ON THE GEOGRAPHICAL DISTRIBUTION OF ORGANIC BEINGS IN PAST AND PRESENT TIMES
For convenience sake I shall divide this chapter into three sections. In the first place I shall endeavour to state the laws of the distribution of existing beings, as far as our present object is concerned; in the second, that of extinct; and in the third section I shall consider how far these laws accord with the theory of allied species having a common descent.
 Chapters XI and XII in the Origin, Ed. i., vi. chs. XII and XIII ("On geographical distribution") show signs of having been originally one, in the fact that one summary serves for both. The geological element is not separately treated there, nor is there a separate section on "how far these laws accord with the theory, &c."
In the MS. the author has here written in the margin "If same species appear at two spot at once, fatal to my theory." See Origin, Ed. i. p. 352, vi. p. 499
Distribution of the inhabitants in the different continents.
In the following discussion I shall chiefly refer to terrestrial mammifers, inasmuch as they are better known; their differences in different countries, strongly marked; and especially as the necessary means of their transport are more evident, and confusion, from the accidental conveyance by man of a species from one district to another district, is less likely to arise. It is known that all mammifers (as well as all other organisms) are united in one great system; but that the different species, genera, or families of the same order inhabit different quarters of the globe. If we divide the land into two divisions, according to the amount of difference, and disregarding the numbers of the terrestrial mammifers inhabiting them, we shall have first Australia including New Guinea; and secondly the rest of the world: if we make a three-fold division, we shall have Australia, S. America, and the rest of the world; I must observe that North America is in some respects neutral land, from possessing some S. American forms, but I believe it is more closely allied (as it certainly is in its birds, plants and shells) with Europe. If our division had been four-fold, we should have had Australia, S. America, Madagascar (though inhabited by few mammifers) and the remaining land: if five-fold, Africa, especially the southern eastern parts, would have to be separated from the remainder of the world. These differences in the mammiferous inhabitants of the several main divisions of the globe cannot, it is well known, be explained by corresponding differences in their conditions; how similar are parts of tropical America and Africa; and accordingly we find some analogous resemblances,--thus both have monkeys, both large feline animals, both large Lepidoptera, and large dung-feeding beetles; both have palms and epiphytes; and yet the essential difference between their productions is as great as between those of the arid plains of the Cape of Good Hope and the grass-covered savannahs of La Plata. Consider the distribution of the Marsupialia, which are eminently characteristic of Australia, and in a lesser degree of S. America; when we reflect that animals of this division, feeding both on animal and vegetable matter, frequent the dry open or wooded plains and mountains of Australia, the humid impenetrable forests of New Guinea and Brazil; the dry rocky mountains of Chile, and the grassy plains of Banda Oriental, we must look to some other cause, than the nature of the country, for their absence in Africa and other quarters of the world.
 This division of the land into regions does not occur in the Origin, Ed. i.
 Origin, Ed. i. p. 346, vi. p. 493.
 Opposite this passage is written "not botanically," in Sir J. D. Hooker's hand. The word palms is underlined three times and followed by three exclamation marks. An explanatory note is added in the margin "singular paucity of palms and epiphytes in Trop. Africa compared with Trop. America and Ind. Or." [[East Indies]].
Furthermore it may be observed that all the organisms inhabiting any country are not perfectly adapted to it; I mean by not being perfectly adapted, only that some few other organisms can generally be found better adapted to the country than some of the aborigines. We must admit this when we consider the enormous number of horses and cattle which have run wild during the three last centuries in the uninhabited parts of St Domingo, Cuba, and S. America; for these animals must have supplanted some aboriginal ones. I might also adduce the same fact in Australia, but perhaps it will be objected that 30 or 40 years has not been a sufficient period to test this power of struggling [[with]] and overcoming the aborigines. We know the European mouse is driving before it that of New Zealand, like the Norway rat has driven before it the old English species in England. Scarcely an island can be named, where casually introduced plants have not supplanted some of the native species: in La Plata the Cardoon covers square leagues of country on which some S. American plants must once have grown: the commonest weed over the whole of India is an introduced Mexican poppy. The geologist who knows that slow changes are in progress, replacing land and water, will easily perceive that even if all the organisms of any country had originally been the best adapted to it, this could hardly continue so during succeeding ages without either extermination, or changes, first in the relative proportional numbers of the inhabitants of the country, and finally in their constitutions and structure.
 This partly corresponds to Origin, Ed. i. p. 337, vi. p. 483.
Inspection of a map of the world at once shows that the five divisions, separated according to the greatest amount of difference in the mammifers inhabiting them, are likewise those most widely separated from each other by barriers which mammifers cannot pass: thus Australia is separated from New Guinea and some small adjoining islets only by a narrow and shallow strait; whereas New Guinea and its adjoining islets are cut off from the other East Indian islands by deep water. These latter islands, I may remark, which fall into the great Asiatic group, are separated from each other and the continent only by shallow water; and where this is the case we may suppose, from geological oscillations of level, that generally there has been recent union. South America, including the southern part of Mexico, is cut off from North America by the West Indies, and the great table-land of Mexico, except by a mere fringe of tropical forests along the coast: it is owing, perhaps, to this fringe that N. America possesses some S. American forms. Madagascar is entirely isolated. Africa is also to a great extent isolated, although it approaches, by many promontories and by lines of shallower sea, to Europe and Asia: southern Africa, which is the most distinct in its mammiferous inhabitants, is separated from the northern portion by the Great Sahara Desert and the table-land of Abyssinia. That the distribution of organisms is related to barriers, stopping their progress, we clearly see by comparing the distribution of marine and terrestrial productions. The marine animals being different on the two sides of land tenanted by the same terrestrial animals, thus the shells are wholly different on the opposite sides of the temperate parts of South America, as they are (?) in the Red Sea and the Mediterranean. We can at once perceive that the destruction of a barrier would permit two geographical groups of organisms to fuse and blend into one. But the original cause of groups being different on opposite sides of a barrier can only be understood on the hypothesis of each organism having been created or produced on one spot or area, and afterwards migrating as widely as its means of transport and subsistence permitted it.
 On the general importance of barriers, see Origin, Ed. i. p. 347, vi. p. 494.
 Origin, Ed. i. p. 348, vi. p. 495.
Relation of range in genera and species.
It is generally found, that where a genus or group ranges over nearly the entire world, many of the species composing the group have wide ranges: on the other hand, where a group is restricted to any one country, the species composing it generally have restricted ranges in that country. Thus among mammifers the feline and canine genera are widely distributed, and many of the individual species have enormous ranges [the genus Mus I believe, however, is a strong exception to the rule]. Mr Gould informs me that the rule holds with birds, as in the owl genus, which is mundane, and many of the species range widely. The rule holds also with land and fresh-water mollusca, with butterflies and very generally with plants. As instances of the converse rule, I may give that division of the monkeys which is confined to S. America, and amongst plants, the Cacti, confined to the same continent, the species of both of which have generally narrow ranges. On the ordinary theory of the separate creation of each species, the cause of these relations is not obvious; we can see no reason, because many allied species have been created in the several main divisions of the world, that several of these species should have wide ranges; and on the other hand, that species of the same group should have narrow ranges if all have been created in one main division of the world. As the result of such and probably many other unknown relations, it is found that, even in the same great classes of beings, the different divisions of the world are characterised by either merely different species, or genera, or even families: thus in cats, mice, foxes, S. America differs from Asia and Africa only in species; in her pigs, camels and monkeys the difference is generic or greater. Again, whilst southern Africa and Australia differ more widely in their mammalia than do Africa and S. America, they are more closely (though indeed very distantly) allied in their plants.
 [[Note in original.]] The same laws seem to govern distribution of species and genera, and individuals in time and space. [[See Origin, Ed. i. p. 350, vi. p. 497, also a passage in the last chapter, p. 146.]]
 Origin, Ed. i. p. 404, vi. p. 559.
Distribution of the inhabitants in the same continent.
If we now look at the distribution of the organisms in any one of the above main divisions of the world, we shall find it split up into many regions, with all or nearly all their species distinct, but yet partaking of one common character. This similarity of type in the subdivisions of a great region is equally well-known with the dissimilarity of the inhabitants of the several great regions; but it has been less often insisted on, though more worthy of remark. Thus for instance, if in Africa or S. America, we go from south to north, or from lowland to upland, or from a humid to a dryer part, we find wholly different species of those genera or groups which characterise the continent over which we are passing. In these subdivisions we may clearly observe, as in the main divisions of the world, that sub-barriers divide different groups of species, although the opposite sides of such sub-barriers may possess nearly the same climate, and may be in other respects nearly similar: thus it is on the opposite sides of the Cordillera of Chile, and in a lesser degree on the opposite sides of the Rocky mountains. Deserts, arms of the sea, and even rivers form the barriers; mere preoccupied space seems sufficient in several cases: thus Eastern and Western Australia, in the same latitude, with very similar climate and soils, have scarcely a plant, and few animals or birds, in common, although all belong to the peculiar genera characterising Australia. It is in short impossible to explain the differences in the inhabitants, either of the main divisions of the world, or of these sub-divisions, by the differences in their physical conditions, and by the adaptation of their inhabitants. Some other cause must intervene.
 Origin, Ed. i. p. 349, vi. p. 496.
We can see that the destruction of sub-barriers would cause (as before remarked in the case of the main divisions) two sub-divisions to blend into one; and we can only suppose that the original difference in the species, on the opposite sides of sub-barriers, is due to the creation or production of species in distinct areas, from which they have wandered till arrested by such sub-barriers. Although thus far is pretty clear, it may be asked, why, when species in the same main division of the world were produced on opposite sides of a sub-barrier, both when exposed to similar conditions and when exposed to widely different influences (as on alpine and lowland tracts, as on arid and humid soils, as in cold and hot climates), have they invariably been formed on a similar type, and that type confined to this one division of the world? Why when an ostrich was produced in the southern parts of America, was it formed on the American type, instead of on the African or on Australian types? Why when hare-like and rabbit-like animals were formed to live on the Savannahs of La Plata, were they produced on the peculiar Rodent type of S. America, instead of on the true hare-type of North America, Asia and Africa? Why when borrowing Rodents, and camel-like animals were formed to tenant the Cordillera, were they formed on the same type with their representatives on the plains? Why were the mice, and many birds of different species on the opposite sides of the Cordillera, but exposed to a very similar climate and soil, created on the same peculiar S. American type? Why were the plants in Eastern and Western Australia, though wholly different as species, formed on the same peculiar Australian types? The generality of the rule, in so many places and under such different circumstances, makes it highly remarkable and seems to demand some explanation.
 The case of the ostrich (Rhea) occurs in the Origin, Ed. i. p. 349, vi. p. 496.
 [[Note in original.]] There is a hare in S. America,--so bad example.
 See Origin, Ed. i. p. 349, vi. p. 497.
If we now look to the character of the inhabitants of small islands, we shall find that those situated close to other land have a similar fauna with that land, whilst those at a considerable distance from other land often possess an almost entirely peculiar fauna. The Galapagos Archipelago is a remarkable instance of this latter fact; here almost every bird, its one mammifer, its reptiles, land and sea shells, and even fish, are almost all peculiar and distinct species, not found in any other quarter of the world: so are the majority of its plants. But although situated at the distance of between 500 and 600 miles from the S. American coast, it is impossible to even glance at a large part of its fauna, especially at the birds, without at once seeing that they belong to the American type. Hence, in fact, groups of islands thus circumstanced form merely small but well-defined sub-divisions of the larger geographical divisions. But the fact is in such cases far more striking: for taking the Galapagos Archipelago as an instance; in the first place we must feel convinced, seeing that every island is wholly volcanic and bristles with craters, that in a geological sense the whole is of recent origin comparatively with a continent; and as the species are nearly all peculiar, we must conclude that they have in the same sense recently been produced on this very spot; and although in the nature of the soil, and in a lesser degree in the climate, there is a wide difference with the nearer part of the S. American coast, we see that the inhabitants have been formed on the same closely allied type. On the other hand, these islands, as far as their physical conditions are concerned, resemble closely the Cape de Verde volcanic group, and yet how wholly unlike are the productions of these two archipelagoes. The Cape de Verde group, to which may be added the Canary Islands, are allied in their inhabitants (of which many are peculiar species) to the coast of Africa and southern Europe, in precisely the same manner as the Galapagos Archipelago is allied to America. We here clearly see that mere geographical proximity affects, more than any relation of adaptation, the character of species. How many islands in the Pacific exist far more like in their physical conditions to Juan Fernandez than this island is to the coast of Chile, distant 300 miles; why then, except from mere proximity, should this island alone be tenanted by two very peculiar species of humming-birds--that form of birds which is so exclusively American? Innumerable other similar cases might be adduced.
 For the general problem of Oceanic Islands, see Origin, Ed. i. p. 388, vi. p. 541.
 This is an illustration of the general theory of barriers (Origin, Ed. i. p. 347, vi. p. 494). At i. p. 391, vi. p. 544 the question is discussed from the point of view of means of transport. Between the lines, above the words "with that land," the author wrote "Cause, formerly joined, no one doubts after Lyell."
 Origin, Ed. i. p. 390, vi. p. 543.
 See Origin, Ed. i. p. 397, vi. p. 552.
 The Cape de Verde and Galapagos Archipelagoes are compared in the Origin, Ed. i. p. 398, vi. p. 553. See also Journal of Researches, 1860, p. 393.
The Galapagos Archipelago offers another, even more remarkable, example of the class of facts we are here considering. Most of its genera are, as we have said, American, many of them are mundane, or found everywhere, and some are quite or nearly confined to this archipelago. The islands are of absolutely similar composition, and exposed to the same climate; most of them are in sight of each other; and yet several of the islands are inhabited, each by peculiar species (or in some cases perhaps only varieties) of some of the genera characterising the archipelago. So that the small group of the Galapagos Islands typifies, and follows exactly the same laws in the distribution of its inhabitants, as a great continent. How wonderful it is that two or three closely similar but distinct species of a mocking-thrush should have been produced on three neighbouring and absolutely similar islands; and that these three species of mocking-thrush should be closely related to the other species inhabiting wholly different climates and different districts of America, and only in America. No similar case so striking as this of the Galapagos Archipelago has hitherto been observed; and this difference of the productions in the different islands may perhaps be partly explained by the depth of the sea between them (showing that they could not have been united within recent geological periods), and by the currents of the sea sweeping straight between them,--and by storms of wind being rare, through which means seeds and birds could be blown, or drifted, from one island to another. There are however some similar facts: it is said that the different, though neighbouring islands of the East Indian Archipelago are inhabited by some different species of the same genera; and at the Sandwich group some of the islands have each their peculiar species of the same genera of plants.
 In the Origin, Ed. i. p. 390, a strong point is made of birds which immigrated "with facility and in a body" not having been modified. Thus the author accounts for the small percentage of peculiar "marine birds."
Islands standing quite isolated within the intra-tropical oceans have generally very peculiar floras, related, though feebly (as in the case of St Helena where almost every species is distinct), with the nearest continent: Tristan d'Acunha is feebly related, I believe, in its plants, both to Africa and S. America, not by having species in common, but by the genera to which they belong. The floras of the numerous scattered islands of the Pacific are related to each other and to all the surrounding continents; but it has been said, that they have more of an Indo-Asiatic than American character. This is somewhat remarkable, as America is nearer to all the Eastern islands, and lies in the direction of the trade-wind and prevailing currents; on the other hand, all the heaviest gales come from the Asiatic side. But even with the aid of these gales, it is not obvious on the ordinary theory of creation how the possibility of migration (without we suppose, with extreme improbability, that each species with an Indo-Asiatic character has actually travelled from the Asiatic shores, where such species do not now exist) explains this Asiatic character in the plants of the Pacific. This is no more obvious than that (as before remarked) there should exist a relation between the creation of closely allied species in several regions of the world, and the fact of many such species having wide ranges; and on the other hand, of allied species confined to one region of the world having in that region narrow ranges.
 "The affinities of the St Helena flora are strongly South African." Hooker's Lecture on Insular Floras in the Gardeners' Chronicle, Jan. 1867.
 It is impossible to make out the precise form which the author intended to give to this sentence, but the meaning is clear.
 This is no doubt true, the flora of the Sandwich group however has marked American affinities.
We will now turn to the floras of mountain-summits which are well known to differ from the floras of the neighbouring lowlands. In certain characters, such as dwarfness of stature, hairiness, &c., the species from the most distant mountains frequently resemble each other,--a kind of analogy like that for instance of the succulency of most desert plants. Besides this analogy, Alpine plants present some eminently curious facts in their distribution. In some cases the summits of mountains, although immensely distant from each other, are clothed by the same identical species which are likewise the same with those growing on the likewise very distant Arctic shores. In other cases, although few or none of the species may be actually identical, they are closely related; whilst the plants of the lowland districts surrounding the two mountains in question will be wholly dissimilar. As mountain-summits, as far as their plants are concerned, are islands rising out of an ocean of land in which the Alpine species cannot live, nor across which is there any known means of transport, this fact appears directly opposed to the conclusion which we have come to from considering the general distribution of organisms both on continents and on islands--namely, that the degree of relationship between the inhabitants of two points depends on the completeness and nature of the barriers between those points. I believe, however, this anomalous case admits, as we shall presently see, of some explanation. We might have expected that the flora of a mountain summit would have presented the same relation to the flora of the surrounding lowland country, which any isolated part of a continent does to the whole, or an island does to the mainland, from which it is separated by a rather wide space of sea. This in fact is the case with the plants clothing the summits of some mountains, which mountains it may be observed are particularly isolated; for instance, all the species are peculiar, but they belong to the forms characteristic of the surrounding continent, on the mountains of Caraccas, of Van Dieman's Land and of the Cape of Good Hope. On some other mountains, for instance [[in]] Tierra del Fuego and in Brazil, some of the plants though distinct species are S. American forms; whilst others are allied to or are identical with the Alpine species of Europe. In islands of which the lowland flora is distinct [[from]] but allied to that of the nearest continent, the Alpine plants are sometimes (or perhaps mostly) eminently peculiar and distinct; this is the case on Teneriffe, and in a lesser degree even on some of the Mediterranean islands.
 See Origin, Ed. i. p. 365, vi. p. 515. The present discussion was written before the publication of Forbes' celebrated paper on the same subject; see Life and Letters, vol. I. p. 88.
 The apparent breakdown of the doctrine of barriers is slightly touched on in the Origin, Ed. i. p. 365, vi. p. 515.
 In the Origin, Ed. i. p. 375, vi. p. 526, the author points out that on the mountains at the Cape of Good Hope "some few representative European forms are found, which have not been discovered in the inter-tropical parts of Africa."
 See Hooker's Lecture on Insular Floras in the Gardeners' Chronicle, Jan. 1867.
If all Alpine floras had been characterised like that of the mountain of Caraccas, or of Van Dieman's Land, &c., whatever explanation is possible of the general laws of geographical distribution would have applied to them. But the apparently anomalous case just given, namely of the mountains of Europe, of some mountains in the United States (Dr Boott) and of the summits of the Himalaya (Royle), having many identical species in common conjointly with the Arctic regions, and many species, though not identical, closely allied, require a separate explanation. The fact likewise of several of the species on the mountains of Tierra del Fuego (and in a lesser degree on the mountains of Brazil) not belonging to American forms, but to those of Europe, though so immensely remote, requires also a separate explanation.
Cause of the similarity in the floras of some distant mountains.
Now we may with confidence affirm, from the number of the then floating icebergs and low descent of the glaciers, that within a period so near that species of shells have remained the same, the whole of Central Europe and of North America (and perhaps of Eastern Asia) possessed a very cold climate; and therefore it is probable that the floras of these districts were the same as the present Arctic one,--as is known to have been to some degree the case with then existing sea-shells, and those now living on the Arctic shores. At this period the mountains must have been covered with ice of which we have evidence in the surfaces polished and scored by glaciers. What then would be the natural and almost inevitable effects of the gradual change into the present more temperate climate? The ice and snow would disappear from the mountains, and as new plants from the more temperate regions of the south migrated northward, replacing the Arctic plants, these latter would crawl up the now uncovered mountains, and likewise be driven northward to the present Arctic shores. If the Arctic flora of that period was a nearly uniform one, as the present one is, then we should have the same plants on these mountain-summits and on the present Arctic shores. On this view the Arctic flora of that period must have been a widely extended one, more so than even the present one; but considering how similar the physical conditions must always be of land bordering on perpetual frost, this does not appear a great difficulty; and may we not venture to suppose that the almost infinitely numerous icebergs, charged with great masses of rocks, soil and brushwood and often driven high up on distant beaches, might have been the means of widely distributing the seeds of the same species?
 In the margin the author has written "(Forbes)." This may have been inserted at a date later than 1844, or it may refer to a work by Forbes earlier than his Alpine paper.
 See Origin, Ed. i. p. 367, vi. p. 517.
 [[Note in original.]] Perhaps vitality checked by cold and so prevented germinating. [[On the carriage of seeds by icebergs, see Origin, Ed. i. p. 363, vi. p. 513.]]
I will only hazard one other observation, namely that during the change from an extremely cold climate to a more temperate one the conditions, both on lowland and mountain, would be singularly favourable for the diffusion of any existing plants, which could live on land, just freed from the rigour of eternal winter; for it would possess no inhabitants; and we cannot doubt that preoccupation is the chief bar to the diffusion of plants. For amongst many other facts, how otherwise can we explain the circumstance that the plants on the opposite, though similarly constituted sides of a wide river in Eastern Europe (as I was informed by Humboldt) should be widely different; across which river birds, swimming quadrupeds and the wind must often transport seeds; we can only suppose that plants already occupying the soil and freely seeding check the germination of occasionally transported seeds.
 A note by the author gives "many authors" apparently as authority for this statement.
At about the same period when icebergs were transporting boulders in N. America as far as 36░ south, where the cotton tree now grows in South America, in latitude 42░ (where the land is now clothed with forests having an almost tropical aspect with the trees bearing epiphytes and intertwined with canes), the same ice action was going on; is it not then in some degree probable that at this period the whole tropical parts of the two Americas possessed (as Falconer asserts that India did) a more temperate climate? In this case the Alpine plants of the long chain of the Cordillera would have descended much lower and there would have been a broad high-road connecting those parts of North and South America which were then frigid. As the present climate supervened, the plants occupying the districts which now are become in both hemispheres temperate and even semi-tropical must have been driven to the Arctic and Antarctic regions; and only a few of the loftiest points of the Cordillera can have retained their former connecting flora. The transverse chain of Chiquitos might perhaps in a similar manner during the ice-action period have served as a connecting road (though a broken one) for Alpine plants to become dispersed from the Cordillera to the highlands of Brazil. It may be observed that some (though not strong) reasons can be assigned for believing that at about this same period the two Americas were not so thoroughly divided as they now are by the West Indies and tableland of Mexico. I will only further remark that the present most singularly close similarity in the vegetation of the lowlands of Kerguelen's Land and of Tierra del Fuego (Hooker), though so far apart, may perhaps be explained by the dissemination of seeds during this same cold period, by means of icebergs, as before alluded to.
 Opposite to this passage, in the margin, the author has written:--"too hypothetical."
 The Cordillera is described as supplying a great line of invasion in the Origin, Ed. i. p. 378.
 This is an approximation to the author's views on trans-tropical migration (Origin, Ed. i. pp. 376-8). See Thiselton-Dyer's interesting discussion in Darwin and Modern Science, p. 304.
 See Hooker's Lecture on Insular Floras in the Gardeners' Chronicle, Jan. 1867.
 [[Note by the author.]] Similarity of flora of coral islands easily explained.
Finally, I think we may safely grant from the foregoing facts and reasoning that the anomalous similarity in the vegetation of certain very distant mountain-summits is not in truth opposed to the conclusion of the intimate relation subsisting between proximity in space (in accordance with the means of transport in each class) and the degree of affinity of the inhabitants of any two countries. In the case of several quite isolated mountains, we have seen that the general law holds good.
Whether the same species has been created more than once.
As the fact of the same species of plants having been found on mountain-summits immensely remote has been one chief cause of the belief of some species having been contemporaneously produced or created at two different points, I will here briefly discuss this subject. On the ordinary theory of creation, we can see no reason why on two similar mountain-summits two similar species may not have been created; but the opposite view, independently of its simplicity, has been generally received from the analogy of the general distribution of all organisms, in which (as shown in this chapter) we almost always find that great and continuous barriers separate distinct series; and we are naturally led to suppose that the two series have been separately created. When taking a more limited view we see a river, with a quite similar country on both sides, with one side well stocked with a certain animal and on the other side not one (as is the case with the Bizcacha on the opposite sides of the Plata), we are at once led to conclude that the Bizcacha was produced on some one point or area on the western side of the river. Considering our ignorance of the many strange chances of diffusion by birds (which occasionally wander to immense distances) and quadrupeds swallowing seeds and ova (as in the case of the flying water-beetle which disgorged the eggs of a fish), and of whirlwinds carrying seeds and animals into strong upper currents (as in the case of volcanic ashes and showers of hay, grain and fish), and of the possibility of species having survived for short periods at intermediate spots and afterwards becoming extinct there; and considering our knowledge of the great changes which have taken place from subsidence and elevation in the surface of the earth, and of our ignorance of the greater changes which may have taken place, we ought to be very slow in admitting the probability of double creations. In the case of plants on mountain-summits, I think I have shown how almost necessarily they would, under the past conditions of the northern hemisphere, be as similar as are the plants on the present Arctic shores; and this ought to teach us a lesson of caution.
 On centres of creation see Origin, Ed. i. p. 352, vi. p. 499.
 In the Journal of Researches, Ed. 1860, p. 124, the distribution of the Bizcacha is described as limited by the river Uruguay. The case is not I think given in the Origin.
 In the Origin, Ed. i. a special section (p. 356, vi. p. 504) is devoted to Means of Dispersal. The much greater prominence given to this subject in the Origin is partly accounted for by the author's experiments being of later date, i.e. 1855 (Life and Letters, vol. II. p. 53). The carriage of fish by whirlwinds is given in the Origin, Ed. i. p. 384, vi. p. 536.
 The case of islands serving as halting places is given in the Origin, Ed. i. p. 357, vi. p. 505. But here the evidence of this having occurred is supposed to be lost by the subsidence of the islands, not merely by the extinction of the species.
But the strongest argument against double creations may be drawn from considering the case of mammifers in which, from their nature and from the size of their offspring, the means of distribution are more in view. There are no cases where the same species is found in very remote localities, except where there is a continuous belt of land: the Arctic region perhaps offers the strongest exception, and here we know that animals are transported on icebergs. The cases of lesser difficulty may all receive a more or less simple explanation; I will give only one instance; the nutria, I believe, on the eastern coast of S. America live exclusively in fresh-water rivers, and I was much surprised how they could have got into rivulets, widely apart, on the coast of Patagonia; but on the opposite coast I found these quadrupeds living exclusively in the sea, and hence their migration along the Patagonian coast is not surprising. There is no case of the same mammifer being found on an island far from the coast, and on the mainland, as happens with plants. On the idea of double creations it would be strange if the same species of several plants should have been created in Australia and Europe; and no one instance of the same species of mammifer having been created, or aboriginally existing, in two as nearly remote and equally isolated points. It is more philosophical, in such cases, as that of some plants being found in Australia and Europe, to admit that we are ignorant of the means of transport. I will allude only to one other case, namely, that of the Mydas, an Alpine animal, found only on the distant peaks of the mountains of Java: who will pretend to deny that during the ice period of the northern and southern hemispheres, and when India is believed to have been colder, the climate might not have permitted this animal to haunt a lower country, and thus to have passed along the ridges from summit to summit? Mr Lyell has further observed that, as in space, so in time, there is no reason to believe that after the extinction of a species, the self-same form has ever reappeared. I think, then, we may, notwithstanding the many cases of difficulty, conclude with some confidence that every species has been created or produced on a single point or area.
 "We find no inexplicable cases of the same mammal inhabiting distant points of the world." Origin, Ed. i. p. 352, vi. p. 500. See also Origin, Ed. i. p. 393, vi. p. 547.
 [[Note by the author.]] Many authors.
 Nutria is the Spanish for otter, and is now a synonym for Lutra. The otter on the Atlantic coast is distinguished by minute differences from the Pacific species. Both forms are said to take to the sea. In fact the case presents no especial difficulties.
 In Origin, Ed. i. p. 394, vi. p. 548, bats are mentioned as an explicable exception to this statement.
 This reference is doubtless to Mydaus, a badger-like animal from the mountains of Java and Sumatra (Wallace, Geographical Distribution, ii. p. 199). The instance does not occur in the Origin but the author remarks (Origin, Ed. i. p. 376, vi. p. 527) that cases, strictly analogous to the distribution of plants, occur among terrestrial mammals.
 See Origin, Ed. i. p. 313, vi. p. 454.
On the number of species, and of the classes to which they belong in different regions.
The last fact in geographical distribution, which, as far as I can see, in any way concerns the origin of species, relates to the absolute number and nature of the organic beings inhabiting different tracts of land. Although every species is admirably adapted (but not necessarily better adapted than every other species, as we have seen in the great increase of introduced species) to the country and station it frequents; yet it has been shown that the entire difference between the species in distant countries cannot possibly be explained by the difference of the physical conditions of these countries. In the same manner, I believe, neither the number of the species, nor the nature of the great classes to which they belong, can possibly in all cases be explained by the conditions of their country. New Zealand, a linear island stretching over about 700 miles of latitude, with forests, marshes, plains and mountains reaching to the limits of eternal snow, has far more diversified habitats than an equal area at the Cape of Good Hope; and yet, I believe, at the Cape of Good Hope there are, of phanerogamic plants, from five to ten times the number of species as in all New Zealand. Why on the theory of absolute creations should this large and diversified island only have from 400 to 500 (? Dieffenbach) phanerogamic plants? and why should the Cape of Good Hope, characterised by the uniformity of its scenery, swarm with more species of plants than probably any other quarter of the world? Why on the ordinary theory should the Galapagos Islands abound with terrestrial reptiles? and why should many equal-sized islands in the Pacific be without a single one or with only one or two species? Why should the great island of New Zealand be without one mammiferous quadruped except the mouse, and that was probably introduced with the aborigines? Why should not one island (it can be shown, I think, that the mammifers of Mauritius and St Iago have all been introduced) in the open ocean possess a mammiferous quadruped? Let it not be said that quadrupeds cannot live in islands, for we know that cattle, horses and pigs during a long period have run wild in the West Indian and Falkland Islands; pigs at St Helena; goats at Tahiti; asses in the Canary Islands; dogs in Cuba; cats at Ascension; rabbits at Madeira and the Falklands; monkeys at St Iago and the Mauritius; even elephants during a long time in one of the very small Sooloo Islands; and European mice on very many of the smallest islands far from the habitations of man. Nor let it be assumed that quadrupeds are more slowly created and hence that the oceanic islands, which generally are of volcanic formation, are of too recent origin to possess them; for we know (Lyell) that new forms of quadrupeds succeed each other quicker than Mollusca or Reptilia. Nor let it be assumed (though such an assumption would be no explanation) that quadrupeds cannot be created on small islands; for islands not lying in mid-ocean do possess their peculiar quadrupeds; thus many of the smaller islands of the East Indian Archipelago possess quadrupeds; as does Fernando Po on the West Coast of Africa; as the Falkland Islands possess a peculiar wolf-like fox; so do the Galapagos Islands a peculiar mouse of the S. American type. These two last are the most remarkable cases with which I am acquainted; inasmuch as the islands lie further from other land. It is possible that the Galapagos mouse may have been introduced in some ship from the S. American coast (though the species is at present unknown there), for the aboriginal species soon haunts the goods of man, as I noticed in the roof of a newly erected shed in a desert country south of the Plata. The Falkland Islands, though between 200 and 300 miles from the S. American coast, may in one sense be considered as intimately connected with it; for it is certain that formerly many icebergs loaded with boulders were stranded on its southern coast, and the old canoes which are occasionally now stranded, show that the currents still set from Tierra del Fuego. This fact, however, does not explain the presence of the Canis antarcticus on the Falkland Islands, unless we suppose that it formerly lived on the mainland and became extinct there, whilst it survived on these islands, to which it was borne (as happens with its northern congener, the common wolf) on an iceberg, but this fact removes the anomaly of an island, in appearance effectually separated from other land, having its own species of quadruped, and makes the case like that of Java and Sumatra, each having their own rhinoceros.
 The comparison between New Zealand and the Cape is given in the Origin, Ed. i. p. 389, vi. p. 542.
 In a corresponding discussion in the Origin, Ed. i. p. 393, vi. p. 546, stress is laid on the distribution of Batrachians not of reptiles.
 The whole argument is given--more briefly than here--in the Origin, Ed. i. p. 394, vi. p. 547.
 See Origin, Ed i. p. 393, vi. p. 547. The discussion is much fuller in the present Essay.
Before summing up all the facts given in this section on the present condition of organic beings, and endeavouring to see how far they admit of explanation, it will be convenient to state all such facts in the past geographical distribution of extinct beings as seem anyway to concern the theory of descent.
Geographical distribution of extinct organisms.
I have stated that if the land of the entire world be divided into (we will say) three sections, according to the amount of difference of the terrestrial mammifers inhabiting them, we shall have three unequal divisions of (1st) Australia and its dependent islands, (2nd) South America, (3rd) Europe, Asia and Africa. If we now look to the mammifers which inhabited these three divisions during the later Tertiary periods, we shall find them almost as distinct as at the present day, and intimately related in each division to the existing forms in that division. This is wonderfully the case with the several fossil Marsupial genera in the caverns of New South Wales and even more wonderfully so in South America, where we have the same peculiar group of monkeys, of a guanaco-like animal, of many rodents, of the Marsupial Didelphys, of Armadilloes and other Edentata. This last family is at present very characteristic of S. America, and in a late Tertiary epoch it was even more so, as is shown by the numerous enormous animals of the Megatheroid family, some of which were protected by an osseous armour like that, but on a gigantic scale, of the recent Armadillo. Lastly, over Europe the remains of the several deer, oxen, bears, foxes, beavers, field-mice, show a relation to the present inhabitants of this region; and the contemporaneous remains of the elephant, rhinoceros, hippopotamus, hyŠna, show a relation with the grand Africo-Asiatic division of the world. In Asia the fossil mammifers of the Himalaya (though mingled with forms long extinct in Europe) are equally related to the existing forms of the Africo-Asiatic division; but especially to those of India itself. As the gigantic and now extinct quadrupeds of Europe have naturally excited more attention than the other and smaller remains, the relation between the past and the present mammiferous inhabitants of Europe has not been sufficiently attended to. But in fact the mammifers of Europe are at present nearly as much Africo-Asiatic as they were formerly when Europe had its elephants and rhinoceroses, etc.; Europe neither now nor then possessed peculiar groups as does Australia and S. America. The extinction of certain peculiar forms in one quarter does not make the remaining mammifers of that quarter less related to its own great division of the world: though Tierra del Fuego possesses only a fox, three rodents, and the guanaco, no one (as these all belong to S. American types, but not to the most characteristic forms) would doubt for one minute
 See Origin, Ed. i. p. 339, vi. p. 485.
 In the Origin, Ed. i. p. 339, vi. p. 485, which corresponds to this part of the present Essay, the author does not make a separate section for such cases as the occurrence of fossil Marsupials in Europe (Origin, Ed. i. p. 340, vi. p. 486) as he does in the present Essay; see the section on Changes in geographical distribution, p. 177.
We find some evidence of the same general fact in a relation between the recent and the Tertiary sea-shells, in the different main divisions of the marine world.
This general and most remarkable relation between the lately past and present mammiferous inhabitants of the three main divisions of the world is precisely the same kind of fact as the relation between the different species of the several sub-regions of any one of the main divisions. As we usually associate great physical changes with the total extinction of one series of beings, and its succession by another series, this identity of relation between the past and the present races of beings in the same quarters of the globe is more striking than the same relation between existing beings in different sub-regions: but in truth we have no reason for supposing that a change in the conditions has in any of these cases supervened, greater than that now existing between the temperate and tropical, or between the highlands and lowlands of the same main divisions, now tenanted by related beings. Finally, then, we clearly see that in each main division of the world the same relation holds good between its inhabitants in time as over space.
 "We can understand how it is that all the forms of life, ancient and recent, make together one grand system; for all are connected by generation." Origin, Ed. i. p. 344, vi. p. 491.
Changes in geographical distribution.
If, however, we look closer, we shall find that even Australia, in possessing a terrestrial Pachyderm, was so far less distinct from the rest of the world than it now is; so was S. America in possessing the Mastodon, horse, [hyŠna,] and antelope. N. America, as I have remarked, is now, in its mammifers, in some respects neutral ground between S. America and the great Africo-Asiatic division; formerly, in possessing the horse, Mastodon and three Megatheroid animals, it was more nearly related to S. America; but in the horse and Mastodon, and likewise in having the elephant, oxen, sheep, and pigs, it was as much, if not more, related to the Africo-Asiatic division. Again, northern India was much more closely related (in having the giraffe, hippopotamus, and certain musk-deer) to southern Africa than it now is; for southern and eastern Africa deserve, if we divide the world into five parts, to make one division by itself. Turning to the dawn of the Tertiary period, we must, from our ignorance of other portions of the world, confine ourselves to Europe; and at that period, in the presence of Marsupials and Edentata, we behold an entire blending of those mammiferous forms which now eminently characterise Australia and S. America.
 The word hyŠna is erased. There appear to be no fossil HyŠnidŠ in S. America.
 See note 1, p. 175, also Origin, Ed. i. p. 340, vi. p. 486.
 [[Note by the author.]] And see Eocene European mammals in N. America.
If we now look at the distribution of sea-shells, we find the same changes in distribution. The Red Sea and the Mediterranean were more nearly related in these shells than they now are. In different parts of Europe, on the other hand, during the Miocene period, the sea-shells seem to have been more different than at present. In the Tertiary period, according to Lyell, the shells of N. America and Europe were less related than at present, and during the Cretaceous still less like; whereas, during this same Cretaceous period, the shells of India and Europe were more like than at present. But going further back to the Carbonaceous period, in N. America and Europe, the productions were much more like than they now are. These facts harmonise with the conclusions drawn from the present distribution of organic beings, for we have seen, that from species being created in different points or areas, the formation of a barrier would cause or make two distinct geographical areas; and the destruction of a barrier would permit their diffusion. And as long-continued geological changes must both destroy and make barriers, we might expect, the further we looked backwards, the more changed should we find the present distribution. This conclusion is worthy of attention; because, finding in widely different parts of the same main division of the world, and in volcanic islands near them, groups of distinct, but related, species;--and finding that a singularly analogous relation holds good with respect to the beings of past times, when none of the present species were living, a person might be tempted to believe in some mystical relation between certain areas of the world, and the production of certain organic forms; but we now see that such an assumption would have to be complicated by the admission that such a relation, though holding good for long revolutions of years, is not truly persistent.
 [[Note by the author.]] All this requires much verification.
 This point seems to be less insisted on in the Origin.
 Origin, Ed. i. p. 356, vi. p. 504.
I will only add one more observation to this section. Geologists finding in the most remote period with which we are acquainted, namely in the Silurian period, that the shells and other marine productions in North and South America, in Europe, Southern Africa, and Western Asia, are much more similar than they now are at these distant points, appear to have imagined that in these ancient times the laws of geographical distribution were quite different than what they now are: but we have only to suppose that great continents were extended east and west, and thus did not divide the inhabitants of the temperate and tropical seas, as the continents now do; and it would then become probable that the inhabitants of the seas would be much more similar than they now are. In the immense space of ocean extending from the east coast of Africa to the eastern islands of the Pacific, which space is connected either by lines of tropical coast or by islands not very distant from each other, we know (Cuming) that many shells, perhaps even as many as 200, are common to the Zanzibar coast, the Philippines, and the eastern islands of the Low or Dangerous Archipelago in the Pacific. This space equals that from the Arctic to the Antarctic pole! Pass over the space of quite open ocean, from the Dangerous Archipelago to the west coast of S. America, and every shell is different: pass over the narrow space of S. America, to its eastern shores, and again every shell is different! Many fish, I may add, are also common to the Pacific and Indian Oceans.
 [[Note by the author.]] D'Orbigny shows that this is not so.
Summary on the distribution of living and extinct organic beings.
Let us sum up the several facts now given with respect to the past and present geographical distribution of organic beings. In a previous chapter it was shown that species are not exterminated by universal catastrophes, and that they are slowly produced: we have also seen that each species is probably only once produced, on one point or area once in time; and that each diffuses itself, as far as barriers and its conditions of life permit. If we look at any one main division of the land, we find in the different parts, whether exposed to different conditions or to the same conditions, many groups of species wholly or nearly distinct as species, nevertheless intimately related. We find the inhabitants of islands, though distinct as species, similarly related to the inhabitants of the nearest continent; we find in some cases, that even the different islands of one such group are inhabited by species distinct, though intimately related to one another and to those of the nearest continent:--thus typifying the distribution of organic beings over the whole world. We find the floras of distant mountain-summits either very similar (which seems to admit, as shown, of a simple explanation) or very distinct but related to the floras of the surrounding region; and hence, in this latter case, the floras of two mountain-summits, although exposed to closely similar conditions, will be very different. On the mountain-summits of islands, characterised by peculiar faunas and floras, the plants are often eminently peculiar. The dissimilarity of the organic beings inhabiting nearly similar countries is best seen by comparing the main divisions of the world; in each of which some districts may be found very similarly exposed, yet the inhabitants are wholly unlike;--far more unlike than those in very dissimilar districts in the same main division. We see this strikingly in comparing two volcanic archipelagoes, with nearly the same climate, but situated not very far from two different continents; in which case their inhabitants are totally unlike. In the different main divisions of the world, the amount of difference between the organisms, even in the same class, is widely different, each main division having only the species distinct in some families, in other families having the genera distinct. The distribution of aquatic organisms is very different from that of the terrestrial organisms; and necessarily so, from the barriers to their progress being quite unlike. The nature of the conditions in an isolated district will not explain the number of species inhabiting it; nor the absence of one class or the presence of another class. We find that terrestrial mammifers are not present on islands far removed from other land. We see in two regions, that the species though distinct are more or less related, according to the greater or less possibility of the transportal in past and present times of species from one to the other region; although we can hardly admit that all the species in such cases have been transported from the first to the second region, and since have become extinct in the first: we see this law in the presence of the fox on the Falkland Islands; in the European character of some of the plants of Tierra del Fuego; in the Indo-Asiatic character of the plants of the Pacific; and in the circumstance of those genera which range widest having many species with wide ranges; and those genera with restricted ranges having species with restricted ranges. Finally, we find in each of the main divisions of the land, and probably of the sea, that the existing organisms are related to those lately extinct.
Looking further backwards we see that the past geographical distribution of organic beings was different from the present; and indeed, considering that geology shows that all our land was once under water, and that where water now extends land is forming, the reverse could hardly have been possible.
Now these several facts, though evidently all more or less connected together, must by the creationist (though the geologist may explain some of the anomalies) be considered as so many ultimate facts. He can only say, that it so pleased the Creator that the organic beings of the plains, deserts, mountains, tropical and temperature forests, of S. America, should all have some affinity together; that the inhabitants of the Galapagos Archipelago should be related to those of Chile; and that some of the species on the similarly constituted islands of this archipelago, though most closely related, should be distinct; that all its inhabitants should be totally unlike those of the similarly volcanic and arid Cape de Verde and Canary Islands; that the plants on the summit of Teneriffe should be eminently peculiar; that the diversified island of New Zealand should have not many plants, and not one, or only one, mammifer; that the mammifers of S. America, Australia and Europe should be clearly related to their ancient and exterminated prototypes; and so on with other facts. But it is absolutely opposed to every analogy, drawn from the laws imposed by the Creator on inorganic matter, that facts, when connected, should be considered as ultimate and not the direct consequences of more general laws.
An attempt to explain the foregoing laws of geographical distribution, on the theory of allied species having a common descent.
First let us recall the circumstances most favourable for variation under domestication, as given in the first chapter--viz. 1st, a change, or repeated changes, in the conditions to which the organism has been exposed, continued through several seminal (i.e. not by buds or divisions) generations: 2nd, steady selection of the slight varieties thus generated with a fixed end in view: 3rd, isolation as perfect as possible of such selected varieties; that is, the preventing their crossing with other forms; this latter condition applies to all terrestrial animals, to most if not all plants and perhaps even to most (or all) aquatic organisms. It will be convenient here to show the advantage of isolation in the formation of a new breed, by comparing the progress of two persons (to neither of whom let time be of any consequence) endeavouring to select and form some very peculiar new breed. Let one of these persons work on the vast herds of cattle in the plains of La Plata, and the other on a small stock of 20 or 30 animals in an island. The latter might have to wait centuries (by the hypothesis of no importance) before he obtained a "sport" approaching to what he wanted; but when he did and saved the greater number of its offspring and their offspring again, he might hope that his whole little stock would be in some degree affected, so that by continued selection he might gain his end. But on the Pampas, though the man might get his first approach to his desired form sooner, how hopeless would it be to attempt, by saving its offspring amongst so many of the common kind, to affect the whole herd: the effect of this one peculiar "sport" would be quite lost before he could obtain a second original sport of the same kind. If, however, he could separate a small number of cattle, including the offspring of the desirable "sport," he might hope, like the man on the island, to effect his end. If there be organic beings of which two individuals never unite, then simple selection whether on a continent or island would be equally serviceable to make a new and desirable breed; and this new breed might be made in surprisingly few years from the great and geometrical powers of propagation to beat out the old breed; as has happened (notwithstanding crossing) where good breeds of dogs and pigs have been introduced into a limited country,--for instance, into the islands of the Pacific.
 This instance occurs in the Essay of 1842, p. 32, but not in the Origin; though the importance of isolation is discussed (Origin, Ed. i. p. 104, vi. p. 127).
 The meaning of the words within parenthesis is obscure.
 It is unusual to find the author speaking of the selection of sports rather than small variations.
Let us now take the simplest natural case of an islet upheaved by the volcanic or subterranean forces in a deep sea, at such a distance from other land that only a few organic beings at rare intervals were transported to it, whether borne by the sea (like the seeds of plants to coral-reefs), or by hurricanes, or by floods, or on rafts, or in roots of large trees, or the germs of one plant or animal attached to or in the stomach of some other animal, or by the intervention (in most cases the most probable means) of other islands since sunk or destroyed. It may be remarked that when one part of the earth's crust is raised it is probably the general rule that another part sinks. Let this island go on slowly, century after century, rising foot by foot; and in the course of time we shall have instead [[of]] a small mass of rock, lowland and highland, moist woods and dry sandy spots, various soils, marshes, streams and pools: under water on the sea shore, instead of a rocky steeply shelving coast, we shall have in some parts bays with mud, sandy beaches and rocky shoals. The formation of the island by itself must often slightly affect the surrounding climate. It is impossible that the first few transported organisms could be perfectly adapted to all these stations; and it will be a chance if those successively transported will be so adapted. The greater number would probably come from the lowlands of the nearest country; and not even all these would be perfectly adapted to the new islet whilst it continued low and exposed to coast influences. Moreover, as it is certain that all organisms are nearly as much adapted in their structure to the other inhabitants of their country as they are to its physical conditions, so the mere fact that a few beings (and these taken in great degree by chance) were in the first case transported to the islet, would in itself greatly modify their conditions. As the island continued rising we might also expect an occasional new visitant; and I repeat that even one new being must often affect beyond our calculation by occupying the room and taking part of the subsistence of another (and this again from another and so on), several or many other organisms. Now as the first transported and any occasional successive visitants spread or tended to spread over the growing island, they would undoubtedly be exposed through several generations to new and varying conditions: it might also easily happen that some of the species on an average might obtain an increase of food, or food of a more nourishing quality. According then to every analogy with what we have seen takes place in every country, with nearly every organic being under domestication, we might expect that some of the inhabitants of the island would "sport," or have their organization rendered in some degree plastic. As the number of the inhabitants are supposed to be few and as all these cannot be so well adapted to their new and varying conditions as they were in their native country and habitat, we cannot believe that every place or office in the economy of the island would be as well filled as on a continent where the number of aboriginal species is far greater and where they consequently hold a more strictly limited place. We might therefore expect on our island that although very many slight variations were of no use to the plastic individuals, yet that occasionally in the course of a century an individual might be born of which the structure or constitution in some slight degree would allow it better to fill up some office in the insular economy and to struggle against other species. If such were the case the individual and its offspring would have a better chance of surviving and of beating out its parent form; and if (as is probable) it and its offspring crossed with the unvaried parent form, yet the number of the individuals being not very great, there would be a chance of the new and more serviceable form being nevertheless in some slight degree preserved. The struggle for existence would go on annually selecting such individuals until a new race or species was formed. Either few or all the first visitants to the island might become modified, according as the physical conditions of the island and those resulting from the kind and number of other transported species were different from those of the parent country--according to the difficulties offered to fresh immigration--and according to the length of time since the first inhabitants were introduced. It is obvious that whatever was the country, generally the nearest from which the first tenants were transported, they would show an affinity, even if all had become modified, to the natives of that country and even if the inhabitants of the same source (?) had been modified. On this view we can at once understand the cause and meaning of the affinity of the fauna and flora of the Galapagos Islands with that of the coast of S. America; and consequently why the inhabitants of these islands show not the smallest affinity with those inhabiting other volcanic islands, with a very similar climate and soil, near the coast of Africa.
 This brief discussion is represented in the Origin, Ed. i. by a much fuller one (pp. 356, 383, vi. pp. 504, 535). See, however, the section in the present Essay, p. 168.
 On the formation of new stations, see Origin, Ed. i. p. 292, vi. p. 429.
 Origin, Ed. i. pp. 390, 400, vi. pp. 543, 554.
 In the MS. some of the species ... nourishing quality is doubtfully erased. It seems clear that he doubted whether such a problematical supply of food would be likely to cause variation.
 At this time the author clearly put more faith in the importance of sport-like variation than in later years.
 Origin, Ed. i. p. 398, vi. p. 553.
To return once again to our island, if by the continued action of the subterranean forces other neighbouring islands were formed, these would generally be stocked by the inhabitants of the first island, or by a few immigrants from the neighbouring mainland; but if considerable obstacles were interposed to any communication between the terrestrial productions of these islands, and their conditions were different (perhaps only by the number of different species on each island), a form transported from one island to another might become altered in the same manner as one from the continent; and we should have several of the islands tenanted by representative races or species, as is so wonderfully the case with the different islands of the Galapagos Archipelago. As the islands become mountainous, if mountain-species were not introduced, as could rarely happen, a greater amount of variation and selection would be requisite to adapt the species, which originally came from the lowlands of the nearest continent, to the mountain-summits than to the lower districts of our islands. For the lowland species from the continent would have first to struggle against other species and other conditions on the coast-land of the island, and so probably become modified by the selection of its best fitted varieties, then to undergo the same process when the land had attained a moderate elevation; and then lastly when it had become Alpine. Hence we can understand why the faunas of insular mountain-summits are, as in the case of Teneriffe, eminently peculiar. Putting on one side the case of a widely extended flora being driven up the mountain-summits, during a change of climate from cold to temperate, we can see why in other cases the floras of mountain-summits (or as I have called them islands in a sea of land) should be tenanted by peculiar species, but related to those of the surrounding lowlands, as are the inhabitants of a real island in the sea to those of the nearest continent.
 See Origin, Ed. i. p. 403, vi. p. 558, where the author speaks of Alpine humming birds, rodents, plants, &c. in S. America, all of strictly American forms. In the MS. the author has added between the lines "As world has been getting hotter, there has been radiation from high-lands,--old view?--curious; I presume Diluvian in origin."
Let us now consider the effect of a change of climate or of other conditions on the inhabitants of a continent and of an isolated island without any great change of level. On a continent the chief effects would be changes in the numerical proportion of the individuals of the different species; for whether the climate became warmer or colder, drier or damper, more uniform or extreme, some species are at present adapted to its diversified districts; if for instance it became cooler, species would migrate from its more temperate parts and from its higher land; if damper, from its damper regions, &c. On a small and isolated island, however, with few species, and these not adapted to much diversified conditions, such changes instead of merely increasing the number of certain species already adapted to such conditions, and decreasing the number of other species, would be apt to affect the constitutions of some of the insular species: thus if the island became damper it might well happen that there were no species living in any part of it adapted to the consequences resulting from more moisture. In this case therefore, and still more (as we have seen) during the production of new stations from the elevation of the land, an island would be a far more fertile source, as far as we can judge, of new specific forms than a continent. The new forms thus generated on an island, we might expect, would occasionally be transported by accident, or through long-continued geographical changes be enabled to emigrate and thus become slowly diffused.
But if we look to the origin of a continent; almost every geologist will admit that in most cases it will have first existed as separate islands which gradually increased in size; and therefore all that which has been said concerning the probable changes of the forms tenanting a small archipelago is applicable to a continent in its early state. Furthermore, a geologist who reflects on the geological history of Europe (the only region well known) will admit that it has been many times depressed, raised and left stationary. During the sinking of a continent and the probable generally accompanying changes of climate the effect would be little, except on the numerical proportions and in the extinction (from the lessening of rivers, the drying of marshes and the conversion of high-lands into low &c.) of some or of many of the species. As soon however as the continent became divided into many isolated portions or islands, preventing free immigration from one part to another, the effect of climatic and other changes on the species would be greater. But let the now broken continent, forming isolated islands, begin to rise and new stations thus to be formed, exactly as in the first case of the upheaved volcanic islet, and we shall have equally favourable conditions for the modification of old forms, that is the formation of new races or species. Let the islands become reunited into a continent; and then the new and old forms would all spread, as far as barriers, the means of transportal, and the preoccupation of the land by other species, would permit. Some of the new species or races would probably become extinct, and some perhaps would cross and blend together. We should thus have a multitude of forms, adapted to all kinds of slightly different stations, and to diverse groups of either antagonist or food-serving species. The oftener these oscillations of level had taken place (and therefore generally the older the land) the greater the number of species [[which]] would tend to be formed. The inhabitants of a continent being thus derived in the first stage from the same original parents, and subsequently from the inhabitants of one wide area, since often broken up and reunited, all would be obviously related together and the inhabitants of the most dissimilar stations on the same continent would be more closely allied than the inhabitants of two very similar stations on two of the main divisions of the world.
 See the comparison between the Malay Archipelago and the probable former state of Europe, Origin, Ed. i. p. 299, vi. p. 438, also Origin, Ed. i. p. 292, vi. p. 429.
 Origin, Ed. i. p. 349, vi. p. 496. The arrangement of the argument in the present Essay leads to repetition of statements made in the earlier part of the book: in the Origin this is avoided.
I need hardly point out that we now can obviously see why the number of species in two districts, independently of the number of stations in such districts, should be in some cases as widely different as in New Zealand and the Cape of Good Hope. We can see, knowing the difficulty in the transport of terrestrial mammals, why islands far from mainlands do not possess them; we see the general reason, namely accidental transport (though not the precise reason), why certain islands should, and others should not, possess members of the class of reptiles. We can see why an ancient channel of communication between two distant points, as the Cordillera probably was between southern Chile and the United States during the former cold periods; and icebergs between the Falkland Islands and Tierra del Fuego; and gales, at a former or present time, between the Asiatic shores of the Pacific and eastern islands in this ocean; is connected with (or we may now say causes) an affinity between the species, though distinct, in two such districts. We can see how the better chance of diffusion, from several of the species of any genus having wide ranges in their own countries, explains the presence of other species of the same genus in other countries; and on the other hand, of species of restricted powers of ranging, forming genera with restricted ranges.
 Origin, Ed. i. p. 389, vi. p. 542.
 Origin, Ed. i. p. 393, vi. p. 547.
 Origin, Ed. i. pp. 350, 404, vi. pp. 498, 559.
As every one would be surprised if two exactly similar but peculiar varieties of any species were raised by man by long continued selection, in two different countries, or at two very different periods, so we ought not to expect that an exactly similar form would be produced from the modification of an old one in two distinct countries or at two distinct periods. For in such places and times they would probably be exposed to somewhat different climates and almost certainly to different associates. Hence we can see why each species appears to have been produced singly, in space and in time. I need hardly remark that, according to this theory of descent, there is no necessity of modification in a species, when it reaches a new and isolated country. If it be able to survive and if slight variations better adapted to the new conditions are not selected, it might retain (as far as we can see) its old form for an indefinite time. As we see that some sub-varieties produced under domestication are more variable than others, so in nature, perhaps, some species and genera are more variable than others. The same precise form, however, would probably be seldom preserved through successive geological periods, or in widely and differently conditioned countries.
 Origin, Ed. i. p. 352, vi. p. 500.
 Origin, Ed. i. p. 313, vi. p. 454.
Finally, during the long periods of time and probably of oscillations of level, necessary for the formation of a continent, we may conclude (as above explained) that many forms would become extinct. These extinct forms, and those surviving (whether or not modified and changed in structure), will all be related in each continent in the same manner and degree, as are the inhabitants of any two different sub-regions in that same continent. I do not mean to say that, for instance, the present Marsupials of Australia or Edentata and rodents of S. America have descended from any one of the few fossils of the same orders which have been discovered in these countries. It is possible that, in a very few instances, this may be the case; but generally they must be considered as merely codescendants of common stocks. I believe in this, from the improbability, considering the vast number of species, which (as explained in the last chapter) must by our theory have existed, that the comparatively few fossils which have been found should chance to be the immediate and linear progenitors of those now existing. Recent as the yet discovered fossil mammifers of S. America are, who will pretend to say that very many intermediate forms may not have existed? Moreover, we shall see in the ensuing chapter that the very existence of genera and species can be explained only by a few species of each epoch leaving modified successors or new species to a future period; and the more distant that future period, the fewer will be the linear heirs of the former epoch. As by our theory, all mammifers must have descended from the same parent stock, so is it necessary that each land now possessing terrestrial mammifers shall at some time have been so far united to other land as to permit the passage of mammifers; and it accords with this necessity, that in looking far back into the earth's history we find, first changes in the geographical distribution, and secondly a period when the mammiferous forms most distinctive of two of the present main divisions of the world were living together.
 Origin, Ed. i. p. 341, vi. p. 487.
 Origin, Ed. i. p. 396, vi. p. 549.
 Origin, Ed. i. p. 340, vi. p. 486.
I think then I am justified in asserting that most of the above enumerated and often trivial points in the geographical distribution of past and present organisms (which points must be viewed by the creationists as so many ultimate facts) follow as a simple consequence of specific forms being mutable and of their being adapted by natural selection to diverse ends, conjoined with their powers of dispersal, and the geologico-geographical changes now in slow progress and which undoubtedly have taken place. This large class of facts being thus explained, far more than counterbalances many separate difficulties and apparent objections in convincing my mind of the truth of this theory of common descent.
Improbability of finding fossil forms intermediate between existing species.
There is one observation of considerable importance that may be here introduced, with regard to the improbability of the chief transitional forms between any two species being found fossil. With respect to the finer shades of transition, I have before remarked that no one has any cause to expect to trace them in a fossil state, without he be bold enough to imagine that geologists at a future epoch will be able to trace from fossil bones the gradations between the Short-Horns, Herefordshire, and Alderney breeds of cattle. I have attempted to show that rising islands, in process of formation, must be the best nurseries of new specific forms, and these points are the least favourable for the embedment of fossils: I appeal, as evidence, to the state of the numerous scattered islands in the several great oceans: how rarely do any sedimentary deposits occur on them; and when present they are mere narrow fringes of no great antiquity, which the sea is generally wearing away and destroying. The cause of this lies in isolated islands being generally volcanic and rising points; and the effects of subterranean elevation is to bring up the surrounding newly-deposited strata within the destroying action of the coast-waves: the strata, deposited at greater distances, and therefore in the depths of the ocean, will be almost barren of organic remains. These remarks may be generalised:--periods of subsidence will always be most favourable to an accumulation of great thicknesses of strata, and consequently to their long preservation; for without one formation be protected by successive strata, it will seldom be preserved to a distant age, owing to the enormous amount of denudation, which seems to be a general contingent of time. I may refer, as evidence of this remark, to the vast amount of subsidence evident in the great pile of the European formations, from the Silurian epoch to the end of the Secondary, and perhaps to even a later period. Periods of elevation on the other hand cannot be favourable to the accumulation of strata and their preservation to distant ages, from the circumstance just alluded to, viz. of elevation tending to bring to the surface the circum-littoral strata (always abounding most in fossils) and destroying them. The bottom of tracts of deep water (little favourable, however, to life) must be excepted from this unfavourable influence of elevation. In the quite open ocean, probably no sediment is accumulating, or at a rate so slow as not to preserve fossil remains, which will always be subject to disintegration. Caverns, no doubt, will be equally likely to preserve terrestrial fossils in periods of elevation and of subsidence; but whether it be owing to the enormous amount of denudation, which all land seems to have undergone, no cavern with fossil bones has been found belonging to the Secondary period.
 Origin, Ed. i. p. 299, vi. p. 437.
 "Nature may almost be said to have guarded against the frequent discovery of her transitional or linking forms," Origin, Ed. i. p. 292. A similar but not identical passage occurs in Origin, Ed. vi. p. 428.
 Origin, Ed. i. p. 291, vi. p. 426.
 Origin, Ed. i. p. 288, vi. p. 422.
 Origin, Ed. i. p. 289, vi. p. 423.
Hence many more remains will be preserved to a distant age, in any region of the world, during periods of its subsidence, than of its elevation.
 Origin, Ed. i. p. 300, vi. p. 439.
But during the subsidence of a tract of land, its inhabitants (as before shown) will from the decrease of space and of the diversity of its stations, and from the land being fully preoccupied by species fitted to diversified means of subsistence, be little liable to modification from selection, although many may, or rather must, become extinct. With respect to its circum-marine inhabitants, although during a change from a continent to a great archipelago, the number of stations fitted for marine beings will be increased, their means of diffusion (an important check to change of form) will be greatly improved; for a continent stretching north and south, or a quite open space of ocean, seems to be to them the only barrier. On the other hand, during the elevation of a small archipelago and its conversion into a continent, we have, whilst the number of stations are increasing, both for aquatic and terrestrial productions, and whilst these stations are not fully preoccupied by perfectly adapted species, the most favourable conditions for the selection of new specific forms; but few of them in their early transitional states will be preserved to a distant epoch. We must wait during an enormous lapse of time, until long-continued subsidence shall have taken the place in this quarter of the world of the elevatory process, for the best conditions of the embedment and the preservation of its inhabitants. Generally the great mass of the strata in every country, from having been chiefly accumulated during subsidence, will be the tomb, not of transitional forms, but of those either becoming extinct or remaining unmodified.
The state of our knowledge, and the slowness of the changes of level, do not permit us to test the truth of these remarks, by observing whether there are more transitional or "fine" (as naturalists would term them) species, on a rising and enlarging tract of land, than on an area of subsidence. Nor do I know whether there are more "fine" species on isolated volcanic islands in process of formation, than on a continent; but I may remark, that at the Galapagos Archipelago the number of forms, which according to some naturalists are true species, and according to others are mere races, is considerable: this particularly applies to the different species or races of the same genera inhabiting the different islands of this archipelago. Furthermore it may be added (as bearing on the great facts discussed in this chapter) that when naturalists confine their attention to any one country, they have comparatively little difficulty in determining what forms to call species and what to call varieties; that is, those which can or cannot be traced or shown to be probably descendants of some other form: but the difficulty increases, as species are brought from many stations, countries and islands. It was this increasing (but I believe in few cases insuperable) difficulty which seems chiefly to have urged Lamarck to the conclusion that species are mutable.