RECORD: Darwin, C. R. 1856. Draft of Natural selection, chapter 11 'Geographical distribution'. CUL-DAR14.(A0-A6,B1-B55,C0-C5,D1-D47). (John van Wyhe ed., 2002-. The Complete Work of Charles Darwin Online, http://darwin-online.org.uk/)

REVISION HISTORY: First transcribed by R. C. Stauffer in Natural selection, F1583. Adapted with some corrections and omissions transcribed to correspond with the manuscript by John van Wyhe 2-3.2026. RN1

NOTE: See record in the Darwin Online manuscript catalogue, enter its Identifier here. Reproduced with permission of Cambridge University Press, the Syndics of Cambridge University Library and William Huxley Darwin.

CUL-DAR14.A0-A6 'Natural Selection', chap. 11, f. 0 and appendix, draft.
CUL-DAR14.B1-B55 'Natural Selection', chap. 11, ff. 1-55, draft.
CUL-DAR14.C0-C5 'Natural Selection', chap. 11, f. 0 [1-5] and appendix, fair copy.
CUL-DAR14.D1-D47 'Natural Selection', chap. 11, ff. 1-55, fair copy.

The transcription from R. C. Stauffer's book has been placed alongside the images with minor additions and corrections. There are a few discrepancies between the edited text and the original manuscript. For more detail about Darwin's chapter, see of course Stauffer, Natural selection.


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As I believe that all organic beings are produced by the ordinary laws of reproduction which includes, according to the theory under discussion, modification of specific forms, & as it is exceedingly improbable that the same species should ever have been generated in one place from one set of parents, & in another place, (especially if under different conditions) from another set of parents specifically dissimilar, the first & most obvious question is whether we can account on the ordinary notion of propagation for the existence of the same identical species in all quarters of the world.— This is the question, which has long agitated naturalists, namely whether the same species has been created once & therefore at a single point, or more than once at different points.

(After giving general reasons in favour of single production; consider the many & grave difficulties. The most prominent of them may be grouped into three following classes.— First, insular productions of the temperate & tropical latitudes.†— Here give reasons for doubting vast continental extensions of Forbes & Co:

† Perhaps allude to A. Decandolle on large-scaled Plants.

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: give condensed means of dispersal.

Secondly, range of Fresh Water Productions.—

Thirdly, as follows.—

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We will now consider some of the most striking cases of difficulty on geological grounds opposed to the theory that closely allied or representative species, are due to the modification of the same species.

Dr. Hooker has given a most curious list1 of representative species, found in New Zealand, Australia & S. America. With respect to Australia there is no greater difficulty (but great enough)

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(a) text.

This theory implies a communication of some kind between the areas occupied by the representative species at some former period, generally not very remote in a geological sense, as distinctly as does the theory of single centres of creation in regard to the same species when found at distant & separated points.

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This fact of half the genera being confined to the South seems to me remarkable considering that out of the 89 species belonging to 76 genera1 absolutely identical in New Zealand & S. America, only two species belong to genera confined to the South, namely Colobanthus subulatus & Rostkovia Magellanica: Goodenia repens need hardly be added as this is an Australian genus with one littoral wandering species.

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& 18 of very general distribution; & the representative species in these 25 genera (bearing in mind the glacial epoch) present nothing more remarkable than representative species in other parts of the world, to which in former times we may imagine the descendants of the same species to have travelled & subsequently to have become modified: but the other 25 genera are strictly confined to the south with all their species extratropical, — a few on the mountains within the Tropics being excepted.

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in single creations, that there has been at some time some channel of communication between the two areas in question. Dr. Hooker believes that   (a) 

(Of the 50 genera which afford the best instances of representative species in New Zealand & extratropical S. America, 7, as I am informed by Dr. Hooker, are northern genera,

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(a) text

there was formerly a communication by more or less continuous land from both Chile & Fuegia to New Zealand. I cannot persuade myself to admit (though far better judges see no difficulty in admitting) such great geographical changes within so recent a period; & I think that a slight modification of Dr. Hooker's view will remove some little of the difficulty.)

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Again it may be noticed in Dr. Hooker's list (which I am aware is not given as perfect) that of the southern genera, which have representatives in New Zealand & S. America, there are five which have none in Australia or Tasmania, & this is what might have been expected considering the greater distance of Australia, than of New Zealand from S. America; but Australia has four southern genera (viz; Eucryphia, Pernettya, Lebetanthus, & Lomatia) with representatives of S. American species, which genera do not occur in New Zealand or the Auckland islands. These facts, together perhaps with the genera Colobanthus, Ac[a]ena & Lagenophora having representative species, both on these two lands & in several of the circumpolar islands, seem to me to indicate some common centre of radiation.)

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representative species of the southern genera which in S. America grow in low land are confined in New Zealand & in Tasmania  to high land; (Flora of New Zealand Introduct. p. xxiv & note D) & as inferred by Dr Hooker (Id. p. xxiii) in regard to the identical species of the two countries similarly situated & associated with them, the climate of New Zealand must have grown cooler since their introduction.

(Now taking the northern hemisphere as our guide, I should look to the circumpolar regions as the centre of radiation for the representative & for some of the species still remaining identical in the above named several lands. If we look to a chart we see in the little explored regions between 62° & 80° several islands, & large tracts of land, with surroundings in one place 100 miles from the shore, & with indications2 of other rocks besides volcanic.

On these islands not one single land plant can now live, but bearing in mind that in the north the space between these same parallels is the home of the whole Arctic Flora, it seems to me a not very improbable supposition that before the glacial epoch came on, these islands might have been covered by a

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Here then I should infer that it was no ways improbable that these lands & islands may have recently been of greater extent & more continuous.

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tolerably uniform. From this source, I am inclined to suppose that some plants, either identical or allied, migrated before the glacial epoch by various accidental means, aided probably by more continuous land & by the several intermediate islands which we see still existing South of New Zealand; & that when the glacial epoch did come on, the seeds of other plants were brought in a N. Easterly course by icebergs from their common home, soon to be converted into an icy desert. The plants which arrived at this latter period, would, on the returning warmth have ascended the mountains of New Zealand & Tasmania, — most of the species especially those brought first, having subsequently undergone modification & now existing as representative species,— a few having remained identical. But those who receive the common view that every species has been created as we now see it, & that the same species has sometimes been created at more than one point of the earth's surface, may truly say with derision what complicated theories are required, such as the one just given, or the more simple one but requiring much greater geographical changes given by Dr. Hooker, to account on the theory of descent for the same, & with subsequent modification, for the representative species in these two distant areas. On the other hand those who believe in simple creation can, in my opinion, give no explanation in the least degree satisfactory of the shades of affinity & degree of identity in the cases which we have been discussing: they in fact simply state so it is.

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not scanty vegetation.3 According to all analogy, this antarctic vegetation from its isolation would have been very peculiar, but would have been in some degree related to that of the two nearest continents, America and Australia; & this antarctic vegetation though perhaps not nearly so uniform, as that now growing on the almost continuous arctic land, would probably have been4 internally related or already pretty closely related. As the glacial epoch came on, we may imagine that the seeds of these antarctic plants were carried in a N. Easterly course from their native home & landed on the southern shores of Australia, New Zealand, S. America & the several antarctic islands,— already chilled & ready to receive southern colonists. After the glacial epoch, as the climate improved, these antarctic plants would ascend the hills, where we now see them, some few remaining the same in these now widely separated colonies, some having undergone modification since their arrival, & some having arrived distinct, as they existed on their antarctic native islands.*(a)

It may be asked whether any other organic […] seem to have radiated from this[…]   [Bottom of folio torn off.]

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*(a)

3 In the southern hemisphere we have no distinct evidence that the climate was warmer during the older pliocene periods. The existence of burnt & silicified trees & thin beds of coal under the streams of lava in Kerguelen's Land & in [New South] Shetland Isld. (V. Dana's letter on Mr. Eights [Amer. J. Sci. 2nd ser., 22 (1856), 391]) where the vegetation is now so scanty should not be forgotten. I may, also, state that in Tierra del Fuego, (Geological Observation, p. 118) I found in beds underlying the drift or glacial deposits, many leaves of trees; which belong, according to Dr. Hooker to three species of Beech, apparently differing from the two species, which now clothe that forest-clad land.

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what complicated theories are required, such as the one just given, or the more simple one but requiring much greater geographical changes given by Dr. Hooker, to account on the theory of descent for the same, & with subsequent modification, for the representative species in these two distant areas. On the other hand those who believe in simple creation can, in my opinion, give no explanation in the least degree satisfactory of the shades of affinity & degree of identity in the cases which we have been discussing: they in fact simply state so it is.—

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them;

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Zealand; o fthese I can only say that they belong, (with the very remarkable exception of a Myosurus) to genera having species in many parts of the world, which offers some slight presumption that the species of such genera possess wide powers of diffusion.—

Dr. Hooker infers from the species in common, & more especially from the number of closely allied or representative species, that these two areas were formerly (Introduction p. xxiii) & xxxvi) connected by intermediate lands, but not necessarily continuous at any one time over the whole distance; he considers concludes that the relationship, indicated by the representative species (p. xxxvi) shows that the parts are "members of a once more extensive flora, wich has been broken up by geological & climatic causes." This land or islands must have been of considerable breath from north to south to or have lasted a long time. This land or other  islands must have been of

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We now come to our third class of facts, namely the existence of the same species of plants & animals on mountains distant from each other, & likewise on the lowlands nearer the pole, where the climate is nearly similar; as alpine plants & animals could not possibly migrate through the lowlands from one distant alpine point to another, until lately this, perhaps, was one of the strongest cases which could have been adduced by those who believe in the same species having been created at more than one point of the earth's surface. It is familiar to every one that several* plants grow, for instance, on the summits of our Scotch mountains, & on the lowlands of Northern Scandinavia & not in the intermediate low country. So the mountain plants of the Alps are separated by a space of [1000] miles1 from the northern land clothed by the same species. Similar observations have been made on insects.2

1 See A. Decandolle, Bot: Geograph: p. [1007-13] for an excellent summary on this subject.

2 Thus the Elaphrus Lapponicus which inhabits Lapland & Kamtschatka was found by Sir Charles Lyell on the Grampians: according to Erichson Tachinus elongatus is common to Sweden, Unalashka & the Alps of Switzerland. [Hooker here added:'Get from Adam White the alpine Himalayan & Tibetan insects of Thomson.']

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(note continued)

[2] So again Latreille (Memoires du Museum Tom. 3, p. 39) says that Prionus depsarius is found in Sweden & on the Swiss mountains; Lycus minutus in the boreal regions & in Cantal. —

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At the height of 8000-9000 feet on the Alps, we meet with northern grouse. But the most striking cases are afforded by mammals, for here we avoid accidental dispersions, as seeds borne by the wind or birds.— It would be a prodigy to find such northern animals as the Steinbock, which lives on the Alps at a height of 8000-9000 7000-9000 feet, or the field-mouse (Arvicola oeconomica) which lives at the height of 10,000 feet, or the Variable hare &c, in the low country at the northern base of these mountains: equally striking it is to observe in Johnston's Physical Atlas of Europe the small brown patches, marking the far separated & Alpine homes of the Chamois.3 Some few northern plants, but far more generally representatives of northern genera, have been observed on the more southern mountains of Spain & Greece; but to this subject we shall return.— †

These facts have been explained with beautiful simplicity by the late Professor E. Forbes: from the presence of the innumerable, ice-borne, great fragments of northern rocks scattered over the temperate zones of Europe,— from the former far lower descent of glaciers,— & more especially from the several Arctic shells imbedded in the drift, we know to a certainty that

3 [Pl]. 28.

† I have not yet noticed migration from N. to So. America [?] [CD.]

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But perhaps the most striking cases are those of the alpine mammifers, for we can here

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Europe, during a quite recent geological period, suffered under a severe climate; & Prof. Forbes believes that at this Glacial epoch, the seas & land were colonised by arctic forms, which when the climate became warmer, remigrated* northward, but where the land was high the plants, insects & mammals ascended the mountain-peaks & have lived there ever since. Hence it has come, that we now see the same forms at distant points, impenetrably separated by wide extents of surface fully occupied by the productions of the now temperate regions.

Prof. Forbes believed1 that the mountains of temperate Europe existed during the Glacial epoch as islands, & the seeds of northern plants were brought to these islands by icebergs.† That the greater part of Europe (& of Northern America) was under sea during some part of the glacial epoch is certain; but it is equally certain that this epoch endured for an enormous period, & that there were great contemporaneous changes of level,— as indeed most geologists would infer from the rocks scratched by floating ice at such different levels. I do not believe that there is any evidence to show

* for the most part migrated. ?perhaps created on the southern land & sea when cold is Forbes exact on this point? [comment by Joseph Dalton Hooker = J.D.H.]

† It is difficult to believe that during glacial epoch the northern land was warm enough for any plants at all. see Note A. [J.D.H.] [See p. 575.]

1 Memoirs of Geological Survey Vol I, p. 399 &c.

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whether the greater part of Europe existed as land at the commencement of the glacial epoch, so as freely to allow of the southward migration of northern forms; but that there was continuous land before the close of the cold period may be safely inferred from the distribution of the arcto-alpine mammals; & this is likewise confirmed by such cases as that remarked on by Mr. H. C. Watson,2 namely that the alpine flora of Britain is much more nearly related to that of the country northward of it, than to the floras of the Alps of middle Europe; if the remigration northward had been by various accidental means, this relation could hardly have been preserved.‡

We may indeed infer that the land in some parts of Europe stood even higher than at present, before the close of the glacial epoch, as is indicated by the presence of the Alpine Hare on the Scotch mountains, showing that Great Britain & Europe were then united. Let no one think that these great migrations south-ward & remigrations northward of a whole body of species are improbable, for it is hardly too strong an expression to say that in the case of the sea-shells the migration has

‡ very good— it is this confounded relation that obtains so much in more distant plants. [J.D.H.]

2 Cybele Britannica, Vol. p. 37.

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been witnessed3 by Prof. E. Forbes, so beautifully do the fossil shells on the shores of England & in the Mediterranean, compared with those living before & since, show the course of the migrations.

In North America we meet with similar facts of distribution. Dr. Asa Gray has most kindly given me a list of 59 plants (only 33 strictly alpine) growing near the summit of the White Mountains in N. Hampshire at the height of 5000-6000 feet. He believes that every one of them, with only two exceptions, inhabit Labrador situated 400 or 500 miles to the North: the great majority of these plants, viz: about 46, inhabit the circumpolar regions of Europe or Asia. No less than 33, or above half, grow on the Alps of Europe! about 22 of these same plants have been found on the summits of the mountains of New York, (separated from the White Mountains by a plain 60 or 70 miles in width) which afford a fitting site of only a few acres in extent:1 Even in the mountains of Carolina2 "a list of the shrubs & [herbaceous] plants [of this mountain] would be found to include a large portion of the common productions of the extreme northern states & Canada." But I do not know whether the Alleghanies do not now afford a highway* by which the plants could have travelled thus far south.

3 Memoirs of Geolog. Survey. Vol I. p. 385.

1 [In the fair copy, Darwin here wrote: '(Briefly contrast these with general proportion of whole U. States Flora compared to Europe.)' He probably wrote this and cancelled the rest of the paragraph, including the next note, in October, 1856, when he found new and better facts in the article by Gray discussed in note 3, p. 538. He added: '(Yes there is)' after 'highway' in the last sentence of the paragraph, probably also at this time.]

2 Dr. Asa Gray in London Journal of Botany. 1843. p. 114.— see Bartram for the Occone Mountains; Travels in N. America p. 335.

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(We shall, perhaps, better realise what formerly took place, if we imagine a glacial epoch now to come on again. The extreme arctic productions of all kinds are at present mostly the same round the pole: as the cold came on, whether or not strictly contemporaneously in Europe & N. America, then similar forms would slowly travel south, along the shores & land; & when the warmth returned they would return to their native north; but where there were mountains, as the ice & snow thawed & left the rocks uncovered, the northern forms would ascend, & would become surrounded by the stream of living beings flowing up from the south. And as the parent circumpolar productions were mostly the same, so would the alpine productions of the Old & New Worlds left on the mountain-summits on the returning warmth be to a great extent the same in the two worlds. We can thus understand the truly wonderful case of more than half the plants of the White Mountains being the same with those on the Alps of Europe, though separated by the whole Atlantic ocean, & on each side by a broad belt of low land, on which these Alpine productions could not possibly exist.— Had not the glacial epoch been brought to

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light & generally recognised, in main part owing to Agassiz, this case of the identical species on the White Mountains & Alps, might have been advanced as a grand proof of double creation.)

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(But in our recent imaginary change of climate we found the circumpolar productions mostly the same; & I should infer that this must have been the case before the real glacial epoch came on. Prof. Forbes believes1 that N. America and Europe were connected by continuous land, situated far to the north, during the glacial epoch, or towards its close: but the identity of several plants on mountains so far south as the Alps of Switzerland with those of America, seems (a) would have been covered by ice & snow, like the Antarctic islands at the present day under corresponding latitudes, & hence would not have allowed of the passage or of the existence of barely a single terrestrial production. Hence it is that I infer that the connexion was anterior to the Glacial epoch.*

* Certainly J.D.H. Note B [See p. 576.]

1 Memoirs of Geolog: Survey Vol I . p. 383 and 402.

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(a)

for plants coming over

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(a)

to show that the connexion between the Old and New Worlds had been established before the close of the cold period, otherwise the species common to America could not have got so far south as the Alps. During the most intense part of the glacial epoch, I can hardly doubt that land lying "far to the north"

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about 5002 of these British plants (approximately 1400 in number) are found both in N. America & Europe; & of these 500, 110 do not either in the Old or New World range into Mr. Watson's arctic & polar regions, i.e.; not further north than nearly the line of the arctic circle: 60 other plants range only on one side of the Atlantic, (generally on the European side, warmed by the Gulf-stream) into the Arctic Zone. Some, but extremely few,* of the plants common to the two worlds, do not range northward of the latitude of the northern point of Great Britain. But Mr. Watson informs me that since his publication in 1835 our knowledge has been much increased, & that the above numbers can be considered only as approximate. With respect to the sea-shells, common to the shores of Europe & N. America, I am informed by Mr. Woodward, that about one-third of their number do not range into Forbes' Arctic Sea,3 which washes the northern shores of Asia & America: but this one-third includes most of the doubtful cases, so that the number of species in common, which do not reach the arctic zone, is probably con-siderably less than one-third. As in the case of the plants, some 4 or 5 shells, which are common to the Old &

* Asa Gray thinks there are not a few plants common to U.S. & Europe, which do not range to Arctic regions. [CD.] "Certainly" J.D.H.

2 [Darwin later underlined 500, and in parentheses between the lines added: 'NO give Asa Gray's facts, far more accurate, and also wrote 'dele' in the margin of the lines giving Watson's figures, evidently preferring those he had found in the article by Gray referred to in the preceding note.]

3 See map, of Marine Life, in Johnston's Physical Atlas 2nd Edit. [pl. 31.]

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⁋ If we now look to a map of the circumpolar regions, we find near the Arctic circle almost continuous land & sea-coast from Lapland to Eastern America, & by going further north even almost to Eastern Greenland. Therefore if all the organisms, which are now common to Europe & America, could flourish under the present climate between the Arctic circle & 70° (& a great majority do now live there) I can see no insuperable difficulty to their having in the course of ages circulated round the polar regions by this course. No doubt the distance is very great, viz in the parallel of 70°, between 6000 & 7000 miles; but we know that most of the productions on this long line are now the same; & many species of fish & marine shells have even a wider range in the Indo-Pacific ocean.1

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New Worlds do not range even into Forbes' boreal province, & are therefore separated by the whole width of the Atlantic. But these shells (a)

 

 

Shortly before the Glacial epoch came on, during an earlier part of the pliocene period, when most of the organic beings were the same as now, we may fairly infer that the temperature of the* northern hemisphere was slightly warmer, perhaps more equable, than now; & as there can be no doubt judging from their southern limits that almost all arctic productions can well withstand a slightly warmer climate, I can see no great difficulty in supposing that all the organic beings, now common to the

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1 For Fish see Report to Brit. Assoc. for 1845 on the Ichthyology of the Seas of China by Sir J. Richardson, p. 190, 191.

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& the few plants similarly circumstanced probably belong to a distinct category, & were common to the old & new worlds long before the Glacial epoch: this, at least might well have been the case with some of the land shells in common, for these as I am informed by Mr. Woodward, are known to occur as older pliocene fossils. With respect to this more ancient connexion between the two worlds we shall have briefly to return.) ⁋

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two worlds, inhabited the long line of shore-land from eastern America to Lapland. As the glacial epoch came on, the species in common, associated with some not in common, would have migrated southward, & subsequently as the warmth returned they would have remigrated northward to their present homes. During these two great migrations, & with the local changes of climate which we might expect to ensue, it would be strange if several of the species did not become locally extinct. Hence we might expect to find in favourable situations, both on the southern high lands & in their latest northern homes, nests of species occuring elsewhere but not in the country immediately adjoining on either hand. Such nests seem to occur on the more temperate promontory of N. W. America, as I am informed by Dr. Hooker; and again near Lake Baikal2 in Lat: 52°, in the very middle of Siberia. In this same country, Gmelin3 gives several strong cases of plants with interrupted ranges & was thus led, even in the year 1747 to infer that

3 Flora Siberica. p. CIX, p. CX.

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*a

2 See M. Turczaninow in Bull.de la Soc. Imp. de Moscow. 1842, p. 15 account of this Flora. He is much surprised at half the Baikal Flora being the same with that round St. Petersburgh: it has, also, 452 species in common with Sweden: He specifies also species in common with N. America. See Gmelin, Flora Siberica p. cxiv, for the relations of the vegetation of the mountains near Baikal with that of Kamtshatka. See Ledebour (in Hooker's Miscellany. Vol 2. p. 241) for the similarity of the vegetation on the lower part of the Altai with that of Europe; & Sir W. Hooker in Linnean Transactions Vol X IV. p. 360.

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would appear from M. Martins' remarks1 on the decreasing number of European plants on the islands the further we go from Europe, that they were colonised by various accidental means (a view, however, which would be rejected by many of the most competent judges) from Europe, aided by the probable greater height & extension of parts of the European continent

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the same species had been created on diverse points of the earth's surface.

I have remarked that during the most intense part of the Glacial epoch, when arctic plants lived at the foot of the Alps, the most northern land & islands, as Spitzbergen, Iceland, Feroe, & Greenland, must probably have been icy deserts, like the Antarctic islands in the same latitude, & could have supported hardly any or no terrestrial productions. Hence these islands must have been colonised at a comparatively late period;* & it                    (              )

 

during the latter end of the Glacial epoch. Considering the lateness of the colonisation of these islands (for as explained in an early chapter on the view of species being formed by selection we can distinctly understand how time comes in as a most important element)† it ceases to be remarkable that islands so isolated as Spitzbergen, or Feroe, and Iceland, & seas like the Baltic should not possess, as I believe, a single endemic or peculiar inhabitant.‡

* Note C [J.D.H.] [See p. 576.]

† Greenland ought also to have alpine species. [CD.]

‡ Do Greenland or Lapland or Scotland even? [J.D.H.]

[repeat of the above]

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* It is possible that some few plants may have survived at the southern extremity of Greenland, in Lat: 60°. Four or five plants in Dr. Asa Gray's M.S. list common to Greenland & the White Mountains of N. Hampshire, & not found in Europe or Asia, perhaps indicate this; without indeed they were brought subsequently by icebergs.§

§ Which way?— great changes of level required. [J.D.H.]

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It may be objected to the foregoing view that Greenland, Iceland & Spitzbergen are inhabited by some mammals which could not have been transported by accidental means.— I have not seen complete lists of the aboriginal quadrupeds of these lands. In regard to wolves, foxes & bears, I am assured by Dr. Sutherland (see also Appendix I. pp 489, 494, by Dr. John Richardson) to Capt. Back's Journey) that they have all been seen alive on icebergs far out at sea, & their introduction therefore offers no serious difficulty; & indeed Mackenzie (Travels p. 341) asserts that the Black Fox is at present thus sometimes introduced into Iceland. Possibly the same explanation may suffice for the Mustela, (likewise carnivorous) said to have been seen on Spitzbergen (Richardson's Report 1836, on N. American Zoology p. 162) & for the Gulo in the Parry Islands. Whether the Rein-deer, & the Lepus glacialis, which is found in Greenland, & the Mus oeconomicus, said to be an inhabitant of Iceland, & the rodents on which the Mustela must feed in Spitzbergen, could have crossed on ice, I know not; but the skeleton of the Lemming, found by Capt: Parry on the ice in Lat: 81 3/4° N. should not be forgotten. Even at present Iceland is connected with Greenland in the spring by continuous pack-ice.

 

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(On the other hand, at the very first return of the warmth, long before the northern islands could have been colonized, the arctic productions at the base of the more Southern mountains, such as Pyrenees and Alps, would have been completely isolated, as on an island in the sea, as soon as they had ascended the lower slopes of the mountains. Previously to the glacial epoch, these mountains must have had their Alpine species, such as Gentians &c which do not inhabit the arctic regions, & these on the returning warmth, after having, as it would appear, spread over the surrounding country* would together with the arctic species have reascended the mountains.   Here then we have, according to the principles laid down in our fourth chapter, all the elements present which tend to modify species, though not in the highest degree,— namely, considerable lapse of time, isolation, & especially association with somewhat different sets of organic beings. Hence we might have expected that there would have been many representative species & strongly marked varieties, on the several alpine summits of Europe,‡ when compared one with another & with the arctic regions. § I infer that this is the case from various scattered notions; & to give one

* Note D [J.D.H.] [See p. 576.]

‡ Dr. Hooker: I wish I knew whether this was so: Forbes thought so, but I do not know whether he is to be trusted. [CD.]

Certainly true J.H.

§ Note E [J.D.H.] [See p. 576.]

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* It would appear from Ledebour's account (Hookers Miscellany, Vol: II p. 241, 249) that the Alpine vegetation of the Altai in about Lat: 50°, had been able to keep itself during the glacial changes of climate unmixed, owing perhaps to the peculiar character of the climate of the steppes to which the old vegetation would have been adapted. Ledebour says that at the height of 4500 feet the vegetation has a greater similarity to that of Europe than on the surrounding plains, though some of the peculiar steppe-forms are yet found: between 4500 and 6500 feet, the European species gradually diminish in number and give place to the proper flora of the Altai.†

† Please give me this reference. I should like to know if these proper species are varieties of Arctic. J.H.

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example, namely the chamois, which some Zoologists think specifically distinct on the Alps & Pyrenees, & others merely a variety of one & the same species. The arctic forms, which during their migration southward and remigration northward, did not become isolated & so differently associated, but kept in a body together under nearly similar conditions, would have undergone according to our principles very little modification.*(a)

The views here given may perhaps be extended further, though in doing so we are trespassing on the next chapter & considering representative species. If we compare the temperate productions of the lowlands of Europe & N. America, we find in all classes, terrestrial & aquatic, a vast number of species of the same genera, many obviously representatives of each other on the two sides of the Atlantic, & filling exactly the same place in the economy of nature, & not a few so closely allied, that the most practiced naturalists doubt whether to consider them as varieties or as true species. Many examples could be given in every single natural class of these doubtful species, & of quite distinct but representative species. If we look further west across the northern Pacific to Japan, we find many

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* note to p. 13.)

Ireland would have been in chief part colonised at a period between the isolation of the [illeg] apline the productions of the alps & the colonisation of the northern islands & the isolation of the production of the more southern alps

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* Ireland, according to Prof: Forbes, was insulated at an earlier period than Great Britain, & indeed it is scarcely possible to look at W. Thompson's tables of distribution & doubt this. Its climate would have become fitted for its present productions at a later period than the isolation of the more southern alpine forms, but at an earlier period than the colonisation of the northern islands. The fact that there are several doubtful cases of representative species, or very strongly marked varieties in the animals & plants of Ireland, seems to accord with this.

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most striking1 representative species of European, but more especially of American* genera of plants, of mammals, birds & other beings. To complete the circle of the temperate zones, I may just allude to the many closely allied & a few perhaps even identical species of Crustacea in the seas of Japan & in the Mediterranean, as remarked on by Prof: Dana in his admirable Report on Crustacea:2 yet the Medi terranean & Japan, even if we submerge the isthmus of Suez, are separated by a hemisphere of equatorial ocean. I may mention that I was myself much struck by finding two very close & obviously representative species of a very rare genus of parasitic cirripedes on crabs, from Madeira & Japan. Some of the fish, also, from Madeira, as I am informed by the Rev: R. B. Lowe represent those of Japan.†   & I am afraid by Sir John Richardson

* & Himalayan! [J.D.H.]

† Note F [J.D.H.] [See p. 576.]

[C.D:'s pencilled addition:] Refer to Asa Gray's most striking tables.

2 United States Exploring Expedition, p. 1552, 1567, 1586.

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* 1 See the account of Zuccarini's observations in Silliman's Journal             [see Amer. J. SCI., 39 (1840), 175-6; 52 (1846), 135-6.] Decandolle has, also, insisted strongly on the representative species of Europe, America & Asia in the Dict. des Sciences naturelles. Art. Geograph. Bot. p. 414 — 1820  

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Now if the view before given is in some degree probable, namely, that just before the glacial epoch when the climate was very slightly more favourable, there was nearly continuous circumpolar land & coast, as at present, inhabited by a nearly uniform flora & fauna; then it is not so very improbable that still earlier, during the older pliocene or even Miocene period, when the climate in these high latitudes was temperate,* there was likewise land & shore to some extent continuous, whence the closely related and often identical organic terrestrial & acquatic productions might have migrated southward, as the temperature fell, but long before the glacial epoch. As soon as this southern movement had taken place, the several existing floras & faunas of the northern hemisphere would have been separated from each other, as at present, would have been differently associated together, & exposed to somewhat different conditions. And as we are now dealing with comparatively ancient times, we might expect, according to the principles which we are testing in this work, that only a few species of those originally in common would have remained absolutely identical, but still we might expect plainly to see in the productions of the land & seas of temperate Europe, N. America, & Eastern Asia, evidence of their descent from a common home & common parentage; and this, I believe we do see, in the many representative species of these now quite separated countries.)

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*a  Whether the abundant vegetable remains found by Sir John Richardson (Geological Journal, Vol [ ]) in the extreme arctic regions of America, belonged to the older pliocene age, or to a later period but anterior to the glacial epoch, is not known.—

(8 (16

(We have seen that Prof. Forbes' theory explains the distribution of the alpine productions of Europe, in a manner which I think must be satisfactory to every one; & can be extended with no very great difficulty, as it seems to me, to N. America. But I believe that the theory is capable of a much greater extension. In South America I was formerly2 much struck with finding numerous boulders on the island of Chiloe in Lat: 42°, where the rankest forests are now intertwined with almost arborescent canes & these gigantic boulders had been carried on ice from the Cordillera across a wide arm of the sea: on the plains of Patagonia, on the opposite side of the Cordillera, in Lat: 50° and over the southern extremity of the continent, & on the Eastern end of the Falkland Islands, boulders are very numerous. In central Chile, on the road to the Portillo Pass, I examined a mound of detritus which at the time never having read of Moraines greatly perplexed me, but now I can hardly doubt that it was a terminal moraine situated thousands of feet below the line where a glacier could now descend.1 On

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* 1 This mound blocking up the valley at the lower end of the lake-like expanse of the "valle del yeso," seemed to be composed wholly of alluvium, & was apparently 800 feet in thickness; "its surface consisted" (to quote my original notes) "of a confused hilly mass of rounded & angular fragments of rock, many of the latter of very large size."

2 Geological Transactions. Vol VI. p. 424 (1841)

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they do not appear to occur; the surface, perhaps, at this period not having been under the sea: but Prof. Forbes' theory, of the community of alpine & arctic species having been caused by a former cold climate bears so strong an impress of truth, that where there is such community, we may almost safely turn round & argue from it, that the climate has been colder: if so, there can be no doubt from the several cases enumerated in a former note that Siberia has suffered from a cold climate.

(9 (17

the Cordillera of equatorial America, the marks of the former lower descent of glaciers have been observed.2

[passage not transcribed by Stauffer:] It seems to me that Prof. Forbes theory of the community of species on distant mountains distant from each other, being due to their having formerly ranged under a cooler climate over the intermediate country is so satisfactory, that where this uniformityof alpine species occurs we may infer former coller climate: if so this may be inferred in regard to the Isthmus of Panama, for Mr. Seeman found this on the mountains, Mexican plants (?).—

In N. America, on the eastern side we have the plainest geological evidence of glacial action, as far south as Lat: 36°-37°. On the high plains near the Rocky Mountains boulders have likewise been observed; & on the shores of the Pacific in Lat: 46°.3

In Europe erratic boulders extend to near the Western base of the Oural, & in parts, southward to Lat: 45°-46°. In Siberia4

2 Bull. Geolog. Soc. [Acosta, Bull. soc. geol. de France Sér. 2: 8 (1851) 493. 9 (1852) 398.

4 [See appendix for passage cancelled in draft here & replaced by fol. 17a.]

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3 Dana: Geology of the United States Exploring Expedition. Vol X. p. 674.

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Looking south we find in the Himalaya abundant evidence5 of the former much lower descent of the Glaciers, which have left behind them enormous Moraines. In India, using Prof: Forbes' theory, we have some evidence of a cooler climate in several plants,6 (& in some mammals according to Mr. Blyth) being the same on the Nilghiri with those on the Khasia mountains & on the Himalaya; & again some* Dr. Hooker believes abundance of plants on the Nilghiri7 are common with those on the mountains of Ceylon & the Himalaya.* There is much affinity, more especially as shown by the many European genera of plants & some species in common, between these Indian mountains1 & those of Sumatra & Java;2 the case of the Mydaus here comes into play, a quadruped found at the height of several thousand feet on the isolated volcanic mountains of Java, and never in the hot and low intermediate country.3

I have never heard of any marks of glacial action in S. Africa or in Australia; but in New

* Dr. Hooker believes [CD.] abundance of plants [J.D.H.]

* Note G [J.D.H.] [See p. 577.]

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5 Dr. Hooker— Himalayan Journal. Vol I. p. 248, 380.

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7 Gardner— in London Journal of Botany, no. 47. 1845. Also Gardner in Journal of Horticultural Soc.— Vol IV. p. 37. Southward of the Nilghiri on the Pulney mountains (Madras Journal of Literature and Science. Vol V. p. 283) according to Dr. Wight, several of the same northern genera are met with at the height of 8000 feet.— [Addenda:] Flora of Pulney Mts. is identical with Nilghiri [J.D.H.] Also peculiar land-shells nearly or quite the same in mountains of Ceylon & Nilghiri [CD.]

2 F. Junghuhn in his Java, Seine Gestalt, Pflanzendecke (1852) Vol I. p. 417, gives a long list of the genera found at the height of about 7000 feet, the names of which are familiar to every European. Dr. C. Reinwardt (Journal Hort. Soc. Vol IV. p. 233.) says the vegetation of the mountains of Java brings strongly to mind "our native home", but he asserts that all the species are distinct. With respect to Sumatra Temminck speaking of one of the mountains (Coup d'Oeil gen. sur les Possessions Neerlandaises Vol 2. p. 82.) says "La végétation sur son sommet porte tous les characteres [sic] des plantes alpestres de Europe"

3 Sir C. Lyell, Principles of Geology. p. [638-9] Temminck says (Coup d'Oeil sur la Faune des iles de la Sonde p. 13) that the Turdus varius is common to the mountains of Java & the lesser heights of Japan.

6 Dr. Hooker— Flora Indica, p. 87, 99.

1 Flora Indica, p. 104 Dr. Hooker says "constantly, during our examination of the temperate as well as tropical plants of the Nilghiri, Khasia, Ceylon & the Himalaya we find them identical in species with Japanese mountain plants."

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Zealand, Mr. W. Mantell has shown me sketches of great fragments of quartz, lying on tertiary strata, which probably are erratic boulders: I saw myself boulders near the Bay of Islands, which appeared to me at the time as possibly of glacial origin. Dr. Hooker, moreover, informs me that there are certainly many plants common to the mountains of New Zealand, & not inhabiting the intermediate plains: some likewise are common to the mountains of New Zealand & Tasmania, & likewise to the lesser heights on the islands lying south of New Zealand: here then at the Antipodes we have the same sort of evidence of a cooler climate4 as in the northern hemisphere.

With respect to the period of the glacial action or of the cooler climate at these several & very distant parts of the world, we can at least say that it has been in a geological sense recent: the phenomena are superficial; the evidence from scored rocks & moraines shows no great changes of surface have taken place since the moving ice covered the rocks. In most of the cases the glacial period has certainly supervened during

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4 See Dr. Hooker's Remarks on this subject in the Introduction to the Flora of New Zealand. p. xxiii.

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the existence of the majority of living plants, & in the case of Europe & N. America, of living sea-shells. In Europe & N. America there certainly seems to have been a close parallelism in the whole phenomena of glacial action & in the coincident changes of land, but I am well aware that this does not prove strict contemporaneity. (a)

[passage not transcribed by Stauffer:] the remarkable fact, shown by Sir C. Lyell that some now extinct mammals survive the glacial epoch in N. America, which is not known at present to have been the case in Europe, seems to indicate some difference. According to the standard of extinct mammifers, there is some evidence* that the glacial epoch of South & North America were more strictly coincident.

 

 As the northward curvature of the lines of equal temperature at the present day in Europe, compared with N. America is due to the warmth of the Gulf-stream, Mr. Hopkins2 has inferred that the glacial epoch of Europe was probably caused by the Gulf Stream having formerly flowed up the central parts of N. America; how far the greater extension southward of the glacial action in N. America than in Europe, & likewise,3 (as

2 [Geol. Soc. Quart. J., 8 (1852), P. lxiii.]

3 Sir C. Lyell Travels in N. America, 1845, Vol I. p. 139. This inference is drawn from the comparison of the tertiary fossil shells with those of Europe.—

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(a) text

In N. America, according to Sir C. Lyell, & in S. America & in Europe some large mammals have become extinct since the glacial period, namely the Mastodon, Cervus & Megaceros in the north & Macrauchenia1 in the south.—

1 Darwin's Geolog: Observations on S. America, p. 97.

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the apparent parallelisms of the miocene isotherms compared with the existing isotherms agrees with this theory may be doubted: for it might be argued from these facts that probably the course of the Gulf-Stream had long been constant. Seeing how similar the superficial glacial phenomena are on both sides of the Cordillera of Southern America with those on both sides of N. America & bearing in mind that there is some evidence of glacial action in central Chile & all along the Cordillera & in the Equatorial Andes, there seems to me a prima facie probability that both Americas were cooler strictly at the same time. No one who has carefully examined the effects of glacial action

The vast number of boulders borne by icebergs & widely scattered— the thick masses of drift,— the great coincident changes of level both up & down,— the enormous amount of denudation,4 — all bring this conviction most forcibly home to the mind. When I state that there seems a prima facie probability that both Americas were at the same time cooler, I am far from wishing to infer that the cold in the North & South either began or ended at the same date, but that a part of the

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*Q Travels in N. America (1845) Vol I. p. 139

 

(a) text

when the phenomena are well developed, can doubt, that the cold, though in a geological sense recent, endured for an enormous lapse of years.

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*b One is always most impressed with what one observes

[21V]

4 At the head of the S. Cruz river in Patagonia ([Darwin] Geolog: Transact: Vol VI. p. 417) the great plain 1200 feet in height is strewed with boulders; this plain has been worn into a vast bay-like depression, facing the distant Cordillera, & is fringed by an 800 ft plain, also strewed with boulders, & the bay-like depression itself 440 feet above the level of the sea, again has great angular blocks on its surface, but different in kind from those on the upper plains. Here we see that the hard basaltic rocks of the upper plains which form on each side exactly corresponding strata, have been cut away; the intermediate plain has been formed: the whole country has been elevated from the height of at least 440 feet to 1200; & all this has taken place, whilst ice was transporting in Lat: 50° great boulders from the Cordillera. How many hundreds of thousands of years must have been required!

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long period in the North & South was strictly co-incident, so that the intermediate zones were at the same time somewhat cooler. It is immaterial for our present discussion, whether the whole world was at one geologically recent time slightly cooler than at present, (as I confess seems to me most probable) or whether the two Americas were at one time cooler; Europe & Africa at another time; Siberia, the Himalaya, the plains of India, the eastern Archipelago & Australia at another time, but all within the age of the pleistocene formations.

Finally to sum up,— it is far from proved that any part of the cold period at these several & distant regions, was strictly co-incident, contemporaneous, either over the whole world or along a few great meridional belts, indeed this is almost beyond the scope of simple geology; but there seems sufficient probability in this view, so that if it will explain several phenomena of organic distribution, otherwise inexplicable, it may be accepted as a theory worthy of consideration.

Let us then assume that at the period when the northern & southern portions of the world were colder than now, that either the whole, or first one & then another meridional belt of the intertropical regions was rendered slightly

(15 (23

cooler, & what would be the result? The inter-tropical productions would retreat into the hottest districts; their proportional numbers would probably be considerably altered; some would become extinct; some, according to the principles which we are testing, would become modified. But according to these same principles, it may be doubted whether there would be very great modification; in as much as the great mass of surrounding organisms would remain the same, & we have seen reason to believe, that although changed conditions will cause variability, the selection of new specific forms is far more intimately related to the surrounding organic beings, amongst which each has to struggle for existence, & to seize on & occupy by selected changes in its structure any vacant place in the economy of nature.

For the same reason I should not anticipate very much extinction during the cold period within the Tropics, for we have seen good reason to believe that extinction depends far more on other organic beings seizing on the place of the dying forms than on changed conditions; & indeed we know that most organic beings, plants for instance, will endure a considerable change of climate, if protected from competing forms.

On the frontiers of the Tropics, the whole body of the temperate productions would invade, from the north & south, the cooled land; & as

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(a) I have previously remarked that on the same principles I should not expect great modification in the arctic species during their migration southward & remigrations northward, for they must have migrated in a body. To explain further by a metaphor what I mean: if a whole nation migrated in a body, each might retain almost his usual habits & business, but if only a few settled in a foreign land each probably would have more or less to change his habits, & occupy a different position in society.

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all the intertropical productions would be in some slight degree distressed, I can see no great improbability in some few temperate forms penetrating even to the equator and holding their own.    text
be eminently liable to have every slight variation, by which they would become still better adapted to struggle with their new compatriots, selected, & their structure thus specifically altered. (B)

It is obvious that chains of mountains & high land would greatly favour the invasion of the temperate forms. One of the most obvious objections to the theory, is the enormous migratory power, though over continuous land, thus attributed to the temperate forms.

(We will consider some of the most obvious objections to this theory, after we have seen its local applications. First, for America: no one doubts that during the glacial epoch the northern portion was inhabited by many old-world forms, the introduction of which we have already discussed. These would have a broad and eminently favourable high-road for migration southward, during this colder period, as far

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* We might expect to see a vegetation like that so strikingly described by Dr. Hooker1 at the base of the Himalaya, where true Tropical forms are mingled with such northern forms as Birches, Maples, whortle-berries, strawberries &c. Chains of mountains and high land running north and south would obviously favour the invasion of the temperate forms.

After the glacial epoch, [passage not transcribed by Stauffer:] when the Tropics resumed their present temperature, these interlopers would be all destroyed, except where they could ascend high land; & no doubt during their first invasion they would always seize on the more temperate portions. Here surrounded by strange forms & placed under new conditions, even since the close of the glacial epoch, & even during early portion of it, they would

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*a (note to back of p. 24)

1 Himalayan Journal, Vol I. p. 109. Vol. II. p. 319. For similar remarks, see Royle's Illustrations of the Botany of the Himalaya, p. 14, The Rev. F. W. Hope (Entomology, in Royle's Illustrations) describes a similar mixture of insect forms: he says (p. 15) we find in the valleys of the Himalaya where tropical forms abound, "European types & species in numbers sufficient to excite our astonishment." I may add as further showing the possibility of the commingling of tropical and temperate forms. that Lichtenstein (          ) [: see Akad. Wiss. Berlin, Abh. 1838, p. 422] states that this is the case with the birds in parts of Mexico.—

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(B)

Though thousands of years might be required for their passage through the Tropics, slowly advancing as the cold came on, I can believe that they would not so much tend to be specifically altered, as afterwards when permanently settled on some isolated mountain associated with new organic beings: for in our chapter on natural selection it has been shown how excessively slow this process must be, counteracted as it must be in many ways; & that under changing conditions it could effect comparatively little, just as a breeder would be infinitely delayed if he changed his object or standard of perfection.)

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as near the isthmus of Panama. As just stated I can see no great difficulty in some temperate forms passing this hot & low barrier;1 but it may have been then higher; we know, at least, that the isthmus has existed since the creation of the two distinct marine faunas on its two sides; & off Yucatan the coral-reefs favour the idea of considerable subsidence. After passing the isthmus the temperate forms would find in the Cordillera a grand line of communication to the southern part of the continent, as suggested by Dr. Hooker,2 who supposes that at the period of migration the Cordillera were loftier; & therefore more temperate: geological evidence, from the equator southward, as far as it goes, is opposed to this view, & I think all the facts are better explained by change of climate, of which we have much independent evidence.

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* 1 In Mr. Seeman's Narrative of the Voyage of the Herald (Vol. I. p. 253.) it is said that on the mountains of Panama, at the height of 2000 feet the vegetation resembles that of Mexico "with forms of the torrid zone harmoniously blended with those of the temperate."

2 Introduction to the Flora of New Zealand, p. XXV

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(As the climate became warmer, towards the close of the glacial epoch, we may readily believe that nearly all the northern temperate species would be destroyed on the mountains of southern Peru & northern Chile,3 owing to the extreme aridity of their present climate; for in Chile even at the greatest heights, glaciers are now hardly formed. As soon as the ice & snow, with which Tierra del Fuego was probably covered during the intensity of the glacial epoch, disappeared, this southern point of the continent would have been clothed with plants, including the northern temperate forms, which had travelled down the Cordillera. Some of these plants would be left on the mountains, where the climate was fitted for them: of this I saw one instance in Chiloe, where at the height of about a thousand feet, the well known antarctic beech of Fuegia lived in a dwarfed condition. Thus, I think,

 

Note

*aa  The alpine vegetation seems now to be very peculiar on these great mountains, but has been only very imperfectly described: see Meyen's Reise vol.          [: see I, 348, 466.] & Poeppig's Reise vol.

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lived with new associates. But those northern forms which found a suitable home on the lofty mountains of America1 must have been associated for a still longer period with new beings, namely with the American alpine forms which we cannot doubt existed previously to the glacial epoch; they must, also, have been exposed to still more different conditions, & hence we might expect that they would have undergone greater modification than those of Tierra del Fuego, and this I suspect is the case.2)

2 [Here CD. scribbled a memorandum:] See some paper in Eding. New Phil. Journal?

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[not transcribed by Stauffer] Chiloe. Still further south they would have migrated to the extreme south of the continent, now rendered comparatively temperate. Hence
we can understand the presence of so many European forms in Tierra del Fuego, as is so forcibly shown to be the case by Dr. Hooker in the Flora-Antarctica, some absolutely identical, some presenting strongly marked varieties & some quite distinct, but still plainly related to their northern congeners. According to the principles discussed in this volume, we might have expected considerable modification in these forms which have wandered so immensely far from their native home, and which have

[not transcribed by Stauffer:] been at different times differently associated with many new &   lived with new associated & with different
But    Those northern forms which found suitable homes on the lofts mountains of equatorial American,* would must have been exposed to every still m  difference conditions & would be associated for a still longer time period with new beings, namely with with American Alpine to American alpine forms, which we cannot doubt existed previously to the glacial epoch ; they must, also, have been exposed to still more different conditions; been these here we might expect great the [illeg] form would have suff   more modification & I
undergone greater modifications in the than in T. del Fuego, & I believe

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* 1 In Johnston's Physical Atlas. Botan: Geography it is said that of the 327 genera of plants found on the declivities of the Andes, at the height of 7000 feet & upwards, 180 genera, or more than half, are common to the temperate zone.—

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(Those few temperate forms which were able to penetrate the lowlands of Tropical America during the Glacial epoch, would be most of all modified, & when the climate again became hot, could only survive on high land: thus, perhaps, we can understand the presence of species of such temperate genera as Vaccinium, Andromeda, Gaultheria, Hypericum, Drosera & Habenaria found by Mr. Gardner3 between 6000-7000 feet on the Organ Mountains of Brazil. It would appear that some truly American alpine forms had descended & spread over the plains of S. America during the cooler period; for thus apparently can only be explained the presence of the Andian genus Bejaria, & even the same species of Thibaudia on the Silla of Caraccas4 & mountains of New Granada, where they are associated with some of the same genera found on the Organ Mountains & on the heights of Jamaica.

Now let us turn to Africa & briefly consider the period whether or not strictly coincident with the cold period of America, when arctic forms were living at the foot of the Alps. At this period I believe those few northern temperate species, which are now found on the highland of Abyssinia1 penetrated to that latitude though so near the equator. In Drege's enormous collection of plants from the Cape of Good Hope

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3 Journal of the Horticult: Soc: Vol I. (1846) p. 281. Mr. Purdie found [on] the mountains of Jamaica (London Journal of Botany Vol. III p. 512.) Vaccinium, Andromeda, Myrica Mexicana, & Viburnum.

4 Humboldt— Personal Narrative (Eng. Translat.) Vol III. p. 494, 500.

1 A. Decandolle, Geograp. Bot. p. [ ] To the West of northern Africa on the heights of Teneriffe a very few northern species, & several northern genera have been found; & lately, as I am informed by Mr. Wollaston, Erica cinerea has been found near the summit of Madeira. To the East in Lycia at heights between 6000 & 10,000 feet (Lieut. Spratt and E. Forbes Travels, [II] p. 157) Draba aizoides, Anemone Appenina, Scilla bifolia &c are found.—

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as described by E. Meyer.2 there are 96 European phanerogams & ferns enumerated. Mr. Bunbury, who has personally collected at the Cape, has kindly looked over the list for me & has added three species: he considers many of these plants as probably naturalized by man. Some are littoral plants which may possibly have travelled by the coast; about 14 are aquatic or marsh plants which seem to have, as we have seen, some special means of diffusion; but 30 plants apparently do not come under either of these categories & I should infer (if really not naturalized by man's agency) had migrated through the tropics during the cold period. Considering the ordeal they must have gone through in having been so long associated with the very distant Cape species, this number is too great for my theory. If there exists, as some have supposed, near the East African Coast nearly continuous high land from Abyssinia to the Cape, their migration at least, into this colony would be rendered more probable. The fact that on the very arid & somewhat isolated mountains of the Cape, at the height of from 6000 7000 to 8000 feet, there are some distinct species of such northern genera3 as Geum, Epilobium, Pimpinella, Galium, Tanacetum, Myosotis, Dianthus & Anemone, associated with many species of Cape genera, harmonises better with the theory.

In the East, at the time when the glaciers descended low on the Himalaya & the prodigious moraines described by Dr. Hooker were forming, & when probably the woolly-covered Rhinoceros tichorinus and Elephas primigenius* were ranging over the

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2 Flora 1843 Band II. Zwei Pflanzengeograph. Doc. p. 9.

3 Flora 1843 B. II p. 53 Mr. Bunbury thinks that the genera Dianthus, Franklinia Statice are the most striking cases of northern genera having representative species at the Cape. The Heaths offer a well known case, abounding at the Cape, & not known to reappear in the north nearer the Equator than Teneriffe & Arabia; but the species from the north & south I believe, show no especial affinity.

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plains of Siberia, I must believe that those plants, already alluded to as common to the Himalaya, the Nilghiri, & to the mountains of Ceylon & of Java, ranged over the intermediate now torrid country; & that during the cool period they reached & subsequently ascended their present isolated & elevated homes. On the Himalaya, Dr. Hooker has shown that many plants are representatives & many specifically the same, (though often presenting varieties) with those of the regions lying north of them & of the European mountains; & this migration might well have happened during the cool period considering the latitude of these great mountains, & more especially the high but broken land to the north and northwest.1 The majority of the species of northern genera on the Nilghiri & on the heights of Ceylon, as I infer from the writings* of Mr. Gardner2 are representatives, as would ensue from their having been differently associated as compared one with another, having been, as compared with the Himalaya, isolated for a longer period, owing to their more southern position. Considering that Java, Sumatra & Borneo lie near each other & arise from a shallow bank3 & that they have

* No I think majority identical [J.D.H.]

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2 Journal of the Horticult. Soc. London Vol IV. p. 37. A short table is given showing that many more species are representative than identical. ['Gardner however could only guess & not compare enough.' (J.D.H.)]

1 We find exactly the same class of facts in the insects of the Himalaya. Mr. Hope (Entomology, in Dr. Royle's Illustrations) seems continually in doubt whether certain insects are identically the same or most close representatives of those of Europe and Siberia. Amongst Birds we have both identical species & some beautiful representatives of those of Europe, as in the bull-finch, goldfinch, shrikes &c, as represented in Mr. Gould's Century of Birds from the Himalaya. So again it is with many mammals, & Mr. Ogleby [Ogilby] seems in doubt (Royle's Illustrations) in regard to some of the mustelae, badgers, hedgehogs &c whether to consider them identical or representatives.

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[line not transcribed by Stauffer:] considering that Java, Sumatra & Borneo lying near each other, all [illeg] from a quite shallow bank.& a & that they have
some few mammals in common there is a strong probability that the whole area within recent times may have stood at a higher level & been continuous; & therefore there is little more difficulty in the heights of these great islands having been colonized by northern forms, since modified, than in Ceylon having been thus colonized.

We now come to a more difficult case. Long since Robert Brown showed that there were several northern plants in Australia, which could not be considered as naturalized by man's aid. Recently, Australia during (B) [deleted passage not transcribed by Stauffer:] ; & recently Dr Hook, also, has given ample details showing that in New Zealand there exist (New Zealnd Flor. Introduct p. XXX) [illeg]      northern species 60 plants, identical (but often under the form of [illeg]) with those of Europe. These  It is possible that there are should [illeg] I should be inclined to suppose to infer that these plants had   migrated into new

the cold period (when the mountain plants in common to Tasmania, the Australian Alps & New Zealand inhabited the low grounds) by the islands of the Malay archipelago: perhaps through New Guinea; but the vegetation of the lofty mountains of this island is unfortunately quite

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3 Windsor Earl, on the Physical structure of the Indian Archipelago. Geograph. Journal, 1845. Vol 15. p. 358. [Cite not precisely in text— could be for map facing this page.]

[31V]

(B) text.

Dr. F. Muller has found on the Australian Alps several European plants, as Lysimachia vulgaris, Turrutes glabra, Veronica serpyllifolia, which species together with some others mentioned to me by Dr. Hooker, are not common in Australia & are found no where else in the southern hemisphere & are not very widely distributed in the northern hemisphere. I should suppose that these plants had migrated into

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unknown. Between New Guinea & Java, where northern forms are found, the sea in parts is deep, but it is studded with an extraordinary number of islands, so that by strides of 50 miles the interspace can be crossed on dry land; & there have good evidence of prodigious recent subsidence in New Caledonia2 (of which the mountain vegetation is unknown) & at the S. E. extremity of New Guinea, & that between New Zealand & New Caledonia (about 800 miles apart) there are some islets. But the number of species & genera of plants common to New Zealand & Australia though nearly 900 miles apart at their nearest points perhaps should lead to the belief that New Zealand had in chief part derived its northern forms through Australia.3 New Zealand undoubtedly offers another great difficulty to the views here advocated, & is a strong case in favour of those who believe in multiple creations.-)

[bottom half of page not transcribed by Stauffer:] (a)

In the whole of  This discussion, we have as yet hardly chiefly considered  has as yet been   almost confined to plants alone, for they are better known & the same species range further than almost any other organic beings   with the exception of    Before passing to marine produtions, I will say a very few words in regards to other terrestrial productions, as far as they relate to distribution during the cold period.

In mammals; land-brids & reptiles, I know of no cases

2 The barrier coral-reefs show that the island formerly extended 150 geographical miles further at its northern end.

[32v]

2 The barrier coral-reefs show that the island formerly extended 150 geographical miles further at its northern end.

3 Dr Hooker believes that New Zealand & Australia were united within the period of existing species; I have already given my reasons for not being able to admit this view, which would remove many difficulties, but at the same time cause some others. But I am far from wishing to deny the possibility that there may have existed larger & more numerous islands in the intermediate sea; or that the main coasts may have formerly extended somewhat nearer to each other.—

[32V]

is some evidence in parts of subsidence. The identity of the above specified & several other Australian plants with those of Europe, is certainly most remarkable, considering their long sojourn amongst foreigners; & is a parallel case with the European plants at the Cape of Good Hope. It might have been anticipated in both instances that fully as many plants would have undergone specific modification as on the mountains of Java or India.)

(In New Zealand Dr. Hooker has found1 60 European plants and in addition several striking cases of representative species or as I should consider them modified forms of northern genera. With respect to the introduction of these species, I will only remark that we

1 New Zealand Flora: Introduct. p. xxx.

[33A]

appear to be many species in common even on mountain-summits as near to each other as the Alps & Pyrenees, or, as I am informed by Mr. Benson, on the Nilghiri & heights of Ceylon. Nor should we be surprised at this, when we hear from so competent a witness, as Prof. Adams that in Jamaica, the collector in the course of every ten miles finds new species. In regard to insects, I carefully collected the beetles of Tierra del Fuego, & Mr. Waterhouse has examined them; but none are identical with, or cIosely

(33

(This discussion has as yet been almost confined to plants, & I will now make a very few remarks on other organic beings in relation to their migration from north to south during the glacial epoch. In mammals & reptiles, I know of no cases of the same or representative species being found in the opposite hemispheres & not in the intermedial Tropics. In Australia, as I am informed by Mr. Gould, there are some striking cases of Birds, chiefly aquatic, as the Australian coot, moor-hen, & some ducks which represent northern forms, & are not known to occur within the Tropics. In land-shells I can hear of no northern & southern identical or representative species; and this could hardly be expected; for land-shells have either been so frequently created, or as I should infer so easily modified, that there do not  [cont. on 33A]

(33

I will now make a very few remarks on the

(25 (34

representative of, northern forms; Carabus, however, must be excepted, as it seems to have travelled, like many Fuegian plants, along the Cordillera from the north. In southern Australia & New Zealand, there are only a few very doubtful cases of representatives of northern forms. But it should be observed that insects are not nearly such wide rangers as might have been anticipated.

[34v]

[text excised] Report on Fishes

[34a]

(Turning now to marine productions, we hear from Sir J. Richardson,1 that Arctic forms of fishes disappear in the seas of Japan & of northern China, are replaced by other assemblages in the warmer latitudes & reappear on the coast of Tasmania, southern New Zealand & the antarctic islands. He further states that the southern cod-fish are "much like those of the north, & Notacanthus & Macrourus, two very remarkable Greenland genera, which inhabit deep water, have recently been discovered on the coasts of New Zealand & S. Australia." In regard to sea-shells, Dr. A. Gould2 says proceeding from the north, across the equatorial seas, "there is not a return to the same species & rarely to the same genera". But he adds: "along our northern seas, some of the most characteristic shells are Buccinum, Tritonium, Fusus &c. Around Cape Horn are shells of the same types, so closely allied that they have not yet been separated

1 Report on Icthyology, Brit. Assoc. 1845, p. 189, 191.

2 Introduction to the Conchological part of the U. States Exploring Expedition p. xii.

[34]

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as distinct genera, though peculiar in many important respects." Whether this resemblance depends on migration during the glacial epoch & subsequent modification, I can form no opinion. Considering the wide ranges of many shells, I am surprised that there is not more identity or very close representation* between the north & south. In the Bryozoa or

* C. of Good Hope [CD.]

[35a]

[top of page not transcribed by Stauffer:] common to the northern & southern [text excised] regions, not being found in the Tropics. There are a very few cases of apparently representative species on record of genera having species in the North & South, & not in the intermediate seas, but Mr Woodward after taking much trouble finds in almost every case some sources of doubt. I am surprised at this; but it may be in some degree connected with the wide range which most genera of Mollusca have. In Brozoa or

Polyzoa, Mr. Busk gives several cases3 of European corallines now inhabiting Tierra del Fuego, New Zealand, & the Cape of Good Hope, not yet found in the Tropics, but it may be objected that the intertropical seas have hardly yet been sufficiently searched. In the Ascideae, the genus Boltenia has allied species in the arctic & antarctic seas, & Prof. Huxley thinks that the genus is not Tropical; but here again from our ignorance much caution is requisite.

[35av]

3 Catalogue of Marine Polyzoa in British Museum. 1852. p. 39, 67, 70, 83, 84, 94.

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In regard to Crustacea we can refer to Prof. Dana's full & admirable memoir on their Geographical Distribution. Many species, belonging to many genera have very wide ranges, compared with most marine animals;1 & this is important for us in allowing extensive migration during the cool period. Prof. Dana states that the sub-torrid shores of Natal, Japan, & even the Sandwich islands have several identical species & several representative species not found in the intervening torrid seas; & Prof. Dana doubts, though granting the possibility of wide migration, whether these species could possibly have passed from the southern to the northern zones;2 but under a cooler climate this difficulty would be greatly lessened.

[36v]

1 Report on Crustacea: United States Exploring Expedit. by James D. Dana. At p. 1551-54, a list of 42 species are given with very wide ranges. At p. 1574, another list of 33 species common to the African coasts, Indian ocean & Pacific. Some few species p. 1585 are common even to the East & West coasts of America.

2 Id. p. 1584: at p. 1574, where a list of 12 species in common to Natal & Japan is given.

[text excised] Id. p. 1552, 1567., 1586.—

[37A]

In Prof. Dana's work and in that of Milne Edwards4 I observe that the genera Cancer, Atelecyclus, Lithodes, Jaera & Anonyx have species on the west coast of S. America in the temperate & colder zones, both to the north & south, but none in the intermediate hotter latitudes.

4 Histoire Naturelle des Crustaces. Tome III p. 588.

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On the west coast of America, Prof. Dana3 states that the Californian sub temperate province has a close resemblance in some of its genera to the subtemperate province of Chile, though separated by 3700 miles of warmer seas; but it does not appear that any of the species are in common.

[deleted passage not transcribed by Stauffer:] & separated by the warmer seas. Mr Milne Edwards, *(b) [illeg] gives several cases of genera having not know to have to been species [illeg] to inhabit the Tropics, which have species inhabiting  in on the shores of Chile & northern hemisphere of N. America or Europe.  & N. [illeg]    Again    () 1579 gives [erased pencil words illeg]

The case of New Zealand again is similar for Prof. Dana5 shows that there is a clear relationship between its Crustacea and those of the northern hemisphere. A Palemon6 is almost identical with a British species: Cancer is not elsewhere known out of the temperate zones of N. & S. America & of Europe. The species of Portunus "are representatives of the most characteristic of European genera, & they belong rather to the cold temperate than sub-temperate regions of the Australian & New Zealand seas," Well does Prof. Dana remark that "it is certainly a wonderful fact that New Zealand should have a closer resemblance in its Crustacea to Great Britain, its antipode, than to any other part of the world," [p. 1587].

[37v]

3 Report on Crustacea, p. 1557, 1561.

4 Histoire Naturelle des Crustaces. Tome III p. 588.

5 Report on Crustacea, p. 1578, 1587.

6 Histoire Naturelle des Crustaces Tome II. p. 391.

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(Finally I may add a most striking case on the authority of Dr. Hooker namely that 25 of the same species of algae or seaweeds, belonging to 20 genera, inhabit the shores of New Zealand & Europe, & have not been found in the intermediate tropical ocean.)

[38a]

[passage not transcribed by Stauffer:] amongst [text excised] through how in this & [text excised]  to extend that [illeg] of modification within the glacial period is very doubtful.   we shall nevertheless see

(In the theory now propounded of the cold of the Glacial epoch having affected at the same time the whole world, or at least broad meridional belts, during which period northern species, both terrestrial & aquatic, crossed the Tropics, (the terrestrial stopping where higher land allowed of their permanent existence);— these species in many cases when thrown amongst foreign associates having become modified, we encounter some serious difficulties. Besides special difficulties, such as how the northern plants got into New Zealand,— why sea-shells do not offer better evidence of migration from north to south &c, we encounter some difficulties of a more general nature. The theory supposes that certain species have migrated over an immense space, during a period considered short by Geologists & sometimes falsely spoken of as mere intercalated fraction of time; but no

(31 39

geologist who has examined the glacial phenomena for himself will doubt that the period measured by years has been enormous. Nor should we forget that by the very theory all tropical productions would be in a somewhat distressed condition, & therefore would not oppose so bold a front, as before or subsequently, to the intrusion of strangers; & we know in the case of naturalised plants how widely some few have spread even in the course of a few hundred years.

Those naturalists who believe in the modification of species, but attribute much to the direct action of external conditions or who believe that there is some law determining all species to change cotemporaneously, will object that the whole body of Tropical productions ought to have become changed; but I believe that this view is erroneous, & that there would be but little tendency to change as long as the great body of tropical productions co-existed; whatever modification there may have been, would have chiefly resulted from the altered proportions of the old forms & the intrusion of strangers, new places being thus made in the polity of nature, which would be better occupied by slight selected changes of structure. So it would be with the northern & southern temperate productions during their advance & retreat in mass from the poles towards the equator. Very different would

[40A]

been far longer. Hence perhaps it is a greater difficulty that several of the northern species which have reached the southern zones, should still remain identically the same, than that they should have not been modified during their migration across the Tropics.)

(I must here observe that in several of the cases

(32 (40

it be with many or most of those forms which either crossed the Tropics & gained the temperate regions on the other side, or remained on the mountain heights within the Tropics, for they would have been associated from a more or less early part of the glacial epoch with new animal & vegetable productions. Undoubtedly it is surprising according to the theory we are here discussing, that any temperate forms should have slowly crossed the Tropics, associating all the time with productions of most different natures & exposed to very different conditions, & yet have retained the same identical character. But during these long journeys variability might have ensued, without any new permanent modification having been selected, adapting the wanderers to the not very permanent conditions which they must have encountered during these migrations. Immensely long as was the Glacial epoch, we know not in the least, whether the subsequent period during which the temperate forms have lived with their new associates may not have been far longer than the glacial epoch itself And we can distinctly understand on the theory of selection how simple time plays a most important part in the modification of specific forms.

(I must here observe that in several of the cases

(32 41

in which we have representative forms in the north & south, it by no means follows that all towards either pole have been modified since the Glacial epoch. A genus may formerly have extended, as many now do, from north to south, & have had species at both extremes, & since have become extinct in the equatorial zones, from causes independent of climate. It is also possible that one or two species of a northern genus might during the glacial epoch have migrated southward across the Tropics, & have left from subsequent extinction no individuals of the same species in the north; & in both these cases, we should falsely be led to attribute to modification during or since the glacial epoch, that which was due to migration & extinction, or to modification at a period no ways connected with the glacial epoch.

There is one other & curious difficulty to

(33 42

the foregoing theory of migration during a late cooler period. Dr. Hooker1 has remarked how singular it is that in America, whilst many northern forms have penetrated to the south, no southern forms can be said to have migrated northwards: M. A. DeCandolle1 has made the same remark

[deleted passage not transcribed by Stauffer:] likewise a very striking case with the in regard to Australia; are the identical same species or indeed any species of Eucalyptus or Banksia in the north would be indeed a prodigy!

[text excised] case of southern

 

[42v]

1 Flora of New Zealand. Introduction p. xxv. note

 1 Geographie Botanique p.  

* Dictionaire des Sciences Nat. Art. Geograph. Bot. p. 413. In Abyssinia

[42a]

in regard to Australia; & indeed the same species or any species of Eucalyptus or Banksia in the north would be a prodigy! But we have a most curious exception to this remark in the recent discovery on a mountain of Borneo, at the height of 8000 feet, of "three of the most peculiar Antarctic, New Zealand & Tasmanian genera",2 associated with Indian, & with Australian, forms, such as the heath-like Epacridae. On the mountains of Java,3 two Australian temperate genera, have been found, namely Leucopogon & Thelymitra; & it would appear from Dr. Hooker's4 observation that some few other Australian genera have travelled up the Malay peninsula, & two or three have even spread over India:* some of these genera, as the above named Leucopogon & Lagenophora, I believe are confined to the southern temperate zones. In Africa, also, there seems to be some faint indication of migration from the south to the north, as well as from the north, southwards: I allude to the two Mediterranean species of the great Cape genus of Mesembryanthemum & the one species

* Stylidium a capital case [J.D.H.]

[42av]

2 Flora of New Zealand p. xxxvi. I observe that one of the three genera, mentioned by Dr. Hooker, Drimys, was found by Mr. Gardner in the Organ Mountains of Brazil, where it, likewise seems to be a wanderer from the south or from the Cordillera.— [Hooker addendum: 'is found all the way to Mexico. See Fl. Antarct. II. sub Drimys.']

3 F. Junghuhn, Java seine Gestalt &c. 1852. Vol I. p. 417.

4 Flora Indica Introduction p. 103, 253.

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of Ixia, compared by the elder Decandolle5 to soldiers driven from their regiments.†

Notwithstanding these partial exceptions, there seems to be no doubt, that many more species & forms have passed from north to south than in the opposite direction. In attempting to explain this singular fact, we should not forget that in the northern temperate hemisphere, there is much more land than in the south, & that the plants inhabiting it are wider rangers than the more isolated species inhabiting the smaller areas in the southern hemisphere;6 & therefore that there would be a better chance for some of the northern species than for the southern species being great wanderers (enabled to cross the Tropics when slightly cooled). To hazard a conjecture unsupported by any facts, I may remark, that if the cold of the glacial epoch first came on north of the equator, the northern forms would first penetrate the Tropics, & any southern species subsequently intruding would be opposed by the great body of tropical productions with the gaps already occupied by northern forms.

† The Pelargonium & Stapelia in Levant & Algiers, & various other cases. [J.DH.]

[43a]

I believe that all the few southern temperate forms occurring on high land near the equator are specifically different from their southern congeners,* in this respect differing from the several species of northern genera found in the south; it may be & probably is accidental, but this fact harmonises with the view so often referred to that the more complete the association with foreigners the greater the probability of specific modifications through selection.)

* not so with various Cordillera species found in Fuegia [J.D.H.]

[43V]

5 Dictionaire des Sciences Nat. Art. Geograph. Bot. p. 413. In Abyssinia, also, Cape forms are found, but the intermediate country is very little known; (see C. J. F. Bunbury's Residence at the Cape of Good Hope. p. 218. also Hooker's Flora Antarctica p. 210) and the southern forms here mingle, as on the mountains of Borneo and Java, with northern forms.

[43V]

6 A. Decandolle (Geograph. Bot. p. [ ].) gives a curious comparison of the greater range of the species of the same Families in the Russian Empire & at the Cape of Good Hope.

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(I have reserved to the last some cases of distribution, the most extraordinary under our present point of view, as yet known, & which have been fully given by Dr. Hooker in his admirable Flora of the southern ocean; & I am greatly indebted to my friend for having endeavoured to make me appreciate the full force of the several difficulties. Kerguelen's Land is inhabited by only 18 phanerogamic plants; of these three are fresh-water plants found almost everywhere, & have been alluded to as most wonderful cases in an early part of this chapter. Two plants are distinct genera, known no where else; they baffle all inquiry, but do not immediately concern our immediate inquiry, Of the remaining 13 plants, 7 are endemic or aborigines, but one of them is too close to a Fuegian species; & five out of the seven genera to which these seven plants belong are genera found in but not confined to Fuegia. The remaining 8 plants are common to Fuegia, but three of them are likewise found in the New Zealand group of islands. Therefore, as remarked by Dr. Hooker, Kerguelen's Land has a much stronger Botanical affinity to Tierra del Fuego than to any other region. But these two points, measured along the parallel of 50° S. Latitude, are separated by no less than about 5000 miles of open ocean; & Kerguelen

(45

situated between Amierca & Africa in Lat 37°, about 700 miles nearer the equator than Kerguelen Land: it is inhabited, as I am informed by Dr. Hooker, by about 33 plants; of these some are not perfectly known to Dr. Hooker; from 7 to 16 are endemic; 12 are common to S. America & of the twelve, six are, in the Southern hemisphere, not found elsewhere. Hence Tristan d'Acunha, like Kerguelen Land, is botanically more nearly related to Fuegia (from which it is almost 2300 miles distant) than to any other country; & this is the more remarkable as it is only about 1700 miles distant from the southern point of Africa, to which it is related by only one or perhaps two forms, & differs in the most striking manner. Lastly, seven of the 33 plants are common to several of the antarctic islands & to the mountains, as I am informed by Dr. Hooker, of New Zealand, Tasmania and South America; so that if derived from those countries their introduction into these several countries & islands probably dates from the glacial epoch.

(45

Land is situated very much nearer to the southern points of Africa & Australia, between which it lies intermediate.

The island of Tristan d'Acunha, is inhabited by 29 phanerogamic plants, some of which are Fuegian species.

Fuego : it is situation between America & Africa, to which latter continent it shows in two of its plants some slight affinity : it lies in Lat 37° about 700 miles nearer to equator than Kergulen land, & is distant from Tierra del Fuego, about 2300 miles.

Dr. Hooker accounts for the close connexion of the floras of these distant points by supposing that within the existence of living species, there was once nearly continuous land. For reasons already given,* it seems to me that those who are inclined to believe in multiple creations, might object to the admission of such enormous changes of land & ocean without the concurrence of the weightiest evidence, both geological, zoological & botanical.

* in previous chapter [CD.]

[45v]

Some account of the Island of Tristan d'Acunha by Capt. D. Carmichael. Linnean Transactions vol XII p. 483.

(38 (46)

[passage not transcribed by Stauffer:] modification of specific forms be allowed, fomr the southern extremity of America. But these two points are separated by measure along to parallel 50° parallel of Latitude by about 5000 miles of open ocean

(46)

Sir Charles Lyell1 has suggested that plants may have been widely disseminated in the antarctic ocean by the agency of ice-bergs. There seems to me much probability in this view, especially if we bear in mind the prodigious number of great blocks of rock, which have been transported from both sides of the Cordillera from its southern extremity up to Latitude 42°.2 The most obvious objection to this view is that the icebergs must have travelled a vast distance in nearly the same latitude; & in the northern hemisphere we know from the scratches on the rocks that the course of the icebergs was formerly, as now, approximately north & south. But the great difference between the northern hemisphere & the quite open ocean of the southern hemisphere must not be overlooked. We have, also, the following fact as a guide: a bottle was thrown overboard by Sir James Ross3 a little northward and eastward

1 [Principles, 9th ed., p. 622.]

3 As stated before the Geographical Society, June 22, 1846. [See The Athenaeum (1846), 656.]

[46v]

2 Geological Transactions Vol VI. C. Darwin on the Distribution of the erratic Boulders of S. America.

(39 (47)

of Cape Horn, and was picked up at Cape Liptrap, the extreme southern point of Australia north of Tasmania;

(With respect to icebergs occasionally carrying seeds, I think it would be quite extraordinary if they did not do so, just in the same way as seeds are carried in the ballast of ships & plants thus naturalised: we should remember the innumerable great fragments of rock which certainly have thus been carried many hundred miles. I have had the particulars given me of two icebergs in the antarctic ocean with great fragments of rock, at least 1200 miles from the nearest known land. Besides stones, "loads of earth", brushwood, live animals of several kinds, the skull of the musk ox which was landed in Greenland, the bones of the Lemming, & even the nest of a bird with its eggs2 have all been observed on icebergs. Can we doubt that seeds of plants, with their vitality well preserved, might likewise be thus carried? Dr. Rae has suggested to me that the gales of winter, which sweep the ground bare of snow, can hardly fail to blow seeds on the fissured glaciers near the coast: stray birds resting on icebergs might occasionally leave,hard seeds of fruit in their droppings: where there are rivers the autumnal frosts would freeze mud & seeds together, & such river-ice,

1 Horsburgh. Philosophical Transactions 1830 p. 117

2 For these latter facts see Crantz, History of Greenland. Vol I. p. 26. Supplement to Parry's voyage by Capt. Sabine p. cxc. Also Richardson's British Assoc. Report for 1836. p. 163. Scoresby estimated the weight of "the beds of earth & rock" on many of the icebergs near Spitzbergen at from 50,000 to 100,000 tons; (Lyell's Principles of Geology. 9th Edit. p. 227.)

[47Vv]

[not transcribed by Stauffer:] (a) text) The extraordinary prevalence of westerly gales in the latitudes must not be forgotten

The course of icebergs would probably be determinedto a considerable extent

[47V]

The extraordinary prevalence of violent westerly gales, comparable in regularity with the trade-winds, in these latitudes must not be forgotten. But the course of the ice-bergs would be very much determined by their depth, & their reaching the underlying stream of cold water flowing to the equator. At the present day icebergs have been observed within a degree of the Cape of Good Hope,1 & could hardly have travelled, bearing in mind the westerly winds, less than 3000 miles.

1 Horsburgh. Philosophical Transactions 1830 p. 117

[47V]

this bottle, during its voyage Eastward of about 9000 miles had gained only about 900 miles northing.

[47V]

* 2 For these latter facts see Crantz, History of Greenland. Vol I. p. 26. Supplement to Parry's voyage by Capt. Sabine p. cxc. Also Richardson's British Assoc. Report for 1836. p. 163. Scoresby estimated the weight of "the beds of earth & rock" on many of the icebergs near Spitzbergen at from 50,000 to 100,000 tons; (Lyell's Principles of Geology. 9th Edit. p. 227.)

[an erased pencil note mentions Kerguelen Land]

[48A]

(I should infer from Capt. Carmichael's account that there were proportionally fewer species in Tristan d'Acunha identically the same with those of Fuegia, than in Kerguelen Land. If this be so these islands present a parallel case to the mountains of Scotland & the Alps of Switzerland compared with the Arctic regions; the cause being, I should suppose the same, namely, the points nearer the equator having been colonized, and the colonists isolated, at an earlier part of the Glacial epoch.)

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as I am informed by Dr. Rae, is sometimes 6 or 8 feet thick, & when in the sea gets packed and crushed together. When an iceberg is stranded, great masses of ice, by the unanimous testimony of Arctic travellers, are pushed up high and dry by the pressure of the pack outside; & Dr. Rae assures me he has seen hundreds of instances of ice driven so high on land, that when it thawed any enclosed seeds would have had a good chance of growing. Seeds in earth, even if discharged in the sea on a shallow coast, would have a chance of being thrown up, like shells from deep water. We must never forget during how many hundred-thousand years this action must have gone on during the glacial period; & that during this period the native plants of many southern coasts would have been distressed & could not have resisted the intrusion of more vigorous southern strangers. Hence I can see no insuperable difficulty in the seeds of Fuegian plants having been carried to Kerguelen Land1 & to Tristan d'Acunha during some part of the glacial period.

[48v]

1 It perhaps deserves notice that the Modiolarca trapezina is common to the Falkland Islands & Kerguelen Land (see Woodward's excellent Supplement to Treatise on Shells p. 371, 378) & as this shell is often attached to the masses of the gigantic Macrocystis, this seaweed is probably drifted from the one place to the other.—

[Hooker added: "We pulled up immense masses with shells & stones attached, fin all parts of Ant. Sea beyond the very icy regions J.H."]

[49A]

namely 13 common to Fuegia & New Zealand (of which 6 likewise occur in the small Aukland & Campbell islands lying between 200 & 300 miles south of New Zealand)

[49A]

Dr Hooker states (+ 2 Flora of New Zealand, Introduction, p. xxxi)

that 89 plants are common to New Zealand & S.America; several of these are very wide ranging species & offer no more special difficulty than in other parallel cases: some few may have travelled from the north during the glacial period, and so got into these two distant southern points of the world. But of the 89, Dr. Hooker has given me a list of about 25 species which are absolutely confined to the southern temperate zone, & yet are identical to these two widely separated points. The interspace of ocean measured along the parallel of 45°, is about 4500 miles, without one single island now existing as a resting place.† From the same species in common

† The Fuschias & Calceolarieas are as great difficulties under your view of modification. [J.D.H.]

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[page not transcribed by Stauffer]

(New Zealand offers in some respects a still more difficult case. Dr. Hooker

 

 

& more especially from the representative species in common Dr Hooker infers (Introduct p. XXIII) that both Chile & Fuegia were formerly connected with New Zealand by intermediate land, but not necessarily continuous at any one time at any one time over the whole distance. The above mentioned 25 species in common may be divided into two classes,

 

 

The remaining 12 are common to Chile

(50

& New Zealand. Of these twelve, it can only be said that they belong (with the very remarkable exception of a Myosurus) to genera having species in many parts of the world; but Dr. Hooker informs me that the species in these genera are neither particularly wide rangers nor particularly restricted.

[deleted passage not transcribed by Stauffer:] With respect to the species in common to Fuego & New Zealand, *a [illeg] possibility of transport during the glacial epoch by iceberg, should not be overlooked.

With respect to the representative species I shall have to allude in the next chapter, from those southern lands now

In the next chapter when remarking on these very interesting representative species,2 I shall have occasion to allude to the possibility of those southern islands which are now wholly covered by ice, having been clothed with vegetation before the commencement of the glacial epoch; & the seeds of some plants now in common to New Zealand, S. America & the other Antarctic islands, as Kerguelen Land &c, may have been carried by icebergs at an early part of the glacial epoch, from a common southern home. & the species not have been subsequently modified. The advocates of multiple creations, may, in my opinion, bring forward the species more especially those found in Chile & New Zealand as a very strong case in favour of their view:

[50v]

that seeds might have been formerly carried from Tierra de Fuego to New Zealand; though the voyage at the rate of 25 miles a day, would take a year!  

[50V]

2 There are, according to Dr. Hooker, several identical & representative species on the heights of New Zealand & Tasmania. The possibility of great icebergs having-been formed during the glacial epoch on the Eastern side of Tasmania, & having thus carried seeds should not be forgotten, for in the very same latitude on the shores of S. America immense blocks of rock (one of granite 15 feet by 11, & 9 in height) have been carried about 40 miles from the Cordillera (not very lofty in this part) to the island of Chiloe.— The case of the bottle carried from near Cape Horn to the southern point of Australia should not be overlooked; for it seems just possible, though very improbable that seeds might have been formerly carried from Tierra de Fuego to New Zealand; though the voyage at the rate of 25 miles a day, would take a year! [Hooker wrote: 'Why strike this out? The berg would perhaps travel faster with winds J.H.']

(51

but it should not be overlooked that they would find it very difficult to give any rational explanation of the community of these few species, for the great mass of organic productions, & all the external conditions are widely different in Chile & New Zealand; & it might well be asked, why should these few plants be identical & so vast a number of other productions widely different: it seems to me safer to rely on our ignorance of the means of diffusion.—

In regard to our general conclusion on the great amount of migration during the glacial epoch, of which epoch we have in many of the areas in question independent & decisive geological evidence, I think it has much probability, notwithstanding the many cases of difficulty enumerated, some special, some general & others probably overlooked. This same view, may I believe be extended to some cases, which have not been here noticed from want of space. It explains in my opinion many anomalies in distribution, & removes some few of the greatest difficulties in admitting, in accordance with

(52

the strong presumption derived from general laws, that each organic form was created or produced in one area. Moreover it strengthens the theory, in as much as it explains to a certain extent several facts otherwise inexplicable, that species under certain given conditions undergo modification. There is much interest in looking at the alpine productions of mountains in the most distant quarters of the world as monuments of not very high antiquity, yet often written in a changed dialect, recording the nature of the organic beings which once, when the world was cooler, surrounded their bases, & there perished. We have on these monuments the evidence of a great tide of life which slowly flowed from either pole towards the equator,— the waters, it may be said, breaking more freely over from the north than from the south. The two great tidal waves then slowly ebbed towards the Poles, but have not yet reached, & perhaps will never reach, their first & native source.

(53

(To sum up this chapter, already much too long, we commenced with showing that many general facts or laws indicate that each species has appeared at one point or rather area of the earth's surface; each species not being necessarily derived from a single pair, but by the very slow modification, through selection, of many individuals of another species. The supposed creation of the same species at more than one point of the earth's surface is admitted, even by those who hold this belief, to be an exceptional & even paradoxical case; yet it must be owned that such exceptional cases are not rare, & often present inexplicable, but not in my opinion overwhelming difficulties.

We see clear indications of a law of single creation, & we cannot honestly deny that we are profoundly ignorant of the many possible means of diffusion, past & present. Who denies that the weather is due to regular laws, yet who can go into detail & say why the sun shone yesterday, or the rain falls today? The cases of the greatest difficulty are mostly included in the three

[54A]

may formerly have had under different conditions a more continuous range & become extinct in the intermediate regions, & secondly that some species have retained the same identical forms since even the commencement of the Miocene period, & this allows time for prodigious geographical changes. Hence I conclude that it has not as yet been absolutely proved that the same species has ever appeared, independently of migration,

(54

great classes discussed in this chapter, namely in the floras & faunas of oceanic islands,— of fresh-water lakes or rivers, — & of mountain-summits with the polar regions; & I have collected together such explanations as have been given by others or have occurred to myself To have collected the several isolated cases, would have been less serviceable & most tedious: yet some such are very curious & quite inexplicable. For instance the presence of Myrsine africana at the Cape of Good Hope, Abyssinia & the Azores & not as far as is known in any intermediate point. Besides our ignorance of the means of dispersion, & the chances of naturalisation by man's agency at some unknown time, we should never forget that a species

(55

on two separate points of the earth's surface: if this were proved or rendered highly probable, the whole of this volume would be useless,* & we should be compelled to admit the truth of the common view of absolute actual creation; & that organic beings are not exclusively produced by ordinary generation, with or without modification.)

* No No— whether or no do not say so— it is not to the purpose. J.H.

[B55v]

[page not transcribed by Stauffer]

(55

on two separate points of the earths surface. But if this could be proved, or rendered probable, then the theory that new species are derived from the modification of other species might be considered as disproved or rendered improbably in the highest degree; for if the production creation of an organic being out of the dead elements in of such nature that it may though to be so little mis[illeg], that is proved in one instance, the case of a being which might in another quarter of the world have been  
which if this were proved or rendered highly probable then [words illeg] to about the [illeg] of the common view that organic beings are absolutely created or that each [illeg] & a generation with [words illeg]
For the question we are [illeg] in the colour the [words illeg]

Nor as much as   It seems so extremely inprobable that the same identical forms shd ever have arisen by ordinary generation & modification from two distinct species at two distinct points, that such a notion may be disregarded, & therefore if it could be proved or by as most as it is rendered probable, that the same species now occur on 2 points of the earths surface, without the possibility of it having migrated from one to the other, then we shd have to admit that [illeg] creation of organic beings

[C0]

(41)

[fair copy, in the hand of Ebenezer Norman]

As I believe that all organic beings are produced by the ordinary laws of reproduction which includes, according to the theory under discussion, modification of specific forms, & as it is exceedingly improbable that the same species should ever have been generated in one place from one set of parents, & in another place, (especially if under different conditions) from another set of parents specifically dissimilar, the first & most obvious question is whether we can account on the ordinary notion of propagation for the existence of the same identical species in all quarters of the world.— This is the question, which has long agitated naturalists, namely whether the same species has been created once & therefore at a single point, or more than once at different points.

After giving general reasons in favour of single production; consider the many & grave difficulties. The most prominent of them may be grouped into three following classes.— First, insular productions of the temperate & tropical latitudes.†— Here give reasons for doubting vast continental extensions of Forbes & Co:  Give condensed means of dispersal.—

Secondly, range of Fresh Water Productions.—

Thirdly, as follows.—

† Perhaps allude to A. Decandolle on large-scaled Plants. [C.D.]

[C1]

1

(This will come towards end of another chapter)

We will now consider some of the most striking cases of difficulty on geological grounds opposed to the theory that closely allied or representative species, are due to the modification of the same species. Dr. Hooker has given a most curious list1 of representative species, found in New Zealand, Australia & S. America. With respect to Australia there is no greater difficulty (but great enough) than in other analogous cases, already discussed when considering the identical species of insular floras & faunas.* But with respect to S. America the case is different, owing to the vast space of open ocean between that continent & New Zealand. Yet even here we find an accordance with the general rule that the productions of an island are more or less allied to those of the land nearest to it; & again we see an accordance with the rule that where there are representative species there are some identical species in common, proving to those who believe in single creations, that there has been at some time some channel of communication between the two areas in question. Dr. Hooker believes that there was formerly a communication

 

* Find out whether any old Rocks in New Zealand; also about soundings, so to give chance of former union. [C.D.]

[1V]

1 New Zealand Flora. Introduction, p. xxxiv.

2

by more or less continuous land from both Chile & Fuegia to New Zealand. I cannot persuade myself to admit (though far better judges see no difficulty in admitting) such great geographical changes within so recent a period; & I think that a slight modification of Dr. Hooker's view will remove some little of the difficulty.

Of the 50 genera which afford the best instances of representative species in New Zealand & extratropical S. America, 7, as I am informed by Dr. Hooker, are northern genera, & 18 of very general distribution; & the representative species in these 25 genera (bearing in mind the glacial epoch) present nothing more remarkable than representative species in other parts of the world, to which in former times we may imagine the descendants of the same species to have travelled & subsequently to have become modified: but the other 25 genera are strictly confined to the south with all their species extratropical,— a few on the mountains within the Tropics being excepted. This fact of half the genera being confined to the South seems to me remarkable considering that out of the 89 species belonging to 76 genera1 absolutely identical in New Zealand & S. America, only two species belong to genera confined to the South, namely Colobanthus subulatus & Rostkovia Magellanica: Goodenia repens          need hardly be added as this is an Australian genus with one littoral wandering species. Again it may be noticed in Dr. Hooker's list (which I am

3

aware is not given as perfect) that of the southern genera, which have representatives in New Zealand & S. America, there are five which have none in Australia or Tasmania, & this is what might have been expected considering the greater distance of Australia, than of New Zealand from S. America; but Australia has four southern genera (viz; Eucryphia, Pernettya, Lebetanthus, & Lomatia) with representatives of S. American species, which genera do not occur in New Zealand or the Auckland islands. These facts, together perhaps with the genera Colobanthus, Ac[a]ena & Lagenophora having representative species, both on these two lands & in several of the circumpolar islands, seem to me to indicate some common centre of radiation.1

Now taking the northern hemisphere as our guide, I should look to the circumpolar regions as the centre of radiation for the representative & for some of the species still remaining identical in the above named several lands. If we look to a chart we see in the little explored regions between 62° & 80° several islands, & large tracts of land, with surroundings in one place 100 miles from the shore, & with indications2 of other rocks besides volcanic. Here then I should infer that it was no ways improbable that these lands & islands may have recently been of greater extent & more continuous. On these islands not one single land plant can now live, but bearing in mind that in the north the space between these same parallels is the home of the whole Arctic Flora, it seems to

[3V]

1 Memoirs of Geological Survey Vol I, p. 399 &c.

2 Cybele Britannica, Vol. p. 37.

[3V]

2 Sir J. Ross Voyage to S. Seas. Vol. 2, p. 421.

4

me a not very improbable supposition that before the glacial epoch came on, these islands might have been covered by a not scanty vegetation.3 According to all analogy, this antarctic vegetation from its isolation would have been very peculiar, but would have been in some degree related to that of the two nearest continents, America and Australia; & this antarctic vegetation though perhaps not nearly so uniform, as that now growing on the almost continuous arctic land, would probably have been4  tolerably uniform. From this source, I am inclined to suppose that some plants, either identical or allied, migrated before the glacial epoch by various accidental means, aided probably by more continuous land & by the several intermediate islands which we see still existing South of New Zealand; & that when the glacial epoch did come on, the seeds of other plants were brought in a N. Easterly course by icebergs from their common home, soon to be converted into an icy desert. The plants which arrived at this latter period, would, on the returning warmth have ascended the mountains of New Zealand & Tasmania, — most of the species especially those brought first, having subsequently undergone modification & now existing as representative species,— a few having remained identical. But those who receive the common view that every species has been created as we now see it, & that the same species has sometimes been created at more than one point of the earth's surface, may truly say with derision what complicated

[4V]

3 In the southern hemisphere we have no distinct evidence that the climate was warmer during the older pliocene periods. The existence of burnt & silicified trees & thin beds of coal under the streams of lava in Kerguelen's Land & in [New South] Shetland Isld. (V. Dana's letter on Mr. Eights [Amer. J. Sci. 2nd ser., 22 (1856), 391]) where the vegetation is now so scanty should not be forgotten. I may, also, state that in Tierra del Fuego, (Geological Observation, p. 118) I found in beds underlying the drift or glacial deposits, many leaves of trees; which belong, according to Dr. Hooker to three species of Beech, apparently differing from the two species, which now clothe that forest-clad land.

5

theories are required, such as the one just given, or the more simple one but requiring much greater geographical changes given by Dr. Hooker, to account on the theory of descent for the same, & with subsequent modification, for the representative species in these two distant areas. On the other hand those who believe in simple creation can, in my opinion, give no explanation in the least degree satisfactory of the shades of affinity & degree of identity in the cases which we have been discussing: they in fact simply state so it is.—

CUL-DAR14.D1r: Ms p 1

[fair copy by Ebenezer Norman of CUL-DAR14.B1-B55, above, with corrections by Darwin. Stauffer gives the annotations by Hooker and Darwin in the transcription of the original draft above]

CUL-DAR14.D1Vv: Inserts * & † to Ms p 1   

CUL-DAR14.D2r: Ms p 2      

CUL-DAR14.D3r: Ms p 3      

CUL-DAR14.D4r: Ms p 4      

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CUL-DAR14.D6V[2]v: [Note by Darwin]      

Dr. Oswald Heer (Band XV der neuen Denkschriften des allgem. Schweizerischen Gesellschaft) boldly supposes that land stretched continuously from West Europe to East America, sending promontories to Iceland in the north & to the Canary Islands in the south & that this land endured till the end of the Tertiary period, which would include the glacial epoch. To those who can freely admit such enormous geographical changes on no other evidence besides distribution, this view will satisfactorily ??? account for many of the phenomena.

(Give argument against continuous land between Europe & N. America from the proportion of plants common to Europe & U. States, not greatly exceeding those in common between Asia & U. States Oct. 10. 56. See A. Gray in Silliman)

CUL-DAR14.D7r: Ms p 7      

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CUL-DAR14.D13r: Ms p 13      

CUL-DAR14.D13v: [Note by Darwin]      

Refer to Asa Grays most striking tables

CUL-DAR14.D13Vv: Inserts * & † to Ms p 13      

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CUL-DAR14.D16r: Ms p 16      

CUL-DAR14.D16v: [Note by Darwin]      

Also peculiar Land-shells nearly or quite the same in mountains of Ceylon & Nilghiri

CUL-DAR14.D16Vv: Inserts * , † , ‡ , & ⊙ to Ms p 16      

CUL-DAR14.D17r: Ms p 17      

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