The Foundations of the Origin of Species Two Essays written in 1842 and 1844 · Charles Darwin
<GEOGRAPHICAL DISTRIBUTION.>
Sec. VI. Let us consider the absolute state of distribution of organisms of earth's face. Referring chiefly, but not exclusively (from difficulty of transport, fewness, and the distinct characteristics of groups) to Mammalia; and first considering the three or four main [regions] divisions; North America, Europe, Asia, including greater part of E. Indian Archipelago and Africa are intimately allied. Africa most distinct, especially most southern parts. And the Arctic regions, which unite N. America, Asia and Europe, only separated (if we travel one way by Behring's St.) by a narrow strait, is most intimately allied, indeed forms but one restricted group. Next comes S. America,--then Australia, Madagascar (and some small islands which stand very remote from the land). Looking at these main divisions separately, the organisms vary according to changes in condition{122} of different parts. But besides this, barriers of every kind seem to separate regions in a greater degree than proportionally to the difference of climates on each side. Thus great chains of mountains, spaces of sea between islands and continents, even great rivers and deserts. In fact the amount <of> difference in the organisms bears a certain, but not invariable relation to the amount of physical difficulties to transit{123}. {122} In the _Origin_, Ed. i. p. 346, vi. p. 493, the author begins his discussion on geographical distribution by minimising the effect of physical conditions. He lays great stress on the effect of _barriers_, as in the present Essay.
The simple geologist can explain many of the foregoing cases of distribution. Subsidence of a continent in which free means of dispersal, would drive the lowland plants up to the mountains, now converted into islands, and the semi-alpine plants would take place of alpine, and alpine be destroyed, if mountains originally were not of great height. So we may see, during gradual changes{126} of climate on a continent, the propagation of species would vary and adapt themselves to small changes causing much extermination{127}. The mountains of Europe were quite lately covered with ice, and the lowlands probably partaking of the Arctic climate and Fauna. Then as climate changed, arctic fauna would take place of ice, and an inundation of plants from different temperate countries <would> seize the lowlands, leaving islands of arctic forms. But if this had happened on an island, whence could the new forms have come,--here the geologist calls in creationists. If island formed, the geologist will suggest <that> many of the forms might have been borne from nearest land, but if peculiar, he calls in creationist,--as such island rises in height &c., he still more calls in creation. The creationist tells one, on a <illegible> spot the American spirit of creation makes _Orpheus_ and _Tyrannus_ and American doves, and in accordance with past and extinct forms, but no persistent relation between areas and distribution, Geologico-Geograph.-Distribution.
Now according to analogy of domesticated animals let us see what would result. Let us take case of farmer on Pampas, where everything approaches nearer to state of nature. He works on organisms having strong tendency to vary: and he knows <that the> only way to make a distinct breed is to select and separate. It would be useless to separate the best bulls and pair with best cows if their offspring run loose and bred with the other herds, and tendency to reversion not counteracted; he would endeavour therefore to get his cows on islands and then commence his work of selection. If several farmers in different _rincons_{128} were to set to work, especially if with different objects, several breeds would soon be produced. So would it be with horticulturist and so history of every plant shows; the number of varieties{129} increase in proportion to care bestowed on their selection and, with crossing plants, separation. Now, according to this analogy, change of external conditions, and isolation either by chance landing <of> a form on an island, or subsidence dividing a continent, or great chain of mountains, and the number of individuals not being numerous will best favour variation and selection{130}.
No doubt change could be effected in same country without any barrier by long continued selection on one species: even in case of a plant not capable of crossing would easier get possession and solely occupy an island{131}. Now we can at once see that <if> two parts of a continent isolated, new species thus generated in them, would have closest affinities, like cattle in counties of England: if barrier afterwards destroyed one species might destroy the other or both keep their ground. So if island formed near continent, let it be ever so different, that continent would supply inhabitants, and new species (like the old) would be allied with that continent. An island generally very different soil and climate, and number and order of inhabitants supplied by chance, no point so favourable for generation of new species{132},--especially the mountains, hence, so it is. As isolated mountains formed in a plain country (if such happens) is an island. As other islands formed, the old species would spread and thus extend and the fauna of distant island might ultimately meet and a continent formed between them. No one doubts continents formed by repeated elevations and depressions{133}. In looking backwards, but not so far that all geographical boundaries are destroyed, we can thus at once see why existing forms are related to the extinct in the same manner as existing ones are in some part of existing continent. By chance we might even have one or two absolute parent fossils.
If island formed continent, some species would emerge and immigrate. Everyone admits continents. We can see why Galapagos and C. Verde differ <see _Origin_, Ed. i. p. 398>], depressed and raised. We can see from this repeated action and the time required for a continent, why many more forms than in New Zealand <see _Origin_, Ed. i. p. 389 for a comparison between New Zealand and the Cape> no mammals or other classes <see however, _Origin_, Ed. i. p. 393 for the case of the frog>. We can at once see how it comes when there has been an old channel of migration,--Cordilleras; we can see why Indian Asiatic Flora,--[why species] having a wide range gives better chance of some arriving at new points and being selected, and adapted to new ends. I need hardly remark no necessity for change. Finally, as continent (most extinction <?> during formation of continent) is formed after repeated elevation and depression, and interchange of species we might foretell much extinction, and that the survivor would belong to same type, as the extinct, in same manner as different part of same continent, which were once separated by space as they are by time <see _Origin_, Ed. i. pp. 339 and 349>.
As all mammals have descended from one stock, we ought to expect that every continent has been at some time connected, hence obliteration of present ranges. I do not mean that the fossil mammifers found in S. America are the lineal successors <ancestors> of the present forms of S. America: for it is highly improbable that more than one or two cases (who will say how many races after Plata bones) should be found. I believe this from numbers, who have lived,--mere <?> chance of fewness. Moreover in every case from very existence of genera and species only few at one time will leave progeny, under form of new species, to distant ages; and the more distant the ages the fewer the progenitors. An observation may be here appended, bad chance of preservation on rising island, the nurseries of new species, appeal to experience <see _Origin_, Ed. i. p. 292>. This observation may be extended, that in all cases, subsiding land must be, in early stages, less favourable to formation of new species; but it will isolate them, and then if land recommences rising how favourable. As preoccupation is bar to diffusion to species, so would it be to a selected variety.
Looking now to the affinities of organisms, without relation to their distribution, and taking all fossil and recent, we see the degrees of relationship are of different degrees and arbitrary,--sub-genera,--genera,--sub-families, families, orders and classes and kingdoms. The kind of classification which everyone feels is most correct is called the natural system, but no can define this. If we say with Whewell <that we have an> undefined instinct of the importance of organs{135}, we have no means in lower animals of saying which is most important, and yet everyone feels that some one system alone deserves to be called natural. The true relationship of organisms is brought before one by considering relations of analogy, an otter-like animal amongst mammalia and an otter amongst marsupials. In such cases external resemblance and habit of life and _the final end of whole organization_ very strong, yet no relation{136}. Naturalists cannot avoid these terms of relation and affinity though they use them metaphorically. If used in simple earnestness the natural system ought to be a genealogical <one>; and our knowledge of the points which are most easily affected in transmission are those which we least value in considering the natural system, and practically when we find they do vary we regard them of less value{137}.
In classifying varieties the same language is used and the same kind of division: here also (in pine-apple){138} we talk of the natural classification, overlooking similarity of the fruits, because whole plant differs. The origin of sub-genera, genera, &c., &c., is not difficult on notion of genealogical succession, and accords with what we know of similar gradations of affinity in domesticated organisms. In the same region the organic beings are <illegible> related to each other and the external conditions in many physical respects are allied{139} and their differences of same kind, and therefore when a new species has been selected and has obtained a place in the economy of nature, we may suppose that generally it will tend to extend its range during geographical changes, and thus, becoming isolated and exposed to new conditions, will slightly alter and its structure by selection become slightly remodified, thus we should get species of a sub-genus and genus,--as varieties of merino-sheep,--varieties of British and Indian cattle. Fresh species might go on forming and others become extinct and all might become extinct, and then we should have <an> extinct genus; a case formerly mentioned, of which numerous cases occur in Palaeontology.
But more often the same advantages which caused the new species to spread and become modified into several species would favour some of the species being preserved: and if two of the species, considerably different, each gave rise to group of new species, you would have two genera; the same thing will go on. We may look at case in other way, looking to future. According to mere chance every existing species may generate another, but if any species, A, in changing gets an advantage and that advantage (whatever it may be, intellect, &c., &c., or some particular structure or constitution) is inherited{140}, A will be the progenitor of several genera or even families in the hard struggle of nature. A will go on beating out other forms, it might come that A would people earth,--we may now not have one descendant on our globe of the one or several original creations{141}. External conditions air, earth, water being same{142} on globe, and the communication not being perfect, organisms of widely different descent might become adapted to the same end and then we should have cases of analogy{143}, [they might even tend to become numerically representative]. From this often happening each of the great divisions of nature would have their representative eminently adapted to earth, to <air>{144}, to water, and to these in <illegible> and then these great divisions would show numerical relations in their classification.
There is another class of allied and almost identical facts, admitted by the soberest physiologists, [from the study of a certain set of organs in a group of organisms] and refers <? referring> to a unity of type of different organs in the same individual, denominated the science of "Morphology." The <? this> discovered by beautiful and regular series, and in the case of plants from monstrous changes, that certain organs in an individual are other organs metamorphosed. Thus every botanist considers petals, nectaries, stamens, pistils, germen as metamorphosed leaf. They thus explain, in the most lucid manner, the position and number of all parts of the flower, and the curious conversion under cultivation of one part into another. The complicated double set of jaws and palpi of crustaceans{150}, and all insects are considered as metamorphosed <limbs> and to see the series is to admit this phraseology. The skulls of the vertebrates are undoubtedly composed of three metamorphosed vertebrae; thus we can understand the strange form of the separate bones which compose the casket holding man's brain. These{151} facts differ but slightly from those of last section, if with wing, paddle, hand and hoof, some common structure was yet visible, or could be made out by a series of occasional monstrous conversions, and if traces could be discovered of <the> whole having once existed as walking or swimming instruments, these organs would be said to be metamorphosed, as it is they are only said to exhibit a common type.
So we see in grey-hound, bull-dog, in race-horse and cart-horse, which have been selected for their form in full-life, there is much less (?) difference in the few first days after birth{163}, than when full-grown: so in cattle, we see it clearly in cases of cattle, which differ obviously in shape and length of horns. If man were during 10,000 years to be able to select, far more diverse animals from horse or cow, I should expect there would be far less differences in the very young and foetal state: and this, I think, throws light on above marvellous fact. In larvae, which have long life selection, perhaps, does much,--in the pupa not so much{164} There is no object gained in varying form &c. of foetus (beyond certain adaptations to mother's womb) and therefore selection will not further act on it, than in giving to its changing tissues a tendency to certain parts afterwards to assume certain forms. {160} The following: "Deaths of brothers <when> old by same peculiar disease" which is written between the lines seems to have been a memorandum which is expanded a few lines lower. I believe the case of the brothers came from Dr R. W. Darwin. {161} See the discussion to this effect in the _Origin_, Ed. i. pp. 443-4, vi. p. 610. The author there makes the distinction between a cause affecting the germ-cell and the reaction occurring at a late period of life.
From arguments before used, and crossing, we may generally suspect in germ; but I repeat it does not follow, that the change should be apparent till life fully developed; any more than fatness depending on heredity should be apparent during early childhood, still less during foetal existence. In case of horns of cattle, which when inherited must depend on germinal vesicle, obviously no effect till cattle full-grown. Practically it would appear that the [hereditary] peculiarities characterising our domestic races, therefore resulting from vesicle, do not appear with their full characters in very early states; thus though two breeds of cows have calves different, they are not so different,--grey-hound and bull-dog. And this is what is <to> be expected, for man is indifferent to characters of young animals and hence would select those full-grown animals which possessed the desirable characteristics. So that from mere chance we might expect that some of the characters would be such only as became fully apparent in mature life. Furthermore we may suspect it to be a law, that at whatever time a new character appears, whether from vesicle, or effects of external conditions, it would appear at corresponding time <see _Origin_, Ed. i. p. 444>. Thus diseases appearing in old age produce children with d^o.,--early maturity,--longevity,--old men, brothers, of same disease--young children of d^o. I said men do not select for quality of young,--calf with big bullocks. Silk-worms, peculiarities which, appear in caterpillar state or cocoon state, are transmitted to corresponding states. The effect of this would be that if some peculiarity was born in a young animal, but never exercised, it might be inherited in young animal; but if exercised that part of structure would be increased and would be inherited in corresponding time of life after such training.
There is another grand class of facts relating to what are called abortive organs. These consist of organs which the same reasoning power that shows us how beautifully these organs in some cases are adapted to certain end, declares in other cases are absolutely useless. Thus teeth in Rhinoceros{165}, whale, narwhal,--bone on tibia, muscles which do not move,--little bone of wing of Apteryx,--bone representing extremities in some snake,--little wings within <?> soldered cover of beetles,--men and bulls, mammae: filaments without anthers in plants, mere scales representing petals in others, in feather-hyacinth whole flower. Almost infinitely numerous. No one can reflect on these without astonishment, can anything be clearer than that wings are to fly and teeth <to bite>, and yet we find these organs perfect in every detail in situations where they cannot possibly be of their normal use{166}. {165} Some of these examples occur in _Origin_, Ed. i. pp. 450-51, vi. pp. 619-20. {166} The two following sentences are written, one down the margin, the other across the page. "Abortive organs eminently useful in classification. Embryonic state of organs. Rudiments of organs."
The term abortive organ has been thus applied to above structure (as _invariable_ as all other parts{167}) from their absolute similarity to monstrous cases, where from _accident_, certain organs are not developed; as infant without arms or fingers with mere stump representing them: teeth represented by mere points of ossification: headless children with mere button,--viscera represented by small amorphous masses, &c.,--the tail by mere stump,--a solid horn by minute hanging one{168}. There is a tendency in all these cases, when life is preserved, for such structures to become hereditary. We see it in tailless dogs and cats. In plants we see this strikingly,--in Thyme, in _Linum flavum_,--stamen in _Geranium pyrenaicum_{169}. Nectaries abort into petals in Columbine <_Aquilegia_>, produced from some accident and then become hereditary, in some cases only when propagated by buds, in other cases by seed. These cases have been produced suddenly by accident in early growth, but it is part of law of growth that when any organ is not used it tends to diminish (duck's wing{170}?) muscles of dog's ears, <and of> rabbits, muscles wither, arteries grow up. When eye born defective, optic nerve (Tuco Tuco) is atrophied. As every part whether useful or not (diseases, double flowers) tends to be transmitted to offspring, the origin of abortive organs whether produced at the birth or slowly acquired is easily understood in domestic races of organisms: [a struggle between the atrophy and hereditariness. Abortive organs in domestic races.] There will always be a struggle between atrophy of an organ rendered useless, and hereditariness{171}. Because we can understand the origin of abortive organs in certain cases, it would be wrong to conclude absolutely that all must have had same origin, but the strongest analogy is in favour of it. And we can by our theory, for during infinite changes some organ, we might have anticipated, would have become useless. <We can> readily explain the fact, so astounding on any other view, namely that organs possibly useless have been formed often with the same exquisite care as when of vital importance.
Let us recapitulate the whole <?> <of> these latter sections by taking case of the three species of Rhinoceros, which inhabit Java, Sumatra, and mainland of Malacca or India. We find these three close neighbours, occupants of distinct but neighbouring districts, as a group having a different aspect from the Rhinoceros of Africa, though some of these latter inhabit very similar countries, but others most diverse stations. We find them intimately related [scarcely <?> differences more than some breeds of cattle] in structure to the Rhinoceros, which for immense periods have inhabited this one, out of three main zoological divisions of the world. Yet some of these ancient animals were fitted to very different stations: we find all three <illegible> of the generic character of the Rhinoceros, which form a [piece of net]{174} set of links in the broken chain representing the Pachydermata, as the chain likewise forms a portion in other and longer chains. We see this wonderfully in dissecting the coarse leg of all three and finding nearly the same bones as in bat's wings or man's hand, but we see the clear mark in solid tibia of the fusion into it of the fibula. In all three we find their heads composed of three altered vertebrae, short neck, same bones as giraffe. In the upper jaws of all three we find small teeth like rabbit's. In dissecting them in foetal state we find at a not very early stage their form exactly alike the most different animals, and even with arteries running as in a fish: and this similarity holds when the young one is produced in womb, pond, egg or spawn. Now these three undoubted species scarcely differ more than breeds of cattle, are probably subject to many the same contagious diseases; if domesticated these forms would vary, and they might possibly breed together, and fuse into something{175} different <from> their aboriginal forms; might be selected to serve different ends.
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