The Foundations of the Origin of Species

Nor can it be admitted as probable that any one Secondary formation contains a fair record even of those organisms which are most easily preserved, namely hard marine bodies. In how many cases have we not certain evidence that between the deposition of apparently closely consecutive beds, the lower one existed for an unknown time as land, covered with trees. Some of the Secondary formations which contain most marine remains appear to have been formed in a wide and not deep sea, and therefore only those marine animals which live in such situations would be preserved[320 - «Note in original.» Neither highest or lowest fish (i. e. Myxina «?» or Lepidosiren) could be preserved in intelligible condition in fossils.]. In all cases, on indented rocky coasts, or any other coast, where sediment is not accumulating, although often highly favourable to marine animals, none can be embedded: where pure sand and pebbles are accumulating few or none will be preserved. I may here instance the great western line of the S. American coast[321 - Origin, Ed. i. p. 290, vi. p. 425.], tenanted by many peculiar animals, of which none probably will be preserved to a distant epoch. From these causes, and especially from such deposits as are formed along a line of coast, steep above and below water, being necessarily of little width, and therefore more likely to be subsequently denuded and worn away, we can see why it is improbable that our Secondary deposits contain a fair record of the Marine Fauna of any one period. The East Indian Archipelago offers an area, as large as most of our Secondary deposits, in which there are wide and shallow seas, teeming with marine animals, and in which sediment is accumulating; now supposing that all the hard marine animals, or rather those having hard parts to preserve, were preserved to a future age, excepting those which lived on rocky shores where no sediment or only sand and gravel were accumulating, and excepting those embedded along the steeper coasts, where only a narrow fringe of sediment was accumulating, supposing all this, how poor a notion would a person at a future age have of the Marine Fauna of the present day. Lyell[322 - See Origin, Ed. i. p. 310, vi. p. 452 for Lyell's metaphor. I am indebted to Prof. Judd for pointing out that Darwin’s version of the metaphor is founded on the first edition of Lyell’s Principles, vol. I. and vol. III.; see the Essay of 1842, p. 27.] has compared the geological series to a work of which only the few latter but not consecutive chapters have been preserved; and out of which, it may be added, very many leaves have been torn, the remaining ones only illustrating a scanty portion of the Fauna of each period. On this view, the records of anteceding ages confirm my theory; on any other they destroy it.

Finally, if we narrow the question into, why do we not find in some instances every intermediate form between any two species? the answer may well be that the average duration of each specific form (as we have good reason to believe) is immense in years, and that the transition could, according to my theory, be effected only by numberless small gradations; and therefore that we should require for this end a most perfect record, which the foregoing reasoning teaches us not to expect. It might be thought that in a vertical section of great thickness in the same formation some of the species ought to be found to vary in the upper and lower parts[323 - See More Letters, vol. I. pp. 344-7, for Darwin’s interest in the celebrated observations of Hilgendorf and Hyatt.], but it may be doubted whether any formation has gone on accumulating without any break for a period as long as the duration of a species; and if it had done so, we should require a series of specimens from every part. How rare must be the chance of sediment accumulating for some 20 or 30 thousand years on the same spot[324 - This corresponds partly to Origin, Ed. i. p. 294, vi. p. 431.], with the bottom subsiding, so that a proper depth might be preserved for any one species to continue living: what an amount of subsidence would be thus required, and this subsidence must not destroy the source whence the sediment continued to be derived. In the case of terrestrial animals, what chance is there when the present time is become a pleistocene formation (at an earlier period than this, sufficient elevation to expose marine beds could not be expected), what chance is there that future geologists will make out the innumerable transitional sub-varieties, through which the short-horned and long-horned cattle (so different in shape of body) have been derived from the same parent stock[325 - Origin, Ed. i. p. 299, vi. p. 437.]? Yet this transition has been effected in the same country, and in a far shorter time, than would be probable in a wild state, both contingencies highly favourable for the future hypothetical geologists being enabled to trace the variation.

CHAPTER V

GRADUAL APPEARANCE AND DISAPPEARANCE OF SPECIES[326 - This chapter corresponds to ch. X of Origin, Ed. i., vi. ch. XI, “On the geological succession of organic beings.”]

In the Tertiary system, in the last uplifted beds, we find all the species recent and living in the immediate vicinity; in rather older beds we find only recent species, but some not living in the immediate vicinity[327 - Origin, Ed. i. p. 312, vi. p. 453.]; we then find beds with two or three or a few more extinct or very rare species; then considerably more extinct species, but with gaps in the regular increase; and finally we have beds with only two or three or not one living species. Most geologists believe that the gaps in the percentage, that is the sudden increments, in the number of the extinct species in the stages of the Tertiary system are due to the imperfection of the geological record. Hence we are led to believe that the species in the Tertiary system have been gradually introduced; and from analogy to carry on the same view to the Secondary formations. In these latter, however, entire groups of species generally come in abruptly; but this would naturally result, if, as argued in the foregoing chapter, these Secondary deposits are separated by wide epochs. Moreover it is important to observe that, with our increase of knowledge, the gaps between the older formations become fewer and smaller; geologists of a few years standing remember how beautifully has the Devonian system[328 - In the margin the author has written “Lonsdale.” This refers to W. Lonsdale’s paper “Notes on the age of the Limestone of South Devonshire,” Geolog. Soc. Trans., Series 2, vol. V. 1840, p. 721. According to Mr H. B. Woodward (History of the Geological Society of London, 1907, p. 107) “Lonsdale’s ‘important and original suggestion of the existence of an intermediary type of Palæozoic fossils, since called Devonian,’ led to a change which was then ‘the greatest ever made at one time in the classification of our English formations’.” Mr Woodward’s quotations are from Murchison and Buckland.] come in between the Carboniferous and Silurian formations. I need hardly observe that the slow and gradual appearance of new forms follows from our theory, for to form a new species, an old one must not only be plastic in its organization, becoming so probably from changes in the conditions of its existence, but a place in the natural economy of the district must [be made,] come to exist, for the selection of some new modification of its structure, better fitted to the surrounding conditions than are the other individuals of the same or other species[329 - «Note in original.» Better begin with this. If species really, after catastrophes, created in showers over world, my theory false. «In the above passage the author is obviously close to his theory of divergence.»].

In the Tertiary system the same facts, which make us admit as probable that new species have slowly appeared, lead to the admission that old ones have slowly disappeared, not several together, but one after another; and by analogy one is induced to extend this belief to the Secondary and Palæozoic epochs. In some cases, as the subsidence of a flat country, or the breaking or the joining of an isthmus, and the sudden inroad of many new and destructive species, extinction might be locally sudden. The view entertained by many geologists, that each fauna of each Secondary epoch has been suddenly destroyed over the whole world, so that no succession could be left for the production of new forms, is subversive of my theory, but I see no grounds whatever to admit such a view. On the contrary, the law, which has been made out, with reference to distinct epochs, by independent observers, namely, that the wider the geographical range of a species the longer is its duration in time, seems entirely opposed to any universal extermination[330 - Opposite to this passage the author has written “d’Archiac, Forbes, Lyell.”]. The fact of species of mammiferous animals and fish being renewed at a quicker rate than mollusca, though both aquatic; and of these the terrestrial genera being renewed quicker than the marine; and the marine mollusca being again renewed quicker than the Infusorial animalcula, all seem to show that the extinction and renewal of species does not depend on general catastrophes, but on the particular relations of the several classes to the conditions to which they are exposed[331 - This passage, for which the author gives as authorities the names of Lyell, Forbes and Ehrenberg, corresponds in part to the discussion beginning on p. 313 of Origin, Ed. i., vi. p. 454.].

Some authors seem to consider the fact of a few species having survived[332 - The author gives Falconer as his authority: see Origin, Ed. i. p. 313, vi. p. 454.] amidst a number of extinct forms (as is the case with a tortoise and a crocodile out of the vast number of extinct sub-Himalayan fossils) as strongly opposed to the view of species being mutable. No doubt this would be the case, if it were presupposed with Lamarck that there was some inherent tendency to change and development in all species, for which supposition I see no evidence. As we see some species at present adapted to a wide range of conditions, so we may suppose that such species would survive unchanged and unexterminated for a long time; time generally being from geological causes a correlative of changing conditions. How at present one species becomes adapted to a wide range, and another species to a restricted range of conditions, is of difficult explanation.

Extinction of species

The extinction of the larger quadrupeds, of which we imagine we better know the conditions of existence, has been thought little less wonderful than the appearance of new species; and has, I think, chiefly led to the belief of universal catastrophes. When considering the wonderful disappearance within a late period, whilst recent shells were living, of the numerous great and small mammifers of S. America, one is strongly induced to join with the catastrophists. I believe, however, that very erroneous views are held on this subject. As far as is historically known, the disappearance of species from any one country has been slow – the species becoming rarer and rarer, locally extinct, and finally lost[333 - This corresponds approximately to Origin, Ed. i. p. 317, vi. p. 458.]. It may be objected that this has been effected by man’s direct agency, or by his indirect agency in altering the state of the country; in this latter case, however, it would be difficult to draw any just distinction between his agency and natural agencies. But we now know in the later Tertiary deposits, that shells become rarer and rarer in the successive beds, and finally disappear: it has happened, also, that shells common in a fossil state, and thought to have been extinct, have been found to be still living species, but very rare ones[334 - The case of Trigonia, a great Secondary genus of shells surviving in a single species in the Australian seas, is given as an example in the Origin, Ed. i. p. 321, vi. p. 463.]. If the rule is that organisms become extinct by becoming rarer and rarer, we ought not to view their extinction, even in the case of the larger quadrupeds, as anything wonderful and out of the common course of events. For no naturalist thinks it wonderful that one species of a genus should be rare and another abundant, notwithstanding he be quite incapable of explaining the causes of the comparative rareness[335 - This point, on which the author laid much stress, is discussed in the Origin, Ed. i. p. 319, vi. p. 461.]. Why is one species of willow-wren or hawk or woodpecker common in England, and another extremely rare: why at the Cape of Good Hope is one species of rhinoceros or antelope far more abundant than other species? Why again is the same species much more abundant in one district of a country than in another district? No doubt there are in each case good causes: but they are unknown and unperceived by us. May we not then safely infer that as certain causes are acting unperceived around us, and are making one species to be common and another exceedingly rare, that they might equally well cause the final extinction of some species without being perceived by us? We should always bear in mind that there is a recurrent struggle for life in every organism, and that in every country a destroying agency is always counteracting the geometrical tendency to increase in every species; and yet without our being able to tell with certainty at what period of life, or at what period of the year, the destruction falls the heaviest. Ought we then to expect to trace the steps by which this destroying power, always at work and scarcely perceived by us, becomes increased, and yet if it continues to increase ever so slowly (without the fertility of the species in question be likewise increased) the average number of the individuals of that species must decrease, and become finally lost. I may give a single instance of a check causing local extermination which might long have escaped discovery[336 - Origin, Ed. i. p. 72, vi. p. 89.]; the horse, though swarming in a wild state in La Plata, and likewise under apparently the most unfavourable conditions in the scorched and alternately flooded plains of Caraccas, will not in a wild state extend beyond a certain degree of latitude into the intermediate country of Paraguay; this is owing to a certain fly depositing its eggs on the navels of the foals: as, however, man with a little care can rear horses in a tame state abundantly in Paraguay, the problem of its extinction is probably complicated by the greater exposure of the wild horse to occasional famine from the droughts, to the attacks of the jaguar and other such evils. In the Falkland Islands the check to the increase of the wild horse is said to be loss of the sucking foals[337 - This case does not occur in the Origin, Ed.], from the stallions compelling the mares to travel across bogs and rocks in search of food: if the pasture on these islands decreased a little, the horse, perhaps, would cease to exist in a wild state, not from the absolute want of food, but from the impatience of the stallions urging the mares to travel whilst the foals were too young.

From our more intimate acquaintance with domestic animals, we cannot conceive their extinction without some glaring agency; we forget that they would undoubtedly in a state of nature (where other animals are ready to fill up their place) be acted on in some part of their lives by a destroying agency, keeping their numbers on an average constant. If the common ox was known only as a wild S. African species, we should feel no surprise at hearing that it was a very rare species; and this rarity would be a stage towards its extinction. Even in man, so infinitely better known than any other inhabitant of this world, how impossible it has been found, without statistical calculations, to judge of the proportions of births and deaths, of the duration of life, and of the increase and decrease of population; and still less of the causes of such changes: and yet, as has so often been repeated, decrease in numbers or rarity seems to be the high-road to extinction. To marvel at the extermination of a species appears to me to be the same thing as to know that illness is the road to death, – to look at illness as an ordinary event, nevertheless to conclude, when the sick man dies, that his death has been caused by some unknown and violent agency[338 - An almost identical sentence occurs in the Origin, Ed. i. p. 320, vi. p. 462.].

In a future part of this work we shall show that, as a general rule, groups of allied species[339 - Origin, Ed. i. p. 316, vi. p. 457.] gradually appear and disappear, one after the other, on the face of the earth, like the individuals of the same species: and we shall then endeavour to show the probable cause of this remarkable fact.

CHAPTER VI

ON THE GEOGRAPHICAL DISTRIBUTION OF ORGANIC BEINGS IN PAST AND PRESENT TIMES

For convenience sake I shall divide this chapter into three sections[340 - 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]. 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.

Section First

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[341 - This division of the land into regions does not occur in the Origin, Ed. i.] 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[342 - Origin, Ed. i. p. 346, vi. p. 493.]; 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[343 - 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».]. 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.

Furthermore it may be observed that all the organisms inhabiting any country are not perfectly adapted to it[344 - This partly corresponds to Origin, Ed. i. p. 337, vi. p. 483.]; 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.

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[345 - On the general importance of barriers, see Origin, Ed. i. p. 347, vi. p. 494.] 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[346 - Origin, Ed. i. p. 348, vi. p. 495.], 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.

Relation of range in genera and species

It is generally[347 - «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 (#Page_146).»] 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[348 - Origin, Ed. i. p. 404, vi. p. 559.]. 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.

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[349 - Origin, Ed. i. p. 349, vi. p. 496.], 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.

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[350 - The case of the ostrich (Rhea) occurs in the Origin, Ed. i. p. 349, vi. p. 496.] 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[351 - «Note in original.» There is a hare in S. America, – so bad example.] 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[352 - See Origin, Ed. i. p. 349, vi. p. 497.] 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.

Insular Faunas

If we now look to the character of the inhabitants of small islands[353 - For the general problem of Oceanic Islands, see Origin, Ed. i. p. 388, vi. p. 541.], we shall find that those situated close to other land have a similar fauna with that land[354 - 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.”], whilst those at a considerable distance from other land often possess an almost entirely peculiar fauna. The Galapagos Archipelago[355 - Origin, Ed. i. p. 390, vi. p. 543.] 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[356 - See Origin, Ed. i. p. 397, vi. p. 552.]. 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[357 - 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.] 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.

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[358 - 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.”] 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.

Islands standing quite isolated within the intra-tropical oceans have generally very peculiar floras, related, though feebly (as in the case of St Helena[359 - “The affinities of the St Helena flora are strongly South African.” Hooker’s Lecture on Insular Floras in the Gardeners’ Chronicle, Jan. 1867.] 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[360 - It is impossible to make out the precise form which the author intended to give to this sentence, but the meaning is clear.]. 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[361 - This is no doubt true, the flora of the Sandwich group however has marked American affinities.]. 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.

Alpine Floras

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[362 - 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.] 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[363 - The apparent breakdown of the doctrine of barriers is slightly touched on in the Origin, Ed. i. p. 365, vi. p. 515.]. 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[364 - 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.”]. 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[365 - See Hooker’s Lecture on Insular Floras in the Gardeners’ Chronicle, Jan. 1867.]; this is the case on Teneriffe, and in a lesser degree even on some of the Mediterranean islands.

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[366 - 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.]? 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[367 - See Origin, Ed. i. p. 367, vi. p. 517.] 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[368 - «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.»] and often driven high up on distant beaches, might have been the means of widely distributing the seeds of the same species?

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[369 - A note by the author gives “many authors” apparently as authority for this statement.] 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.

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[370 - Opposite to this passage, in the margin, the author has written: – “too hypothetical.”] (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[371 - The Cordillera is described as supplying a great line of invasion in the Origin, Ed. i. p. 378.] 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[372 - 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.] 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[373 - See Hooker’s Lecture on Insular Floras in the Gardeners’ Chronicle, Jan. 1867.] 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[374 - «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[375 - On centres of creation see Origin, Ed. i. p. 352, vi. p. 499.], 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[376 - 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.] 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[377 - 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.]), and of the possibility of species having survived for short periods at intermediate spots and afterwards becoming extinct there[378 - 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.]; 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.

But the strongest argument against double creations may be drawn from considering the case of mammifers[379 - “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.] 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[380 - «Note by the author.» Many authors. «See Origin, Ed. i. p. 394, vi. p. 547.»]. The cases of lesser difficulty may all receive a more or less simple explanation; I will give only one instance; the nutria[381 - 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.], 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[382 - In Origin, Ed. i. p. 394, vi. p. 548, bats are mentioned as an explicable exception to this statement.]. 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[383 - 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.], 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[384 - See Origin, Ed. i. p. 313, vi. p. 454.]. 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.

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[385 - The comparison between New Zealand and the Cape is given in the Origin, Ed. i. p. 389, vi. p. 542.], 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[386 - 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.] 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[387 - The whole argument is given – more briefly than here – in the Origin, Ed. i. p. 394, vi. p. 547.]. 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[388 - See Origin, Ed i. p. 393, vi. p. 547. The discussion is much fuller in the present Essay.]; 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.

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.

Section Second

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[389 - See Origin, Ed. i. p. 339, vi. p. 485.]. 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, hyæna, 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 «as to» classifying this district with S. America; and if fossil Edentata, Marsupials and monkeys were to be found in Tierra del Fuego, it would not make this district more truly S. American than it now is. So it is with Europe[390 - 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.], and so far as is known with Asia, for the lately past and present mammifers all belong to the Africo-Asiatic division of the world. In every case, I may add, the forms which a country has is of more importance in geographical arrangement than what it has not.

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[391 - “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, [hyæna,][392 - The word hyæna is erased. There appear to be no fossil Hyænidæ in S. America.] 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[393 - See note 1, p. 175 (#cn_389), also Origin, Ed. i. p. 340, vi. p. 486.] and Edentata, we behold an entire blending of those mammiferous forms which now eminently characterise Australia and S. America[394 - «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[395 - «Note by the author.» All this requires much verification.] 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[396 - This point seems to be less insisted on in the Origin.]. 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[397 - Origin, Ed. i. p. 356, vi. p. 504.]. 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.