the mind feebly to comprehend the lapse of time. He who can read Sir Charles Lyell’s grand work on the Principles of Geology...

On The Poorness Of Our Palaeontological Collections.

That our palaeontological collections are very imperfect, is admitted by every one. The remark of that admirable palaeontologist, the late Edward Forbes, should not be forgotten, namely, that numbers of our 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 Eur...

On The Sudden Appearance Of Whole Groups Of Allied Species.

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 by none more forcibly than by Professor 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 all at once, the fact would be fatal to the theory of descent with slow modification through natural selection. For t...

On The Sudden Appearance Of Groups Of Allied Species In The Lowest Known Fossiliferous Strata.

There is another and allied difficulty, which is much graver. I allude to the manner in which numbers of species of the same group, suddenly appear in the lowest known fossiliferous rocks. Most of the arguments which have convinced me that all the existing species of the same group have descended from one progenitor, apply with nearly equal force to the earliest known species. For instance, I cannot doubt that all the Silurian trilobites have descended from some one crustacean, which must have l...

On the slow and successive appearance of new species. On their different rates of change. Species once lost do not reappear. Groups of species follow the same general rules in their appearance and disappearance as do single species. On Extinction. On simultaneous changes in the forms of life throughout the world. On the affinities of extinct species to each other and to living species. On the state of development of ancient forms. On the succession of the same types within the same areas. Summar...

On Extinction.

We have as yet spoken only incidentally of the disappearance of species and of groups of species. On the theory of natural selection the extinction of old forms and the production of new and improved forms are intimately connected together. The old notion of all the inhabitants of the earth having been swept away at successive periods by catastrophes, is very generally given up, even by those geologists, as Elie de Beaumont, Murchison, Barrande, etc., whose general views would naturally lead the...

On The Forms Of Life Changing Almost Simultaneously Throughout The World.

Scarcely any palaeontological discovery is more striking than the fact, that the forms of life change almost simultaneously throughout the world. Thus our European Chalk formation can be recognised in many distant parts of the world, under the most different climates, where not a fragment of the mineral chalk itself can be found; namely, in North America, in equatorial South America, in Tierra del Fuego, at the Cape of Good Hope, and in the peninsula of India. For at these distant points, the or...

On The Affinities Of Extinct Species To Each Other, And To Living Forms.

Let us now look to the mutual affinities of extinct and living species. They all fall into one grand natural system; and this fact is at once explained on the principle of descent. The more ancient any form is, the more, as a general rule, it differs from living forms. But, as Buckland long ago remarked, all fossils can be classed either in still existing groups, or between them. That the extinct forms of life help to fill up the wide intervals between existing genera, families, and orders, cann...

On The State Of Development Of Ancient Forms.

There has been much discussion whether recent forms are more highly developed than ancient. I will not here enter on this subject, for naturalists have not as yet defined to each other’s satisfaction what is meant by high and low forms. But in one particular sense the more recent forms must, on my theory, be higher than the more ancient; for each new species is formed by having had some advantage in the struggle for life over other and preceding forms. If under a nearly similar climate, the eoce...

On The Succession Of The Same Types Within The Same Areas, During The Later Tertiary Periods.

Mr. Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that continent. In South America, a similar relationship is manifest, even to an uneducated eye, in the gigantic pieces of armour like those of the armadillo, found in several parts of La Plata; and Professor Owen has shown in the most striking manner that most of the fossil mammals, buried there in such numbers, are related to South American types. This relationship ...

Summary Of The Preceding And Present Chapters.

I have attempted to show that the geological record is extremely imperfect; that only a small portion of the globe has been geologically explored with care; that only certain classes of organic beings have been largely preserved in a fossil state; that the number both of specimens and of species, preserved in our museums, is absolutely as nothing compared with the incalculable number of generations which must have passed away even during a single formation; that, owing to subsidence being necess...

Present distribution cannot be accounted for by differences in physical conditions. Importance of barriers. Affinity of the productions of the same continent. Centres of creation. Means of dispersal, by changes of climate and of the level of the land, and by occasional means. Dispersal during the Glacial period co-extensive with the world.

In considering the distribution of organic beings over the face of the globe, the first great fact which strikes us is, that neither the similarity nor the di...

Means Of Dispersal.

Sir C. Lyell and other authors have ably treated this subject. I can give here only the briefest abstract of the more important facts. Change of climate must have had a powerful influence on migration: a region when its climate was different may have been a high road for migration, but now be impassable; I shall, however, presently have to discuss this branch of the subject in some detail. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two mar...

Dispersal During The Glacial Period.

The identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where the Alpine species could not possibly exist, is one of the most striking cases known of the same species living at distant points, without the apparent possibility of their having migrated from one to the other. It is indeed a remarkable fact to see so many of the same plants living on the snowy regions of the Alps or Pyrenees, and in the extreme northern parts of Europ...

On The Inhabitants Of Oceanic Islands.

We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty, on the view that all the individuals both of the same and of allied species have descended from a single parent; and therefore have all proceeded from a common birthplace, notwithstanding that in the course of time they have come to inhabit distant points of the globe. I have already stated that I cannot honestly admit Forbes’s view on continental extensions, which, if le...

Summary Of Last And Present Chapters.

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 ...

CLASSIFICATION, groups subordinate to groups. Natural system. Rules and difficulties in classification, explained on the theory of descent with modification. Classification of varieties. Descent always used in classification. Analogical or adaptive characters. Affinities, general, complex and radiating. Extinction separates and defines groups. MORPHOLOGY, between members of the same class, between parts of the same individual. EMBRYOLOGY, laws of, explained by variations not supervening at an ea...

Morphology.

We have seen that the members of the same class, independently of their habits of life, resemble each other in the general plan of their organisation. This resemblance is often expressed by the term “unity of type;” or by saying that the several parts and organs in the different species of the class are homologous. The whole subject is included under the general name of Morphology. This is the most interesting department of natural history, and may be said to be its very soul. What can be more c...

Embryology.

It has already been casually remarked that certain organs in the individual, which when mature become widely different and serve for different purposes, are in the embryo exactly alike. The embryos, also, of distinct animals within the same class are often strikingly similar: a better proof of this cannot be given, than a circumstance mentioned by Agassiz, namely, that having forgotten to ticket the embryo of some vertebrate animal, he cannot now tell whether it be that of a mammal, bird, or rep...

Rudimentary, Atrophied, Or Aborted Organs.

Organs or parts in this strange condition, bearing the stamp of inutility, are extremely common throughout nature. For instance, rudimentary mammae are very general in the males of mammals: I presume that the “bastard-wing” in birds may be safely considered as a digit in a rudimentary state: in very many snakes one lobe of the lungs is rudimentary; in other snakes there are rudiments of the pelvis and hind limbs. Some of the cases of rudimentary organs are extremely curious; for instance, the pr...

Summary.

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 im...

Causes of Variability. Effects of Habit. Correlation of Growth. Inheritance. Character of Domestic Varieties. Difficulty of distinguishing between Varieties and Species. Origin of Domestic Varieties from one or more Species. Domestic Pigeons, their Differences and Origin. Principle of Selection anciently followed, its Effects. Methodical and Unconscious Selection. Unknown Origin of our Domestic Productions. Circumstances favourable to Man’s power of Selection.

When we look to the individuals of ...

On The Breeds Of The Domestic Pigeon.

Believing that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every breed which I could purchase or obtain, and have been most kindly favoured with skins from several quarters of the world, more especially by the Honourable W. Elliot from India, and by the Honourable C. Murray from Persia. Many treatises in different languages have been published on pigeons, and some of them are very important, as being of considerable antiquity....

Selection.

Let us now briefly consider the steps by which domestic races have been produced, either from one or from several allied species. Some little effect may, perhaps, be attributed to the direct action of the external conditions of life, and some little to habit; but he would be a bold man who would account by such agencies for the differences of a dray and race horse, a greyhound and bloodhound, a carrier and tumbler pigeon. One of the most remarkable features in our domesticated races is that we s...

Natural Selection: its power compared with man’s selection, its power on characters of trifling importance, its power at all ages and on both sexes. Sexual Selection. On the generality of intercrosses between individuals of the same species. Circumstances favourable and unfavourable to Natural Selection, namely, intercrossing, isolation, number of individuals. Slow action. Extinction caused by Natural Selection. Divergence of Character, related to the diversity of inhabitants of any small area, ...

Sexual Selection.

Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex, the same fact probably occurs under nature, and if so, natural selection will be able to modify one sex in its functional relations to the other sex, or in relation to wholly different habits of life in the two sexes, as is sometimes the case with insects. And this leads me to say a few words on what I call Sexual Selection. This depends, not on a struggle for existence, but on a s...

Illustrations Of The Action Of Natural Selection.

In order to make it clear how, as I believe, natural selection acts, I must beg permission to give one or two imaginary illustrations. Let us take the case of a wolf, which preys on various animals, securing some by craft, some by strength, and some by fleetness; and let us suppose that the fleetest prey, a deer for instance, had from any change in the country increased in numbers, or that other prey had decreased in numbers, during that season of the year when the wolf is hardest pressed for fo...

On The Intercrossing Of Individuals.

I must here introduce a short digression. In the case of animals and plants with separated sexes, it is of course obvious that two individuals must always unite for each birth; but in the case of hermaphrodites this is far from obvious. Nevertheless I am strongly inclined to believe that with all hermaphrodites two individuals, either occasionally or habitually, concur for the reproduction of their kind. This view, I may add, was first suggested by Andrew Knight. We shall presently see its impor...

Circumstances Favourable To Natural Selection.

This is an extremely intricate subject. A large amount of inheritable and diversified variability is favourable, but I believe mere individual differences suffice for the work. A large number of individuals, by giving a better chance for the appearance within any given period of profitable variations, will compensate for a lesser amount of variability in each individual, and is, I believe, an extremely important element of success. Though nature grants vast periods of time for the work of natura...

Extinction.

This subject will be more fully discussed in our chapter on Geology; but it must be here alluded to from being intimately connected with natural selection. Natural selection acts solely through the preservation of variations in some way advantageous, which consequently endure. But as from the high geometrical powers of increase of all organic beings, each area is already fully stocked with inhabitants, it follows that as each selected and favoured form increases in number, so will the less favou...

Divergence Of Character.

The principle, which I have designated by this term, is of high importance on my theory, and explains, as I believe, several important facts. In the first place, varieties, even strongly-marked ones, though having somewhat of the character of species–as is shown by the hopeless doubts in many cases how to rank them–yet certainly differ from each other far less than do good and distinct species. Nevertheless, according to my view, varieties are species in the process of formation, or are, as I ha...

Summary Of Chapter.

If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an ...

Effects of external conditions. Use and disuse, combined with natural selection; organs of flight and of vision. Acclimatisation. Correlation of growth. Compensation and economy of growth. False correlations. Multiple, rudimentary, and lowly organised structures variable. Parts developed in an unusual manner are highly variable: specific characters more variable than generic: secondary sexual characters variable. Species of the same genus vary in an analogous manner. Reversions to long lost char...

Effects Of Use And Disuse.

From the facts alluded to in the first chapter, I think there can be little doubt that use in our domestic animals strengthens and enlarges certain parts, and disuse diminishes them; and that such modifications are inherited. Under free nature, we can have no standard of comparison, by which to judge of the effects of long-continued use or disuse, for we know not the parent-forms; but many animals have structures which can be explained by the effects of disuse. As Professor Owen has remarked, th...

Acclimatisation.

Habit is hereditary with plants, as in the period of flowering, in the amount of rain requisite for seeds to germinate, in the time of sleep, etc., and this leads me to say a few words on acclimatisation. As it is extremely common for species of the same genus to inhabit very hot and very cold countries, and as I believe that all the species of the same genus have descended from a single parent, if this view be correct, acclimatisation must be readily effected during long-continued descent. It i...

Correlation Of Growth.

I mean by this expression that the whole organisation is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified. This is a very important subject, most imperfectly understood. The most obvious case is, that modifications accumulated solely for the good of the young or larva, will, it may safely be concluded, affect the structure of the adult; in the same manner as any malcon...

Summary.

Our ignorance of the laws of variation is profound. Not in one case out of a hundred can we pretend to assign any reason why this or that part differs, more or less, from the same part in the parents. But whenever we have the means of instituting a comparison, the same laws appear to have acted in producing the lesser differences between varieties of the same species, and the greater differences between species of the same genus. The external conditions of life, as climate and food, etc., seem t...

Difficulties on the theory of descent with modification. Transitions. Absence or rarity of transitional varieties. Transitions in habits of life. Diversified habits in the same species. Species with habits widely different from those of their allies. Organs of extreme perfection. Means of transition. Cases of difficulty. Natura non facit saltum. Organs of small importance. Organs not in all cases absolutely perfect. The law of Unity of Type and of the Conditions of Existence embraced by the theo...

On The Absence Or Rarity Of Transitional Varieties.

As natural selection acts solely by the preservation of profitable modifications, each new form will tend in a fully-stocked country to take the place of, and finally to exterminate, its own less improved parent or other less-favoured forms with which it comes into competition. Thus extinction and natural selection will, as we have seen, go hand in hand. Hence, if we look at each species as descended from some other unknown form, both the parent and all the transitional varieties will generally ...

On The Origin And Transitions Of Organic Beings With Peculiar Habits And Structure.

It has been asked by the opponents of such views as I hold, how, for instance, a land carnivorous animal could have been converted into one with aquatic habits; for how could the animal in its transitional state have subsisted? It would be easy to show that within the same group carnivorous animals exist having every intermediate grade between truly aquatic and strictly terrestrial habits; and as each exists by a struggle for life, it is clear that each is well adapted in its habits to its place...

Organs Of Extreme Perfection And Complication.

To suppose that the eye, with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest possible degree. Yet reason tells me, that if numerous gradations from a perfect and complex eye to one very imperfect and simple, each grade being useful to its possessor, can be shown to exist...

Organs Of Little Apparent Importance.

As natural selection acts by life and death,–by the preservation of individuals with any favourable variation, and by the destruction of those with any unfavourable deviation of structure,–I have sometimes felt much difficulty in understanding the origin of simple parts, of which the importance does not seem sufficient to cause the preservation of successively varying individuals. I have sometimes felt as much difficulty, though of a very different kind, on this head, as in the case of an organ ...

Summary Of Chapter.

We have in this chapter discussed some of the difficulties and objections which may be urged against my theory. Many of them are very grave; but I think that in the discussion light has been thrown on several facts, which on the theory of independent acts of creation are utterly obscure. We have seen that species at any one period are not indefinitely variable, and are not linked together by a multitude of intermediate gradations, partly because the process of natural selection will always be ve...

Instincts comparable with habits, but different in their origin. Instincts graduated. Aphides and ants. Instincts variable. Domestic instincts, their origin. Natural instincts of the cuckoo, ostrich, and parasitic bees. Slave-making ants. Hive-bee, its cell-making instinct. Difficulties on the theory of the Natural Selection of instincts. Neuter or sterile insects. Summary.

The subject of instinct might have been worked into the previous chapters; but I have thought that it would be more conveni...

Slave-Making Instinct.

This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber, a better observer even than his celebrated father. This ant is absolutely dependent on its slaves; without their aid, the species would certainly become extinct in a single year. The males and fertile females do no work. The workers or sterile females, though most energetic and courageous in capturing slaves, do no other work. They are incapable of making their own nests, or of feeding their own l...

Cell-Making Instinct Of The Hive-Bee.

I will not here enter on minute details on this subject, but will merely give an outline of the conclusions at which I have arrived. He must be a dull man who can examine the exquisite structure of a comb, so beautifully adapted to its end, without enthusiastic admiration. We hear from mathematicians that bees have practically solved a recondite problem, and have made their cells of the proper shape to hold the greatest possible amount of honey, with the least possible consumption of precious wa...

Summary.

I have endeavoured briefly in this chapter to show that the mental qualities of our domestic animals vary, and that the variations are inherited. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. No one will dispute that instincts are of the highest importance to each animal. Therefore I can see no difficulty, under changing conditions of life, in natural selection accumulating slight modifications of instinct to any extent, in any useful direction. I...

Distinction between the sterility of first crosses and of hybrids. Sterility various in degree, not universal, affected by close interbreeding, removed by domestication. Laws governing the sterility of hybrids. Sterility not a special endowment, but incidental on other differences. Causes of the sterility of first crosses and of hybrids. Parallelism between the effects of changed conditions of life and crossing. Fertility of varieties when crossed and of their mongrel offspring not universal. Hy...

Laws Governing The Sterility Of First Crosses And Of Hybrids.

We will now consider a little more in detail the circumstances and rules governing the sterility of first crosses and of hybrids. Our chief object will be to see whether or not the rules indicate that species have specially been endowed with this quality, in order to prevent their crossing and blending together in utter confusion. The following rules and conclusions are chiefly drawn up from Gartner’s admirable work on the hybridisation of plants. I have taken much pains to ascertain how far the...

Causes Of The Sterility Of First Crosses And Of Hybrids.

We may now look a little closer at the probable causes of the sterility of first crosses and of hybrids. These two cases are fundamentally different, for, as just remarked, in the union of two pure species the male and female sexual elements are perfect, whereas in hybrids they are imperfect. Even in first crosses, the greater or lesser difficulty in effecting a union apparently depends on several distinct causes. There must sometimes be a physical impossibility in the male element reaching the ...

Fertility Of Varieties When Crossed, And Of Their Mongrel Offspring.

It may be urged, as a most forcible argument, that there must be some essential distinction between species and varieties, and that there must be some error in all the foregoing remarks, inasmuch as varieties, however much they may differ from each other in external appearance, cross with perfect facility, and yield perfectly fertile offspring. I fully admit that this is almost invariably the case. But if we look to varieties produced under nature, we are immediately involved in hopeless difficu...

Hybrids And Mongrels Compared, Independently Of Their Fertility.

Independently of the question of fertility, the offspring of species when crossed and of varieties when crossed may be compared in several other respects. Gartner, whose strong wish was to draw a marked line of distinction between species and varieties, could find very few and, as it seems to me, quite unimportant differences between the so-called hybrid offspring of species, and the so-called mongrel offspring of varieties. And, on the other hand, they agree most closely in very many important ...

Summary Of Chapter.

First crosses between forms sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally, sterile. The sterility is of all degrees, and is often so slight that the two most careful experimentalists who have ever lived, have come to diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible of favourable and unfavourable conditions. The degree of ...

On the absence of intermediate varieties at the present day. On the nature of extinct intermediate varieties; on their number. On the vast lapse of time, as inferred from the rate of deposition and of denudation. On the poorness of our palaeontological collections. On the intermittence of geological formations. On the absence of intermediate varieties in any one formation. On the sudden appearance of groups of species. On their sudden appearance in the lowest known fossiliferous strata.

In the s...

On The Lapse Of Time.

Independently of our not finding fossil remains of such infinitely numerous connecting links, it may be objected, that time will not have sufficed for so great an amount of organic change, all changes having been effected very slowly through natural selection. It is hardly possible for me even to recall to the reader, who may not be a practical geologist, the facts leading the mind feebly to comprehend the lapse of time. He who can read Sir Charles Lyell’s grand work on the Principles of Geology...

On The Poorness Of Our Palaeontological Collections.

That our palaeontological collections are very imperfect, is admitted by every one. The remark of that admirable palaeontologist, the late Edward Forbes, should not be forgotten, namely, that numbers of our 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 Eur...

On The Sudden Appearance Of Whole Groups Of Allied Species.

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 by none more forcibly than by Professor 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 all at once, the fact would be fatal to the theory of descent with slow modification through natural selection. For t...

On The Sudden Appearance Of Groups Of Allied Species In The Lowest Known Fossiliferous Strata.

There is another and allied difficulty, which is much graver. I allude to the manner in which numbers of species of the same group, suddenly appear in the lowest known fossiliferous rocks. Most of the arguments which have convinced me that all the existing species of the same group have descended from one progenitor, apply with nearly equal force to the earliest known species. For instance, I cannot doubt that all the Silurian trilobites have descended from some one crustacean, which must have l...

On the slow and successive appearance of new species. On their different rates of change. Species once lost do not reappear. Groups of species follow the same general rules in their appearance and disappearance as do single species. On Extinction. On simultaneous changes in the forms of life throughout the world. On the affinities of extinct species to each other and to living species. On the state of development of ancient forms. On the succession of the same types within the same areas. Summar...

On Extinction.

We have as yet spoken only incidentally of the disappearance of species and of groups of species. On the theory of natural selection the extinction of old forms and the production of new and improved forms are intimately connected together. The old notion of all the inhabitants of the earth having been swept away at successive periods by catastrophes, is very generally given up, even by those geologists, as Elie de Beaumont, Murchison, Barrande, etc., whose general views would naturally lead the...

On The Forms Of Life Changing Almost Simultaneously Throughout The World.

Scarcely any palaeontological discovery is more striking than the fact, that the forms of life change almost simultaneously throughout the world. Thus our European Chalk formation can be recognised in many distant parts of the world, under the most different climates, where not a fragment of the mineral chalk itself can be found; namely, in North America, in equatorial South America, in Tierra del Fuego, at the Cape of Good Hope, and in the peninsula of India. For at these distant points, the or...

On The Affinities Of Extinct Species To Each Other, And To Living Forms.

Let us now look to the mutual affinities of extinct and living species. They all fall into one grand natural system; and this fact is at once explained on the principle of descent. The more ancient any form is, the more, as a general rule, it differs from living forms. But, as Buckland long ago remarked, all fossils can be classed either in still existing groups, or between them. That the extinct forms of life help to fill up the wide intervals between existing genera, families, and orders, cann...

On The State Of Development Of Ancient Forms.

There has been much discussion whether recent forms are more highly developed than ancient. I will not here enter on this subject, for naturalists have not as yet defined to each other’s satisfaction what is meant by high and low forms. But in one particular sense the more recent forms must, on my theory, be higher than the more ancient; for each new species is formed by having had some advantage in the struggle for life over other and preceding forms. If under a nearly similar climate, the eoce...

On The Succession Of The Same Types Within The Same Areas, During The Later Tertiary Periods.

Mr. Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that continent. In South America, a similar relationship is manifest, even to an uneducated eye, in the gigantic pieces of armour like those of the armadillo, found in several parts of La Plata; and Professor Owen has shown in the most striking manner that most of the fossil mammals, buried there in such numbers, are related to South American types. This relationship ...

Summary Of The Preceding And Present Chapters.

I have attempted to show that the geological record is extremely imperfect; that only a small portion of the globe has been geologically explored with care; that only certain classes of organic beings have been largely preserved in a fossil state; that the number both of specimens and of species, preserved in our museums, is absolutely as nothing compared with the incalculable number of generations which must have passed away even during a single formation; that, owing to subsidence being necess...

Present distribution cannot be accounted for by differences in physical conditions. Importance of barriers. Affinity of the productions of the same continent. Centres of creation. Means of dispersal, by changes of climate and of the level of the land, and by occasional means. Dispersal during the Glacial period co-extensive with the world.

In considering the distribution of organic beings over the face of the globe, the first great fact which strikes us is, that neither the similarity nor the di...

Means Of Dispersal.

Sir C. Lyell and other authors have ably treated this subject. I can give here only the briefest abstract of the more important facts. Change of climate must have had a powerful influence on migration: a region when its climate was different may have been a high road for migration, but now be impassable; I shall, however, presently have to discuss this branch of the subject in some detail. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two mar...

Dispersal During The Glacial Period.

The identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where the Alpine species could not possibly exist, is one of the most striking cases known of the same species living at distant points, without the apparent possibility of their having migrated from one to the other. It is indeed a remarkable fact to see so many of the same plants living on the snowy regions of the Alps or Pyrenees, and in the extreme northern parts of Europ...

On The Inhabitants Of Oceanic Islands.

We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty, on the view that all the individuals both of the same and of allied species have descended from a single parent; and therefore have all proceeded from a common birthplace, notwithstanding that in the course of time they have come to inhabit distant points of the globe. I have already stated that I cannot honestly admit Forbes’s view on continental extensions, which, if le...

Summary Of Last And Present Chapters.

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 ...

CLASSIFICATION, groups subordinate to groups. Natural system. Rules and difficulties in classification, explained on the theory of descent with modification. Classification of varieties. Descent always used in classification. Analogical or adaptive characters. Affinities, general, complex and radiating. Extinction separates and defines groups. MORPHOLOGY, between members of the same class, between parts of the same individual. EMBRYOLOGY, laws of, explained by variations not supervening at an ea...

Morphology.

We have seen that the members of the same class, independently of their habits of life, resemble each other in the general plan of their organisation. This resemblance is often expressed by the term “unity of type;” or by saying that the several parts and organs in the different species of the class are homologous. The whole subject is included under the general name of Morphology. This is the most interesting department of natural history, and may be said to be its very soul. What can be more c...

Embryology.

It has already been casually remarked that certain organs in the individual, which when mature become widely different and serve for different purposes, are in the embryo exactly alike. The embryos, also, of distinct animals within the same class are often strikingly similar: a better proof of this cannot be given, than a circumstance mentioned by Agassiz, namely, that having forgotten to ticket the embryo of some vertebrate animal, he cannot now tell whether it be that of a mammal, bird, or rep...

Rudimentary, Atrophied, Or Aborted Organs.

Organs or parts in this strange condition, bearing the stamp of inutility, are extremely common throughout nature. For instance, rudimentary mammae are very general in the males of mammals: I presume that the “bastard-wing” in birds may be safely considered as a digit in a rudimentary state: in very many snakes one lobe of the lungs is rudimentary; in other snakes there are rudiments of the pelvis and hind limbs. Some of the cases of rudimentary organs are extremely curious; for instance, the pr...

Summary.

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 im...