Darwin's On the Origin of Species: A Modern Rendition

INTRODUCTION

WHEN ON BOARD HMS BEAGLE AS NATURALIST, I WAS STRUCK by the distribution of South America’s organisms and the geological relationships between its past and present inhabitants. These observations seemed to me to illuminate that mystery of mysteries: the origin of species. After my return home, it occurred to me, in 1837, that this question may be clarified by patiently accumulating and reflecting on all sorts of relevant facts. Following five years’ work I began to speculate on the subject and drew up some short notes, which I enlarged in 1844 into a sketch of probable conclusions. Since then I have steadily pursued the same object. I hope the reader will excuse me for entering on these personal details; I just want to show that I have not been hasty in coming to a decision.

My work is nearly finished, but I will require two or three more years to complete it, and as my health is far from strong I have been urged to publish this abstract. I was further prompted to do this because last year Alfred Russel Wallace, who is now studying the natural history of the Malay Archipelago, sent me a memoir in which he arrives at conclusions very similar to mine. He requested that I forward it to Sir Charles Lyell. Lyell and Dr. Hooker, who both knew of my work – Dr. Hooker having read my 1844 sketch – honored me by advising publication of extracts from my own manuscript alongside Mr. Wallace’s excellent piece. Both appear in the third volume of the Journal of the Linnean Society.

This abstract is necessarily imperfect. I cannot here provide references and must trust the reader’s confidence in my accuracy. Errors have no doubt crept in, although I have tried to use only reliable authorities. I give only general conclusions with a few illustrative examples, which I hope will suffice. And I entirely appreciate the necessity of describing all the information upon which I ground my conclusions in a future work. I am well aware that little is discussed to which additional observations could not be given, often leading to apparently contradictory conclusions. A fair assessment can only be reached by a full statement of the facts and balancing arguments on both sides of each issue; this cannot possibly be done here.

I regret that a lack of space prevents acknowledgment of the generous assistance I have received from many naturalists, some of them personally unknown to me. However, I will not pass up this opportunity to thank Dr. Hooker, who has aided me over the past fifteen years in every possible way with his excellent judgment and large stores of knowledge.

A naturalist considering the origin of species and reflecting on the affinities among organisms, their embryological relationships, their geographic distribution, geological succession, and other factors might conclude that each species had not been created independently but had descended as varieties do from other species. But even if such a conclusion were well founded, it would be unsatisfactory until it were shown how the innumerable species inhabiting the earth have been modified to a perfection of structure and coadaptation that justly excites our admiration. Naturalists refer to external conditions, such as climate and food supply, as the only possible cause of variation. This may be true in one very limited sense, as we will see, but it’s absurd to claim, for example, that the woodpecker’s feet, tail, beak, and tongue have become so well adapted to catch insects under tree bark simply due to external conditions. Or consider the mistletoe, which draws nourishment from certain trees, has seeds that must be transported by specific birds, and has flowers with separate sexes requiring particular insects to bring pollen from one flower to the other. It is equally absurd to claim that the structure of this parasitic plant, with its relationships to distinct organisms, results from habits or volition or external conditions. The author of the Vestiges of the Natural History of Creation would presumably argue that after an indeterminate number of generations some bird had given birth to a perfectly formed woodpecker and some plant had brought forth a perfectly formed mistletoe. But this assumption is no explanation, leaving the coadaptations of organisms to one another and their physical environments untouched.

So it is very important to learn how modification and coadaptation happen. As I began my observations, it seemed that a careful study of domesticated plants and animals would provide the best chance for insight into this problem. I have not been disappointed; our knowledge of variation under domestication, though imperfect, invariably affords the best and safest clue to these and other perplexing problems. I suggest that this field is highly valuable, though it has commonly been neglected by naturalists.

I therefore devote the first chapter of this abstract to variation under domestication. I demonstrate that a large amount of hereditary modification is at least possible and, perhaps more importantly, that humans have caused huge changes in domesticated plants and animals through the selection and accumulation of slight successive variations. I then briefly discuss the variability of species in the wild. This topic could only have been treated properly by long catalogs of facts, but I nevertheless discuss the circumstances favorable to variation. The third chapter treats the struggle for existence among all organisms, which follows inevitably from their ability to proliferate geometrically: the doctrine of Malthus applied to all living things. Because many more individuals of each species are born than can possibly survive, any individual possessing even a slightly favorable variation enjoys a better chance of surviving the complex and sometimes fluctuating environment and is naturally selected. The principle of inheritance ensures that a selected variety will tend to propagate its new and modified form.

This fundamental subject of natural selection is treated at length in the fourth chapter, where I discuss how it often causes extinction of less improved life forms and induces divergence of character. In the following chapter I address the complex and poorly understood rules of variation and correlated growth. In the four succeeding chapters I present the most obvious and serious challenges to the theory: (1) how a simple organism or simple organ can be changed and perfected into something highly developed or elaborately constructed, (2) the mental power of animals (instinct), (3) the infertility of species but fertility of varieties when crossed (hybridism), and (4) the imperfection of the geological record. Then I consider the geological succession of organisms through time; in the eleventh and twelfth chapters, their geographic distribution; in the thirteenth, their classification, based on affinities in both embryonic and fully developed states; and in the last chapter I give a brief summary of the work and a few concluding remarks.

Given our ignorance of the relationships among organisms, it is not surprising that much remains to be explained about the origin of species. Who can explain why one species is numerous with a wide range while a related species is rare with a narrow range? And yet such questions are important because their answers explain the present state and future modifications and success of all organisms. We know even less about the relationships among the innumerable past inhabitants of the earth. Although much remains obscure, and will long remain obscure, the most deliberate study and dispassionate judgment of which I am capable have dissuaded me from the view commonly held by naturalists, and previously held by me, that each species has been independently created. I am fully convinced that species are mutable and that species within a genus are linearly descended from some usually extinct species, in the same way that a variety of a given species is a descendant of that species. I am also convinced that natural selection has been the main, though not exclusive, means of modification.

1

VARIATION UNDER DOMESTICATION

IN CONSIDERING THE INDIVIDUALS OF A DOMESTICATED plant or animal variety, it is striking that they are generally more diverse than those belonging to varieties or species in the wild. The vast diversity of domesticated organisms, which have varied under many different climates and treatments, suggests that greater variability results from the conditions under which domestication occurs – conditions unlike those encountered by the parent species in the wild. This variability may partly be connected with excess food, as proposed by Andrew Knight. It seems clear that organisms must be exposed to a new environment over several generations for it to cause appreciable variation, and once organization begins to vary, it usually continues to do so for many generations. There is no case of a variable organism ceasing to be variable under domestication. Established domesticated plants such as wheat still often yield new varieties, and animals domesticated long ago are still capable of rapid improvement or modification.

It is disputed whether the causes of variation – whatever they may be – act during the early or late stage of embryonic development or at the instant of conception. Isidore Geoffroy St. Hilaire’s experiments show that unnatural treatment of the embryo causes monstrosities, which cannot be clearly differentiated from mere variations. I strongly suspect that variability is most frequently caused by effects on the egg or sperm before conception, mainly because of the remarkable influence of cultivation or confinement on the functions of the reproductive system, which appear far more susceptible to environmental changes than any other component of organization. Nothing is easier than taming an animal and nothing more difficult than getting it to reproduce in confinement, even when the male and female mate. This is generally attributed to impaired instincts, but many cultivated plants are vigorous yet do not seed. In some cases, minor changes, like a little more or less water at a particular period of growth, determine whether or not a plant will produce seeds. I will not go into the copious details I have collected on this curious subject, but to illustrate the strangeness of the rules that govern the reproduction of captive animals, consider that, with the exception of bears, carnivorous mammals, even from the tropics, breed freely in Britain under confinement, whereas carnivorous birds rarely lay fertile eggs. Many exotic plants have pollen as useless as that of the most sterile hybrids. Some domesticated plants and animals that are otherwise weak and sickly breed freely under confinement; but tame, long-lived, and healthy individuals taken young from the wild may have reproductive systems so seriously affected by unknown causes that they are nonfunctional. Unsurprisingly, then, when the reproductive system actually works under confinement, it does so irregularly, producing offspring that are different from the parents. Finally, some organisms breed under very unnatural conditions – like rabbits and ferrets kept in hutches – demonstrating that their reproductive systems have not been affected. So some organisms withstand domestication and vary only slightly, perhaps hardly more than in the wild.

Sterility is a horticultural nuisance, but variability, the source of all the choicest productions of the garden, shares a cause with sterility. There are many plants (called sporting plants by gardeners) that produce single buds or offshoots with novel characteristics, sometimes very different from the rest of the plant. Such buds can be propagated by grafting or other techniques, and sometimes by seed. These sports are rare in the wild but common under cultivation. In this case, manipulation of the parent affects a bud or offshoot but not the ovules or pollen. According to most physiologists, however, there is no essential difference between a bud and an ovule in the earliest stages of formation. Therefore, sports show that variability may be largely attributed to the effect on the ovules, pollen, or both by treatment of the parent prior to conception. In any case, these examples demonstrate that variation is not necessarily connected with the act of generation, as some authors have suggested.

Seedlings from the same fruit and young from the same litter sometimes differ considerably from each other even though both parent and offspring have apparently been exposed to the same conditions, as Müller has remarked. This shows how unimportant direct environmental effects are in comparison to the laws governing reproduction, growth, and inheritance. If the influence of environment were direct, then variation would be the same among offspring. Judging the extent to which heat, moisture, light, food, and other factors have an impact on variation is difficult. My impression is that such agents produce very little direct effect on animals, but apparently more on plants. (Mr. Buckman’s recent experiments on plants are valuable here.) When all or nearly all individuals exposed to certain conditions are identically affected, the resultant changes appear to flow directly from the conditions. But in some cases opposite conditions generate similar structural changes. Nevertheless, some slight amount of change may be attributed to direct environmental action, as in certain cases of increased size from greater food intake, altered coloration from particular kinds of food and light, and perhaps the thickness of fur from climate.

Habit also has a deciding influence, as with the flowering period of plants transported from one climate to another. The effect is greater in animals. For example, I find that the wing bones of a domestic duck weigh less and the leg bones weigh more in proportion to the whole skeleton than do those of a wild duck. I presume this results from the domestic duck flying much less and walking more than its wild parent. The large inherited udders of cows and goats in countries where they are habitually milked is another example of the effect of use. There is no domestic animal that in some region does not have drooping ears. As suggested by some authors, this probably results from the disuse of ear muscles, the animals being rarely alarmed.

Many rules regulate variation; some of them can be dimly seen and will be briefly mentioned. Here I will only allude to correlated growth. For example, a change in the embryo or larva often entails changes in the mature animal. With monstrosities, correlations between distinct parts are very curious.¹ Breeders maintain that long limbs are often accompanied by an elongated head. Some correlations are whimsical; for example, cats with blue eyes are invariably deaf. There are many remarkable cases among plants and animals of coloration and constitutional peculiarities going together. Observations collected by Heusinger suggest that white sheep and pigs are affected differently by poisonous vegetables than individuals with coloration. Hairless dogs have imperfect teeth; long-haired and coarse-haired animals tend to have long or many horns; pigeons with feathered feet have skin between their outer toes; pigeons with short beaks have small feet, and those with long beaks have large feet. If humans select and augment a peculiarity, they will probably unintentionally modify other parts as a consequence of the mysterious rules of correlated growth.

The dimly seen or unknown rules of variation yield infinitely complex and diverse results. Treatises on some established domesticated plants, such as the hyacinth, potato, and even the dahlia, reveal a surprising number of slight structural and constitutional differences between varieties and subvarieties. The whole organization seems to have become plastic, with a tendency to depart somewhat from the parental type.

Variations that cannot be inherited are unimportant to this argument. What nevertheless remains is an endless number of diverse heritable structural deviations of both slight and considerable physiological importance. (Dr. Prosper Lucas’s two-volume work is the best treatment of this subject.) The strong propensity for inheritance is known by breeders, whose fundamental belief is that like produces like. (Only theoretical writers have thrown doubt on this principle.) When a commonly occurring deviation is observed in both parent and offspring, it may result from the same cause acting on both. But when a very rare deviation due to some extraordinary combination of circumstances appears, say, once in several million individuals all apparently exposed to the same conditions, and it reappears in an offspring, the mere doctrine of chance almost compels us to attribute its reappearance to inheritance. Everyone has heard of albinism, prickly skin, hairy bodies, and other such peculiar characteristics reappearing in several members of the same family. If strange and rare deviations really are heritable, then surely commonplace deviations are also heritable. Perhaps the correct view is to take inheritance of every characteristic as the rule and non-inheritance as the anomaly.

The laws of inheritance are unknown. It is unknown why some given peculiarity of individuals within the same species, or of individuals among different species, is sometimes inherited and sometimes not, why a child reverts to characteristics found in a grandparent or more remote ancestor, or why some peculiarities are inherited in a gender-dependent manner.² Peculiarities appearing in the males of domestic breeds are often transmitted exclusively, or more strongly, to male progeny. A more important rule is that the age at which a peculiarity first appears tends to be the same in the parent and in its offspring (although sometimes earlier in the offspring). In many cases this cannot be otherwise. For example, the inherited peculiarities of cattle horns can appear only as the offspring mature, and peculiarities in the silkworm are known to appear at the corresponding caterpillar or cocoon stage. Hereditary diseases and some other examples suggest that this rule is generally applicable: even when there is no apparent reason for a peculiarity to appear at a particular stage, it tends to appear in the offspring at the same period of development as in the parent. This is very important to illuminating the rules of embryology. These remarks are, of course, confined to the first appearance of a peculiarity and not to its primary cause, which may have acted on the egg or sperm. If the offspring of a short-horned cow and a long-horned bull develops long horns, then it’s clearly due to the sperm.

Naturalists often argue that when domestic varieties run wild, their characteristics gradually but surely revert to those found in the original stocks, and that, consequently, deductions drawn from domestic varieties cannot be applied to species in nature. I have tried without success to find the decisive facts on which this statement is so often and so boldly made; it would be very difficult to prove, because many established domestic varieties could not possibly survive in the wild. In many cases we do not know what the original stock was and could not tell whether or not reversion had ensued. It would also be necessary to turn loose only one variety to avoid the effects of intercrossing. Nevertheless, varieties sometimes do partially revert to the parental form. For example, if various strains of cabbage were cultivated in very poor soil for many generations, they would probably revert wholly or largely to the wild stock. (However, some effect would have to be attributed to the direct action of the poor soil.) Whether or not the experiment would succeed is not particularly important to the argument, because the experiment necessarily alters the environment. If a strong tendency for reversion – that is, a loss of acquired characteristics under constant conditions in a large population so that free crossing, by blending, checks slight deviations of structure – were demonstrated in domesticated varieties, I would grant that nothing deduced from domestic varieties would apply to species. But there is not a shadow of evidence in favor of this view. To assert that we could not breed cart and racehorses, long- and short-haired cattle, and poultry of various breeds, and cultivate edible vegetables for an almost infinite number of generations is contrary to all experience. When the environment changes in nature, variations and reversions probably do occur, but natural selection, as will be explained, determines how far such new characteristics are preserved.

As already mentioned, there is less uniformity of character among individuals of a domestic variety than among individuals of a true species. Also, domestic varieties of the same species often have a monstrous character, by which I mean that although they differ in some minor respects from one another and members of the same genus, they often differ extremely in some one part. With these exceptions and that of the perfect fertility of crossed varieties (discussed later), domestic varieties of the same species differ from one another in a manner similar to the way closely related species of the same genus differ in the wild. There are very few domestic varieties of plant or animal that have not been classified by some competent judges as just varieties and by others as descendants of distinct parent species; if there were any significant distinction between domestic varieties and species, this source of doubt would be less common. Contrary to frequently made assertions, I think domestic varieties differ from one another in generic characteristics,³ which naturalists disagree in defining because all such valuations are currently empirical. Given the following examination of the origin of genera, there is no reason to often expect generic differences in domesticated organisms.

Attempts to estimate the amount of structural difference between domestic varieties of the same species are hampered by our ignorance of whether they have descended from one parent species or several; it would be interesting to clear up this problem. For example, if it were shown that the greyhound, bloodhound, terrier, spaniel, and bulldog, which propagate their kind truly, are derived from a single species, the supposed immutability of the many closely related natural species (such as the foxes) would be brought under considerable doubt. I do not believe that all dog breeds have descended from one wild species (see below),⁴ but there is tentative or even strong evidence that some other domestic varieties have.

Humans are often assumed to have chosen for domestication those plants and animals that possess an extraordinary inherent tendency to vary and to withstand diverse climates. Although such capacities have added significantly to the value of many domesticated productions, how could primitive humans have possibly known when first taming an animal that it would vary in succeeding generations and endure other climates? The limited variability of the ass and the guinea fowl, and the low tolerance for warmth by the reindeer and for cold by the common camel did not prevent their domestication. If plants and animals equal in number and belonging to equally diverse classes and regions to existing domesticated organisms were taken from the wild and bred for an equal number of generations under domestication, they would vary on average as much as the parent species of already domesticated organisms have varied.

I think it is impossible to ascertain with complete certainty whether established domesticated plants and animals have descended from one or multiple species. Those who believe in the multiple origin of domestic animals argue mainly that ancient records, especially on the monuments of Egypt, reveal a great diversity of breeds, some of which resemble or are identical to existing ones. Even if this were found to be more strictly and generally true than I believe is the case, it suggests only that some of our breeds originated there four or five thousand years ago. Based on Mr. Horner’s research, civilization advanced enough to manufacture pottery probably existed in the Nile valley thirteen or fourteen thousand years ago; it is not known how long before these ancient periods peoples like those of Tierra del Fuego or Australia, who possess a semidomesticated dog, may have existed in Egypt.

I think the whole subject must remain vague. Nevertheless, without going into details – but based on geographic and other considerations – I think it is likely that domestic dogs have descended from several wild species. I cannot form an opinion with respect to goats and sheep. Information about the habits, voice, constitution, and other features of humped Indian cattle, communicated to me by Mr. Blyth, indicate that it descended from a different stock than European cattle, which, in turn, have more than one parent, according to several judges. And for reasons I cannot cover here, I am doubtfully inclined to believe, in opposition to several authors, that all the varieties of horse have descended from one wild stock. Mr. Blyth – whose opinion I value highly, drawn as it is from his large and varied stores of knowledge – thinks that all poultry breeds have proceeded from the common wild Indian fowl. Duck and rabbit breeds, which differ considerably from one another in structure, have all descended from the common wild duck and rabbit.

Some authors carry the doctrine of plural descent to an absurd extreme, believing that every variety that breeds true, even those possessing only very slight distinctive characteristics, has a distinct wild prototype. At this rate there must have existed twenty species of wild cattle and sheep – and several goats – in Europe alone, and even several within Great Britain. One author believes that there once existed eleven unique wild sheep species in Great Britain! Britain barely has any unique mammals, France only a few distinct from Germany, and vice versa, and the same is true of Hungary, Spain, and other localities. Yet each of these kingdoms possesses several peculiar breeds of cattle, sheep, and other animals, meaning that many domestic breeds originated in Europe because it lacks sufficient unique species as parent stocks. This is also true for India. Even in the case of the domestic dogs of the whole world, which I admit probably descended from multiple wild species, there has been immense heritable variation. Who can believe that animals closely resembling the Italian greyhound, bloodhound, bulldog, or Blenheim spaniel – so unlike all wild dogs – ever existed in the wild? Crossing a few original dog species would result in only intermediate forms, and accounting for domestic dogs by this process requires the previous existence of extreme forms, similar to the breeds mentioned, in a wild state. In addition, the possibility of generating distinct varieties by crossing is exaggerated. Obviously a variety can be modified by occasional crossing if mongrels with desired characteristics are carefully selected, but a variety intermediate between two extremely different varieties or species could not be obtained. (Sir J. Sebright experimentally attempted precisely this and failed.) The offspring from the first cross between two pure breeds is fairly uniform – and as I have found with pigeons, sometimes very uniform – and everything seems simple enough. But when these mongrels are crossed with one another for several generations, few will be alike and the utter hopelessness of the task becomes apparent. An intermediate between two very distinct breeds could be obtained only with extreme care and continuous long-term selection, but I cannot find a single recorded case of a variety formed this way.

Believing that it is best to study one particular group, after deliberation I took up domestic pigeons. Pigeons have been watched, tended with the utmost care, and loved by many people. They have been domesticated for thousands of years in several parts of the world. Professor Lepsius has pointed out to me that the earliest known record of pigeons is in the fifth Egyptian dynasty (ca. 3000 BC), but Mr. Birch informs me that pigeons are given a bill of fare in the previous dynasty. In Roman times pigeons fetched a high price. Pliny writes, Nay, they are come to this pass, that they can reckon up their pedigree and race. The court of India’s Akber Khan (ca. 1600) always traveled with at least twenty thousand pigeons. The monarchs of Iran and Turan sent him some very rare birds, and, continues the court historian, His Majesty, by crossing the breeds, which method was never practiced before, has improved them astonishingly. At about this same period, the Dutch were as enthusiastic about pigeons as the Romans had been.

I have kept every pigeon breed that I could purchase or obtain and have been kindly favored with skins from several parts of the world, especially by the Hon. W. Elliot from India and the Hon. C. Murray from Persia. Many treatises in several languages have been written about pigeons, some of them very important because of their age. I have consulted several eminent breeders and joined two of the London Pigeon Clubs. The diversity of breeds is astonishing. Compare the English carrier to the short-faced tumbler and see the wonderful differences in their beaks, with corresponding differences in their skulls. The carrier, especially the male, is remarkable for the carunculated skin about its head and the accompanying greatly elongated eyelids, large nostrils, and widely gaping mouth. The outline of the short-faced tumbler’s beak is like that of the finch’s beak, while the common tumbler has the strictly inherited singular habit of flying at a great height in a compact flock and tumbling in the air head-over-heels. The runt is large with a long, massive beak and big feet. Some runt sub-breeds have long necks; others, long wings and tails; still others, short tails. The barb is related to the carrier but has a short and broad beak instead of a long one. The pouter’s body, wings, and legs are elongated; it glories in inflating its enormously developed crop, which may elicit astonishment and even laughter. The turbit has a short and conical beak, with a line of reversed feathers down its breast and a habit of slightly expanding the upper part of its esophagus. The Jacobin has feathers along the back of the neck that are so reversed that they form a hood; for its size, it has relatively elongated wing and tail feathers. The trumpeter and laugher, as their names suggest, coo very differently from the other breeds. The fantail has thirty or even forty tail feathers – even though the normal number in all members of the pigeon family is twelve or fourteen – and they are kept expanded and carried so erect that in good specimens the head and tail touch; the oil gland is aborted. Several other less distinct breeds could be