I have now read your paper, and I hope that you will not think me presumptuous in writing another line to say how excellent it seems to me. I believe that you have largely solved the problem of the affinities of the inhabitants of this most interesting little island, and this is a delightful triumph.
LETTER 395. TO J.D. HOOKER. Down, July 22nd [1879].
I have just read Ball"s Essay. (395/1. The late John Ball"s lecture "On the Origin of the Flora of the Alps" in the "Proceedings of the R.
Geogr. Soc." 1879. Ball argues (page 18) that "during ancient Palaeozoic times, before the deposition of the Coal-measures, the atmosphere contained twenty times as much carbonic acid gas and considerably less oxygen than it does at present." He further a.s.sumes that in such an atmosphere the percentage of CO2 in the higher mountains would be excessively different from that at the sea-level, and appends the result of calculations which gives the amount of CO2 at the sea-level as 100 per 10,000 by weight, at a height of 10,000 feet as 12.5 per 10,000.
Darwin understands him to mean that the Vascular Cryptogams and Gymnosperms could stand the sea-level atmosphere, whereas the Angiosperms would only be able to exist in the higher regions where the percentage of CO2 was small. It is not clear to us that Ball relies so largely on the condition of the atmosphere as regards CO2. If he does he is clearly in error, for everything we know of a.s.similation points to the conclusion that 100 per 10,000 (1 per cent.) is by no means a hurtful amount of CO2, and that it would lead to an especially vigorous a.s.similation. Mountain plants would be more likely to descend to the plains to share in the rich feast than ascend to higher regions to avoid it. Ball draws attention to the imperfection of our plant records as regards the floras of mountain regions. It is, he thinks, conceivable that there existed a vegetation on the Carboniferous mountains of which no traces have been preserved in the rocks. See "Fossil Plants as Tests of Climate," page 40, A.C. Seward, 1892.
Since the first part of this note was written, a paper has been read (May 29th, 1902) by Dr. H.T. Brown and Mr. F. Es...o...b.., before the Royal Society on "The Influence of varying amounts of Carbon Dioxide in the Air on the Photosynthetic Process of Leaves, and on the Mode of Growth of Plants." The author"s experiments included the cultivation of several dicotyledonous plants in an atmosphere containing in one case 180 to 200 times the normal amount of CO2, and in another between three and four times the normal amount. The general results were practically identical in the two sets of experiments. "All the species of flowering plants, which have been the subject of experiment, appear to be accurately "tuned" to an atmospheric environment of three parts of CO2 per 10,000, and the response which they make to slight increases in this amount are in a direction altogether unfavourable to their growth and reproduction." The a.s.similation of carbon increases with the increase in the partial pressure of the CO2. But there seems to be a disturbance in metabolism, and the plants fail to take advantage of the increased supply of CO2. The authors say:--"All we are justified in concluding is, that if such atmospheric variations have occurred since the advent of flowering plants, they must have taken place so slowly as never to outrun the possible adaptation of the plants to their changing conditions."
Prof. Farmer and Mr. S.E. Chandler gave an account, at the same meeting of the Royal Society, of their work "On the Influence of an Excess of Carbon Dioxide in the Air on the Form and Internal Structure of Plants."
The results obtained were described as differing in a remarkable way from those previously recorded by Teodoresco ("Rev. Gen. Botanique,"
II., 1899
It is hoped that Dr. Horace Brown and Mr. Es...o...b.. will extend their experiments to Vascular Cryptogams, and thus obtain evidence bearing more directly upon the question of an increased amount of CO2 in the atmosphere of the Coal-period forests.) It is pretty bold. The rapid development as far as we can judge of all the higher plants within recent geological times is an abominable mystery. Certainly it would be a great step if we could believe that the higher plants at first could live only at a high level; but until it is experimentally [proved] that Cycadeae, ferns, etc., can withstand much more carbonic acid than the higher plants, the hypothesis seems to me far too rash. Saporta believes that there was an astonishingly rapid development of the high plants, as soon [as] flower-frequenting insects were developed and favoured intercrossing. I should like to see this whole problem solved. I have fancied that perhaps there was during long ages a small isolated continent in the S. Hemisphere which served as the birthplace of the higher plants--but this is a wretchedly poor conjecture. It is odd that Ball does not allude to the obvious fact that there must have been alpine plants before the Glacial period, many of which would have returned to the mountains after the Glacial period, when the climate again became warm. I always accounted to myself in this manner for the gentians, etc.
Ball ought also to have considered the alpine insects common to the Arctic regions. I do not know how it may be with you, but my faith in the glacial migration is not at all shaken.
LETTER 396. A.R. WALLACE TO CHARLES DARWIN.
(396/1. This letter is in reply to Mr. Darwin"s criticisms on Mr.
Wallace"s "Island Life," 1880.)
Pen-y-Bryn, St. Peter"s Road, Croydon, November 8th, 1880.
Many thanks for your kind remarks and notes on my book. Several of the latter will be of use to me if I have to prepare a second edition, which I am not so sure of as you seem to be.
1. In your remark as to the doubtfulness of paucity of fossils being due to coldness of water, I think you overlook that I am speaking only of water in the lat.i.tude of the Alps, in Miocene and Eocene times, when icebergs and glaciers temporarily descended into an otherwise warm sea; my theory being that there was no Glacial epoch at that time, but merely a local and temporary descent of the snow-line and glaciers owing to high excentricity and winter in aphelion.
2. I cannot see the difficulty about the cessation of the Glacial period.
Between the Miocene and the Pleistocene periods geographical changes occurred which rendered a true Glacial period possible with high excentricity. When the high excentricity pa.s.sed away the Glacial epoch also pa.s.sed away in the temperate zone; but it persists in the arctic zone, where, during the Miocene, there were mild climates, and this is due to the persistence of the changed geographical conditions. The present arctic climate is itself a comparatively new and abnormal state of things, due to geographical modification.
As to "epoch" and "period," I use them as synonyms to avoid repeating the same word.
3. Rate of deposition and geological time. Here no doubt I may have gone to an extreme, but my "28 million years" may be anything under 100 millions, as I state. There is an enormous difference between mean and maximum denudation and deposition. In the case of the great faults the upheaval along a given line would itself facilitate the denudation (whether sub-aerial or marine) of the upheaved portion at a rate perhaps a hundred times above the average, just as valleys have been denuded perhaps a hundred times faster than plains and plateaux. So local subsidence might itself lead to very rapid deposition. Suppose a portion of the Gulf of Mexico, near the mouths of the Mississippi, were to subside for a few thousand years, it might receive the greater portion of the sediment from the whole Mississippi valley, and thus form strata at a very rapid rate.
4. You quote the Pampas thistles, etc., against my statement of the importance of preoccupation. But I am referring especially to St.
Helena, and to plants naturally introduced from the adjacent continents.
Surely if a certain number of African plants reached the island, and became modified into a complete adaptation to its climatic conditions, they would hardly be expelled by other African plants arriving subsequently. They might be so, conceivably, but it does not seem probable. The cases of the Pampas, New Zealand, Tahiti, etc., are very different, where highly developed aggressive plants have been artificially introduced. Under nature it is these very aggressive species that would first reach any island in their vicinity, and, being adapted to the island and colonising it thoroughly, would then hold their own against other plants from the same country, mostly less aggressive in character.
I have not explained this so fully as I should have done in the book.
Your criticism is therefore useful.
5. My Chapter XXIII. is no doubt very speculative, and I cannot wonder at your hesitating at accepting my views. To me, however, your theory of hosts of existing species migrating over the tropical lowlands from the N. temperate to the S. temperate zone appears more speculative and more improbable. For where could the rich lowland equatorial flora have existed during a period of general refrigeration sufficient for this?
and what became of the wonderfully rich Cape flora, which, if the temperature of tropical Africa had been so recently lowered, would certainly have spread northwards, and on the return of the heat could hardly have been driven back into the sharply defined and very restricted area in which it now exists.
As to the migration of plants from mountain to mountain not being so probable as to remote islands, I think that is fully counterbalanced by two considerations:--
a. The area and abundance of the mountain stations along such a range as the Andes are immensely greater than those of the islands in the N.
Atlantic, for example.
b. The temporary occupation of mountain stations by migrating plants (which I think I have shown to be probable) renders time a much more important element in increasing the number and variety of the plants so dispersed than in the case of islands, where the flora soon acquires a fixed and endemic character, and where the number of species is necessarily limited.
No doubt direct evidence of seeds being carried great distances through the air is wanted, but I am afraid can hardly be obtained. Yet I feel the greatest confidence that they are so carried. Take, for instance, the two peculiar orchids of the Azores (Habenaria sp.) What other mode of transit is conceivable? The whole subject is one of great difficulty, but I hope my chapter may call attention to a hitherto neglected factor in the distribution of plants.
Your references to the Mauritius literature are very interesting, and will be useful to me; and I again thank you for your valuable remarks.
LETTER 397. TO J.D. HOOKER.
(397/1. The following letters were written to Sir J.D. Hooker when he was preparing his Address as President of the Geographical Section of the British a.s.sociation at its fiftieth meeting, at York. The second letter (August 12th) refers to an earlier letter of August 6th, published in "Life and Letters," III., page 246.)
4, Bryanston Street, W., Sat.u.r.day, 26th [February, 1881].
I should think that you might make a very interesting address on Geographical Distribution. Could you give a little history of the subject. I, for one, should like to read such history in petto; but I can see one very great difficulty--that you yourself ought to figure most prominently in it; and this you would not do, for you are just the man to treat yourself in a dishonourable manner. I should very much like to see you discuss some of Wallace"s views, especially his ignoring the all-powerful effects of the Glacial period with respect to alpine plants. (397/2. "Having been kindly permitted by Mr. Francis Darwin to read this letter, I wish to explain that the above statement applies only to my rejection of Darwin"s view that the presence of arctic and north temperate plants in the SOUTHERN HEMISPHERE was brought about by the lowering of the temperature of the tropical regions during the Glacial period, so that even "the lowlands of these great continents were everywhere tenanted under the equator by a considerable number of temperate forms ("Origin of Species," Edition VI., page 338). My own views are fully explained in Chapter XXIII. of my "Island Life,"
published in 1880. I quite accept all that Darwin, Hooker, and Asa Gray have written about the effect of the Glacial epoch in bringing about the present distribution of alpine and arctic plants in the NORTHERN HEMISPHERE."--Note by Mr. Wallace.) I do not know what you think, but it appears to me that he exaggerates enormously the influence of debacles or slips and new surface of soil being exposed for the reception of wind-blown seeds. What kinds of seeds have the plants which are common to the distant mountain-summits in Africa? Wallace lately wrote to me about the mountain plants of Madagascar being the same with those on mountains in Africa, and seemed to think it proved dispersal by the wind, without apparently having inquired what sorts of seeds the plants bore. (397/3. The affinity with the flora of the Eastern African islands was long ago pointed out by Sir J.D. Hooker, "Linn. Soc. Journal," VI., 1861, page 3. Speaking of the plants of Clarence Peak in Fernando Po, he says, "The next affinity is with Mauritius, Bourbon, and Madagascar: of the whole 76 species, 16 inhabit these places and 8 more are closely allied to plants from there. Three temperate species are peculiar to Clarence Peak and the East African islands..." The facts to which Mr. Wallace called Darwin"s attention are given by Mr. J.G. Baker in "Nature," December 9th, 1880, page 125. He mentions the Madagascar Viola, which occurs elsewhere only at 7,000 feet in the Cameroons, at 10,000 feet in Fernando Po and in the Abyssinian mountains; and the same thing is true of the Madagascar Geranium. In Mr. Wallace"s letter to Darwin, dated January 1st, 1881, he evidently uses the expression "pa.s.sing through the air" in contradistinction to the migration of a species by gradual extension of its area on land. "Through the air"
would moreover include occasional modes of transport other than simple carriage by wind: e.g., the seeds might be carried by birds, either attached to the feathers or to the mud on their feet, or in their crops or intestines.)
I suppose it would be travelling too far (though for the geographical section the discussion ought to be far-reaching), but I should like to see the European or northern element in the Cape of Good Hope flora discussed. I cannot swallow Wallace"s view that European plants travelled down the Andes, tenanted the hypothetical Antarctic continent (in which I quite believe), and thence spread to South Australia and the Cape of Good Hope.
Moseley told me not long ago that he proposed to search at Kerguelen Land the coal beds most carefully, and was absolutely forbidden to do so by Sir W. Thomson, who said that he would undertake the work, and he never once visited them. This puts me in a pa.s.sion. I hope that you will keep to your intention and make an address on distribution. Though I differ so much from Wallace, his "Island Life" seems to me a wonderful book.
Farewell. I do hope that you may have a most prosperous journey. Give my kindest remembrances to Asa Gray.
LETTER 398. TO J.D. HOOKER. Down, August 12th, 1881.
...I think that I must have expressed myself badly about Humboldt.
I should have said that he was more remarkable for his astounding knowledge than for originality. I have always looked at him as, in fact, the founder of the geographical distribution of organisms. I thought that I had read that extinct fossil plants belonging to Australian forms had lately been found in Australia, and all such cases seem to me very interesting, as bearing on development.
I have been so astonished at the apparently sudden coming in of the higher phanerogams, that I have sometimes fancied that development might have slowly gone on for an immense period in some isolated continent or large island, perhaps near the South Pole. I poured out my idle thoughts in writing, as if I had been talking with you.
No fact has so interested me for a heap of years as your case of the plants on the equatorial mountains of Africa; and Wallace tells me that some one (Baker?) has described a.n.a.logous cases on the mountains of Madagascar (398/1. See Letter 397, note.)...I think that you ought to allude to these cases.
I most fully agree that no problem is more interesting than that of the temperate forms in the southern hemisphere, common to the north. I remember writing about this after Wallace"s book appeared, and hoping that you would take it up. The frequency with which the drainage from the land pa.s.ses through mountain-chains seems to indicate some general law--viz., the successive formation of cracks and lines of elevation between the nearest ocean and the already upraised land; but that is too big a subject for a note.
I doubt whether any insects can be shown with any probability to have been flower feeders before the middle of the Secondary period. Several of the a.s.serted cases have broken down.
Your long letter has stirred many pleasant memories of long past days, when we had many a discussion and many a good fight.
LETTER 399. TO J.D. HOOKER. Down, August 21st, 1881.
I cannot aid you much, or at all. I should think that no one could have thought on the modification of species without thinking of representative species. But I feel sure that no discussion of any importance had been published on this subject before the "Origin,"
for if I had known of it I should a.s.suredly have alluded to it in the "Origin," as I wished to gain support from all quarters. I did not then know of Von Buch"s view (alluded to in my Historical Introduction in all the later editions). Von Buch published his "Isles Canaries" in 1836, and he here briefly argues that plants spread over a continent and vary, and the varieties in time come to be species. He also argues that closely allied species have been thus formed in the SEPARATE valleys of the Canary Islands, but not on the upper and open parts. I could lend you Von Buch"s book, if you like. I have just consulted the pa.s.sage.