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Darwin had been working for several years on his major species book, but was still years away from finishing it. On July 20, 1858, he decided to write an abstract of this work for quicker publication, to be called An Abstract of an Essay on the Origin of Species and Varieties Through Natural Selection, a title the publisher (John Murray) rightly considered too clunky. The full title of the book in its first edition was On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life, but by the sixth edition this had been shortened to just The Origin of Species.
Title Page of the First Edition of On the Origin of Species. Races in the long subtitle (typical of Victorian books) had the meaning of varieties and not the modern meaning. The first use of races in the book is a reference to races of cabbage. Darwin is identified as the author of his journal about the voyage of the Beagle.
The author is a literary man & German scholar. He has read my book attentively; but what is very remarkable, it seems that he is a profound naturalist. He knows my Barnacle book, & appreciates it too highly. Lastly he writes & thinks with uncommon force & clearness; & what is even still rarer his writing is seasoned with most pleasant wit Who can it be? Certainly I should have said that there was only one man in England who could have written this essay & that you were the man. But I suppose that I am wrong, & that there is some hidden genius of great calibre. For how could you influence Jupiter Olympus & make him give 3 columns to pure science. The old Fogies will think the world will come to an end. Well whoever the man is, he has done great service to the cause, far more than by a dozen reviews in common periodicals. If you should happen to be acquainted with the author for Heaven-sake tell me who he is.
5 Separate Theses of Darwins Views of Evolution These are the major theses of what has often been called Darwinism.
(1) Evolution as such. This is the thesis that the world is not constant or recently created nor perpetually cycling but rather is steadily changing and that organisms are transformed in time. (2) Common descent (3) Multiplication of species (4) Gradualism (5) Natural selection
Examples:
A common ancestor of chimpanzees and bonobos (formerly pygmy chimpanzees or hippie chimpanzees) split into these two lines several million years ago. A common ancestor of chimpanzees, bonobos, and humans split about 5 to 7 million years ago. Many splits seem to have occurred on the hominid line, with only one species remaining today, Homo sapiens. The apple maggot (Rhagoletis pomonella) originally fed on hawthorn apples, but in North America has split since 1800 into two species, one feeding on hawthorns and one on apples, which the maggot also found palatable.
(4) Gradualism
(5) Natural selection
(1) Evolution as such (2) Common descent (3) Multiplication of species (4) Gradualism. According to this thesis, evolutionary change takes place through the gradual change of populations and not by the sudden (saltational) production of new individuals that represent a new type. Many of the older Darwinians, including Huxley, preferred saltation; for one thing, it was thought that the Earth was not old enough for such gradual processes to have occurred. (5) Natural selection
Fact 2. The size of populations, except for temporary annual fluctuations, remains stable over time (observed steady-state stability). This we see from our own observations again. A gardener who plants a few tomato plants one year doesnt necessarily find thousands of tomato plants in the garden the next year. A homeowner with a prolific crabapple tree doesnt find tens of thousands of crabapple seedlings on his property the next year (or at least, will get rid of most of them!). Rabbits breed quickly dont usually multiply as wildly as they could, in principle.
Fact 3. The resources available to every species are limited. This point was made by Malthus, but is evident from our own observations, whether we are talking about plants or animals.
Inference 1. There is intense competition (struggle for existence) among the members of a species. This point was made by Malthus with regard to human populations, de Candolle with regard to all organisms. It is most evident for animals, but is also true for plants and bacteria.
Fact 4. No two individuals of a population are exactly the same (population thinking). (This was evident to animal breeders, who deliberately chose the best specimens to propagate, to horticulturists, for the same reason, and, in general, to all taxonomists but not generally appreciated by non-scientists.) Inference 2. Individuals of a population differ from each other in the probability of survival (i.e., natural selection). This inference was made by Darwin, for populations of all types of organisms, but had previously been noted by others, usually with respect to a particular species or variety.
Fact 5. Many of the differences among the individuals of a population are, at least in part, heritable. This information came from animal breeders (breeders of pigeons, livestock, etc.) Inference 3. Natural selection, continued over many generations, results in evolution. This was Darwins conclusion from the five facts and two inferences listed above.
This line of reasoning makes natural selection pretty unavoidable and thus pretty certain. What is not clear is how important it is in nature and at what rate it occurs. Also, there could be other types of selection, and Darwin identified one: sexual selection (to be discussed later).
Pigeons
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. I have associated with several eminent fanciers, and have been permitted to join two of the London Pigeon Clubs.
The diversity of the breeds is something astonishing. Compare the English carrier and the short-faced tumbler, and see the wonderful difference in their beaks, entailing corresponding differences in their skulls. The carrier, more especially the male bird, is also remarkable from the wonderful development of the carunculated skin about the head, and this is accompanied by greatly elongated eyelids, very large external orifices to the nostrils, and a wide gape of mouth. The short-faced tumbler has a beak in outline almost like that of a finch; and the common tumbler has the singular and strictly inherited habit of flying at a great height in a compact flock, and tumbling in the air head over heels.
The runt is a bird of great size, with long, massive beak and large feet; some of the sub-breeds of runts have very long necks, others very long wings and tails, others singularly short tails. The barb is allied to the carrier, but, instead of a very long beak, has a very short and very broad one. The pouter has a much elongated body, wings, and legs; and its enormously developed crop, which it glories in inflating, may well excite astonishment and even laughter. The turbit has a very short and conical beak, with a line of reversed feathers down the breast; and it has the habit of continually expanding slightly the upper part of the oesophagus.
The Jacobin has the feathers so much reversed along the back of the neck that they form a hood, and it has, proportionally to its size, much elongated wing and tail feathers. The trumpeter and laugher, as their names express, utter a very different coo from the other breeds. The fantail has thirty or even forty tailfeathers, instead of twelve or fourteen, the normal number in all members of the great pigeon family; and these feathers are kept expanded, and are carried so erect that in good birds the head and tail touch; the oil-gland is quite aborted. Several other less distinct breeds might have been specified.
Chapter 1 continued
Darwin, who had discussed pigeons with many pigeonbreeders, described their immense variability, but concluded that Great as are the differences between the breeds of the pigeon, I am fully convinced that all are descended from the rock-pigeon Columba livia. Pigeon breeders, who thought they were just perfecting each breed, did not believe Darwin.
Variations in domestic fowl: Hamburg fowl (upper left); Spanish fowl (upper right); Polish fowl (bottom)
Chapter 2 continued Darwin regarded the line between species and varieties as somewhat arbitrary, although variation between species would have to be greater than variation between varieties. But larger samples of organisms of a species exhibited greater variation, so species or genera that extended over greater geographical areas would exhibit greater variation (have more species or subspecies) than those confined to a smaller area.
I look at individual differences, though of small interest to the systematist, as of high importance for us, as being the first step towards such slight varieties as are barely thought worth recording in works on natural history. And I look at varieties which are in any degree more distinct and permanent, as steps leading to more strongly marked and more permanent varieties; and at these latter, as leading to sub-species, and to species. I attribute the passage of a variety, from a state in which it differs very slightly from its parent to one in which it differs more, to the action of natural selection in accumulating (as will hereafter be more fully explained) differences of structure in certain definite directions. Hence I believe a well-marked variety may be justly called an incipient species
Darwin points out that the struggle for existence is easy to overlook or forget about, despite its overwhelming importance:
We behold the face of nature bright with gladness, we often see superabundance of food; we do not see, or we forget, that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always bear in mind, that though food may be now superabundant, it is not so at all seasons of each recurring year.
Owing to this struggle for life, any variation, however slight and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations to other organic beings and to external nature, will tend to the preservation of that individual, and will generally be inherited by its offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term of Natural Selection, in order to mark its relation to man's power of selection.
Chapter 4 contains the only diagram in On the Origin of Species, a tree of life diagram, reproduced below.
Secondly, is it possible that an animal having, for instance, the structure and habits of a bat, could have been formed by the modification of some animal with wholly different habits? Can we believe that natural selection could produce, on the one hand, organs of trifling importance, such as the tail of a giraffe, which serves as a fly-flapper, and, on the other hand, organs of such wonderful structure, as the eye, of which we hardly as yet fully understand the inimitable perfection? Darwin admitted he could not give complete answers to any specific questions of this sort, but that with greater knowledge of present and former organisms the evolution of such traits and organs would be understood.
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; if further, the eye does vary ever so slightly, and the variations be inherited, which is certainly the case; and if any variation or modification in the organ be ever useful to an animal under changing conditions of life, then the difficulty of believing that a perfect and complex eye could be formed by natural selection, though insuperable by our imagination, can hardly be considered real.
Darwin went on to point out numerous example of things that look like steps in the evolution of the eye in a variety of different animals, very close to our current understanding of how, in fact, the eye evolved. He adds,
If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my theory would absolutely break down. But I can find out no such case.
In recent years intelligent design advocates have listed such complex organs, but evolutionary biologists have been successful in determining how they evolved, often from organs that originally served other purposes.
We should be extremely cautious in concluding that an organ could not have been formed by transitional gradations of some kind. Numerous cases could be given amongst the lower animals of the same organ performing at the same time wholly distinct functions.
Darwin cites, as an example, the swimbladder of fish, whose purpose was to permit flotation at different depths in the ocean, but which clearly evolved into the lung of the higher vertebrates, which is used for respiration. I can, indeed, hardly doubt that all vertebrate animals having true lungs have descended by ordinary generation from an ancient prototype, of which we know nothing, furnished with a floating apparatus or swimbladder.
Chapter 7: Instinct
This chapter deals with another difficulty Darwin had stated at the beginning of Chapter 6: Thirdly, can instincts be acquired and modified through natural selection? What shall we say to so marvellous an instinct as that which leads the bee to make cells, which have practically anticipated the discoveries of profound mathematicians? Darwin was writing about the hexagonal shape of the wax cells made by the honeybee. He also mentions birds that lay their eggs in other birds nests (avoiding having to care for them) and certain ants that act as slaves to other ants, and other examples.
Darwin indicated that instinct is difficult to define, but that it related to habitual activities that were not learned. He was sure that instinct was also the result of natural selection, which then acted on behavior as well as physical characteristics. Instinctive behavior would have developed very slowly, over long periods of time.
Many chapters in On the Origin of Species were greatly abridged compared to what they would have been had Darwin written his big species book and not been rushed into writing On the Origin of Species. He returned to instinct in other books, notably The Expression of the Emotions in Man and Animals, since he regarded emotions as instincts that evolved through natural selection.
Chapter 8: Hybridisation
In Chapter 8 Darwin discussed hybridization, the crossing of two species (i.e., one parent from each species). It was thought at the time that if two individuals were of the same species, their offspring would be fertile, but if they were of different species, their offspring would be sterile. The hybrids are usually sterile, and when they are fertile the offspring of two hybrids are usually sterile. Darwin argued that this may just be because of dissimilarities in their reproductive organs, not because the individuals were or were not of the same species or same variety. Often, however, the reasons for sterility were not known. Darwin believed their was no good way to distinguish between species and varieties.
In many cases, intermediate forms were never very numerous or widespread, as natural selection was constantly operating on these forms. Darwin accepted Lyells theory of uniformitarianism, that geological processes had been slowly but constantly acting on the earths surface, in many cases probably removing some of the intermediate forms. Indeed, millions of years of strata have disappeared completely in many parts of the world. [I]t is highly important for us to gain some notion, however imperfect, of the lapse of years. During each of these years, over the whole world, the land and the water has been peopled by hosts of living forms. What an infinite number of generations, which the mind cannot grasp, must have succeeded each other in the long roll of years! Now turn to our richest geological museums, and what a paltry display we behold!
Modern geology was still a young science when On the Origin of Species was published barely 75 years after James Huttons first published work. Since its time there has been much more work carried out and many new fossils discovered. Many intermediate forms have been discovered, but, clearly, not all. Many parts of the world that were not explored in Darwins time have been since. Most remarkably, think of all the paleoanthropological studies that have been carried out on the African continent and that have begun to fill in the tree of life beginning with the common ancestor of chimpanzees and humans. This particular difficulty cited by Darwin is not presently regarded as a difficulty.
4. Darwin believed species originated in one place and then migrated to other places, rather than appearing (being created) differently in different places. 5. Some species migrate easily (birds and plants whose seeds can migrate) while others do not (such as most land animals). 6. Geological processes caused land and water levels to fluctuate, affording opportunities for species to evolve due to the appearance and disappearance of water barriers. Darwin discussed the effect of the ice ages in leading to new and different species and the extinction of some.
Chapter 12 of The Origin of Species continues the discussion of the effect of geographical distribution and separation by focusing on island life. The Galpagos Islands are used as the prime example, because Darwin had visited them on the voyage of the HMS Beagle and written about their species of animals, which exhibited the effects of natural selection. I have carefully searched the oldest voyages, but have not finished my search; as yet I have not found a single instance, free from doubt, of a terrestrial mammal (excluding domesticated animals kept by the natives) inhabiting an island situated above 300 miles from a continent or great continental island; and many islands situated at a much less distance are equally barren.
The evolutionary relationships are evident in homologous structures: What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include the same bones, in the same relative positions? The real affinities of all organic beings are due to inheritance or community of descent. The natural system is a genealogical arrangement, in which we have to discover the lines of descent by the most permanent characters, however slight their vital importance may be.
Although I am fully convinced of the truth of the views given in this volume under the form of an abstract, I by no means expect to convince experienced naturalists whose minds are stocked with a multitude of facts all viewed, during a long course of years, from a point of view directly opposite to mine. It is so easy to hide our ignorance under such expressions as the plan of creation, unity of design, etc., and to think that we give an explanation when we only restate a fact. Any one whose disposition leads him to attach more weight to unexplained difficulties than to the explanation of a certain number of facts will certainly reject my theory. A few naturalists, endowed with much flexibility of mind, and who have already begun to doubt on the immutability of species, may be influenced by this volume; but I look with confidence to the future, to young and rising naturalists, who will be able to view both sides of the question with impartiality. Whoever is led to believe that species are mutable will do good service by conscientiously expressing his conviction; for only thus can the load of prejudice by which this subject is overwhelmed be removed.
Chapter 14 continued
Darwins last chapter refers to the whole book as one long argument. He summarizes his theory and its difficulties, and states that he believes it will lead to a revolution in natural history. The closing paragraph of On the Origin of Species:
It is interesting to contemplate an entangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner, have all been produced by laws acting around us.
Perhaps there were several early life forms maybe some with different genetic material, not the DNA or RNA we are familiar with. But if so, all but one disappeared (unless there are other life forms on Earth, with different genetic material, that we have not yet discovered).
Of the common ancestors of all known life on Earth, the last one before the first is the Last Universal Common Ancestor, or LUCA.
In 1864 Darwin complained that Wallace always spoke of Darwins theory without giving himself credit, and Wallace wrote back:
As to the theory of Natural Selection I shall always maintain it to be yours and yours only. You had worked it out in details I had never thought of, years before I had a ray of light on the subject, and my paper would never have convinced anybody or been noticed as more than an ingenious speculation, whereas your book has revolutionized the study of natural history, and carried away captive the best men of the present age. William Irvine, in Apes, Angels, and Victorians, called Wallace so retiring, so reassuring, so generous.
Darwins only reference to humans apart from occasional mentions like their hand bones in the first edition of On the Origin of Species was this: Light will be thrown on the origin of man and his history. In later editions this became Much light will be thrown He apparently felt there was enough in his book already to disturb many readers without discussing humans but this was the subject most readers were interested in. It was not long, however, before Huxley and Lyell wrote books on this subject: Huxleys Evidence as to Man's Place in Nature and Lyells Geological Evidences of the Antiquity of Man, both appearing in 1863.
Darwin eventually dealt with the topic of humans and their history in The Descent of Man, published in 1871.