Darwinism (1889) Part 28

Murray more nearly indicate the limit, but the very small portions of matter brought up by the dredge, as compared with the enormous areas of sea-bottom, over which the atmospheric dust must have been scattered, render it in the highest degree improbable that the maximum limit either of size of particles, or of distance from land has been reached.

Let us, however, a.s.sume that the quartz grains, found by Mr. Murray in the deep-sea ooze 700 miles from land, give us the extreme limit of the power of the atmosphere as a carrier of solid particles, and let us compare with these the weights of some seeds. From a small collection of the seeds of thirty species of herbaceous plants sent me from Kew, those in the above table were selected, and small portions of eight of them carefully weighed in a chemical balance.[175] By counting these portions I was able to estimate the number of seeds weighing one grain. The three very minute species, whose numbers are marked with an asterisk (*), were estimated by the comparison of their sizes with those of the smaller weighed seeds.

No| Species. |Approximate | Approximate | Remarks.

| |No. of Seeds| Dimensions. | | |In one Grain| | | | | in. in. in. | 1|Draba verna | 1,800 |1/60 x 1/90 x 1/160|Oval, flat.

2|Hyperic.u.m perforatum | 520 | 1/30 x 1/80 |Cylindrical.

3|Astilbe rivularis | 4,500 | 1/50 x 1/100 |Elongate, flat, tailed, | | | | wavy.

4|Saxifraga coriophylla| 750 | 1/40 x 1/75 |Surface rough, adhere | | | | to the dry capsules.

5|Oenothera rosea | 640 | 1/40 x 1/80 |Ovate.

6|Hyperic.u.m hirsutum | 700 | 1/30 x 1/100 |Cylindrical, rough.

7|Mimulus luteus | 2,900 | 1/60 x 1/100 |Oval, minute.

8|Penthorum sedoides | 8,000* | 1/70 x 1/150 |Flattened, very minute.

9|Sagina proc.u.mbens | 12,000* | 1/120 |Sub-triangular, flat.

10|Orchis maculata | 15,000* | --- |Margined, flat, | | | | very minute.

11|Gentiana purpurea | 35 | 1/25 |Wavy, rough, with this | | | | coriaceous margins.

12|Silene alpina | --- | 1/30 |Flat, with fringed | | | | margins.

13|Adenophora communis | --- | 1/20 x 1/40 |Very thin, wavy, light.

|Quartz grains | 25,000 | 1/250 |Deep sea ... 700 miles.

|Do. |200,000 | 1/500 |Genoa ... 600 miles.

If now we compare the seeds with the quartz grains, we find that several are from twice to three times the weight of the grains found by Mr. Murray, and others five times, eight times, and fifteen times as heavy; but they are proportionately very much larger, and, being usually irregular in shape or compressed, they expose a very much larger surface to the air. The surface is often rough, and several have dilated margins or tailed appendages, increasing friction and rendering the uniform rate of falling through still air immensely less than in the case of the smooth, rounded, solid quartz grains. With these advantages it is a moderate estimate that seeds ten times the weight of the quartz grains could be carried quite as far through the air by a violent gale and under the most favourable conditions. These limits will include five of the seeds here given, as well as hundreds of others which do not exceed them in weight; and to these we may add some larger seeds which have other favourable characteristics, as is the case with numbers 11-13, which, though very much larger than the rest, are so formed as in all probability to be still more easily carried great distances by a gale of wind. It appears, therefore, to be absolutely certain that every autumnal gale capable of conveying solid mineral particles to great distances, must also carry numbers of small seeds at least as far; and if this is so, the wind alone will form one of the most effective agents in the dispersal of plants.

Hitherto this mode of conveyance, as applying to the transmission of seeds for great distances across the ocean, has been rejected by botanists, for two reasons. In the first place, there is said to be no direct evidence of such conveyance; and, secondly, the peculiar plants of remote oceanic islands do not appear to have seeds specially adapted for aerial transmission. I will consider briefly each of these objections.

_Objection to the Theory of Wind-Dispersal._

To obtain direct evidence of the transmission of such minute and perishable objects, which do not exist in great quant.i.ties, and are probably carried to the greatest distances but rarely and as single specimens, is extremely difficult. A bird or insect can be seen if it comes on board s.h.i.+p, but who would ever detect the seeds of Mimulus or Orchis even if a score of them fell on a s.h.i.+p's deck? Yet if but one such seed per century were carried to an oceanic island, that island might become rapidly overrun by the plant, if the conditions were favourable to its growth and reproduction. It is further objected that search has been made for such seeds, and they have not been found.

Professor Kerner of Innsbruck examined the snow on the surface of glaciers, and a.s.siduously collected all the seeds he could find, and these were all of plants which grew in the adjacent mountains or in the same district. In like manner, the plants growing on moraines were found to be those of the adjacent mountains, plateaux, or lowlands. Hence he concluded that the prevalent opinion that seeds may be carried through the air for very great distances "is not supported by fact."[176] The opinion is certainly not supported by Kerner's facts, but neither is it opposed by them. It is obvious that the seeds that would be carried by the wind to moraines or to the surface of glaciers would be, first and in the greatest abundance, those of the immediately surrounding district; then, very much more rarely, those from more remote mountains; and lastly, in extreme rarity, those from distant countries or altogether distinct mountain ranges. Let us suppose the first to be so abundant that a single seed could be found by industrious search on each square yard of the surface of the glacier; the second so scarce that only one could possibly be found in a hundred yards square; while to find one of the third cla.s.s it would be necessary exhaustively to examine a square mile of surface. Should we expect that _one_ ever to be found, and should the fact that it could not be found be taken as a proof that it was not there? Besides, a glacier is altogether in a bad position to receive such remote wanderers, since it is generally surrounded by lofty mountains, often range behind range, which would intercept the few air-borne seeds that might have been carried from a distant land. The conditions in an oceanic island, on the other hand, are the most favourable, since the land, especially if high, will intercept objects carried by the wind, and will thus cause more of the solid matter to fall on it than on an equal area of ocean. We know that winds at sea often blow violently for days together, and the rate of motion is indicated by the fact that 72 miles an hour was the average velocity of the wind observed during twelve hours at the Ben Nevis observatory, while the velocity sometimes rises to 120 miles an hour. A twelve hours' gale might, therefore, carry light seeds a thousand miles as easily and certainly as it could carry quartz-grains of much greater specific gravity, rotundity, and smoothness, 500 or even 100 miles; and it is difficult even to imagine a sufficient reason why they should not be so carried--perhaps very rarely and under exceptionally favourable conditions,--but this is all that is required.

As regards the second objection, it has been observed that orchideae, which have often exceedingly small and light seeds, are remarkably absent from oceanic islands. This, however, may be very largely due to their extreme specialisation and dependence on insect agency for their fertilisation; while the fact that they do occur in such very remote islands as the Azores, Tahiti, and the Sandwich Islands, proves that they must have once reached these localities either by the agency of birds or by transmission through the air; and the facts I have given above render the latter mode at least as probable as the former. Sir Joseph Hooker remarks on the composite plant of Kerguelen Island (Cotula plumosa) being found also on Lord Auckland and MacQuarrie Islands, and yet having no pappus, while other species of the genus possess it. This is certainly remarkable, and proves that the plant must have, or once have had, some other means of dispersal across wide oceans.[177] One of the most widely dispersed species in the whole world (Sonchus oleraceus) possesses pappus, as do four out of five of the species which are common to Europe and New Zealand, all of which have a very wide distribution.

The same author remarks on the limited area occupied by most species of Compositae, notwithstanding their facilities for dispersal by means of their feathered seeds; but it has been already shown that limitations of area are almost always due to the compet.i.tion of allied forms, facilities for dispersal being only one of many factors in determining the wide range of species. It is, however, a specially important factor in the case of the inhabitants of remote oceanic islands, since, whether they are peculiar species or not, they or their remote ancestors must at some time or other have reached their present position by natural means.

I have already shown elsewhere, that the flora of the Azores strikingly supports the view of the species having been introduced by aerial transmission only, that is, by the agency of birds and the wind, because all plants that could not possibly have been carried by these means are absent.[178] In the same way we may account for the extreme rarity of Leguminosae in all oceanic islands. Mr. Hemsley, in his Report on Insular Floras, says that they "are wanting in a large number of oceanic islands where there is no true littoral flora," as St. Helena, Juan Fernandez, and all the islands of the South Atlantic and South Indian Oceans. Even in the tropical islands, such as Mauritius and Bourbon, there are no endemic species, and very few in the Galapagos and the remoter Pacific Islands. All these facts are quite in accordance with the absence of facilities for transmission through the air, either by birds or the wind, owing to the comparatively large size and weight of the seeds; and an additional proof is thus afforded of the extreme rarity of the successful floating of seeds for great distances across the ocean.[179]

_Explanation of North Temperate Plants in the Southern Hemisphere._

If we now admit that many seeds which are either minute in size, of thin texture or wavy form, or so fringed or margined as to afford a good hold to the air, are capable of being carried for many hundreds of miles by exceptionally violent and long-continued gales of wind, we shall not only be better able to account for the floras of some of the remotest oceanic islands, but shall also find in the fact a sufficient explanation of the wide diffusion of many genera, and even species, of arctic and north temperate plants in the southern hemisphere or on the summits of tropical mountains. Nearly fifty of the flowering plants of Tierra-del-Fuego are found also in North America or Europe, but in no intermediate country; while fifty-eight species are common to New Zealand and Northern Europe; thirty-eight to Australia, Northern Europe, and Asia; and no less than seventy-seven common to New Zealand, Australia, and South America.[180] On lofty mountains far removed from each other, identical or closely allied plants often occur. Thus the fine Primula imperialis of a single mountain peak in Java has been found (or a closely allied species) in the Himalayas; and many other plants of the high mountains of Java, Ceylon, and North India are either identical or closely allied forms. So, in Africa, some species, found on the summits of the Cameroons and Fernando Po in West Africa, are closely allied to species in the Abyssinian highlands and in Temperate Europe; while other Abyssinian and Cameroons species have recently been found on the mountains of Madagascar. Some peculiar Australian forms have been found represented on the summit of Kini Balu in Borneo. Again, on the summit of the Organ mountains in Brazil there are species allied to those of the Andes, but not found in the intervening lowlands.

_No Proof of Recent Lower Temperature in the Tropics._

Now all these facts, and numerous others of like character, were supposed by Mr. Darwin to be due to a lowering of temperature during glacial epochs, which allowed these temperate forms to migrate across the intervening tropical lowlands. But any such change within the epoch of existing species is almost inconceivable. In the first place, it would necessitate the extinction of much of the tropical flora (and with it of the insect life), because without such extinction alpine herbaceous plants could certainly never spread over tropical forest lowlands; and, in the next place, there is not a particle of direct evidence that any such lowering of temperature in inter-tropical lowlands ever took place. The only alleged evidence of the kind is that adduced by the late Professor Aga.s.siz and Mr. Hartt; but I am informed by my friend, Mr. J.C. Branner (now State Geologist of Arkansas, U.S.), who succeeded Mr. Hartt, and spent several years completing the geological survey of Brazil, that the supposed moraines and glaciated granite rocks near Rio Janeiro and elsewhere, as well as the so-called boulder-clay of the same region, are entirely explicable as the results of sub-aerial denudation and weathering, and that there is no proof whatever of glaciation in any part of Brazil.

_Lower Temperature not needed to Explain the Facts._

But any such vast physical change as that suggested by Darwin, involving as it does such tremendous issues as regards its effects on the tropical fauna and flora of the whole world, is really quite uncalled for, because the facts to be explained are of the same essential nature as those presented by remote oceanic islands, between which and the nearest continents no temperate land connection is postulated. In proportion to their limited area and extreme isolation, the Azores, St. Helena, the Galapagos, and the Sandwich Islands, each possess a fairly rich--the last a very rich--indigenous flora; and the means which sufficed to stock them with a great variety of plants would probably suffice to transmit others from mountain-top to mountain-top in various parts of the globe. In the case of the Azores, we have large numbers of species identical with those of Europe, and others closely allied, forming an exactly parallel case to the species found on the various mountain summits which have been referred to. The distances from Madagascar to the South African mountains and to Kilimandjaro, and from the latter to Abyssinia, are no greater than from Spain to the Azores, while there are other equatorial mountains forming stepping-stones at about an equal distance to the Cameroons. Between Java and the Himalayas we have the lofty mountains of Sumatra and of North-western Burma, forming steps at about the same distance apart; while between Kini Balu and the Australian Alps we have the unexplored snow mountains of New Guinea, the b.e.l.l.e.n.den Ker mountains in Queensland, and the New England and Blue Mountains of New South Wales. Between Brazil and Bolivia the distances are no greater; while the unbroken range of mountains from Arctic America to Tierra-del-Fuego offers the greatest facilities for transmission, the partial gap between the lofty peak of Chiriqui and the high Andes of New Grenada being far less than from Spain to the Azores.

Thus, whatever means have sufficed for stocking oceanic islands must have been to some extent effective in transmitting northern forms from mountain to mountain, across the equator, to the southern hemisphere; while for this latter form of dispersal there are special facilities, in the abundance of fresh and unoccupied surfaces always occurring in mountain regions, owing to avalanches, torrents, mountain-slides, and rock-falls, thus affording stations on which air-borne seeds may germinate and find a temporary home till driven out by the inroads of the indigenous vegetation. These temporary stations may be at much lower alt.i.tudes than the original habitat of the species, if other conditions are favourable. Alpine plants often descend into the valleys on glacial moraines, while some arctic species grow equally well on mountain summits and on the seash.o.r.e. The distances above referred to between the loftier mountains may thus be greatly reduced by the occurrence of suitable conditions at lower alt.i.tudes, and the facilities for transmission by means of aerial currents proportionally increased.[181]

_Facts Explained by the Wind-Carriage of Seeds._

But if we altogether reject aerial transmission of seeds for great distances, except by the agency of birds, it will be difficult, if not impossible, to account for the presence of so many identical species of plants on remote mountain summits, or for that "continuous current of vegetation" described by Sir Joseph Hooker as having apparently long existed from the northern to the southern hemisphere. It may be admitted that we can, possibly, account for the greater portion of the floras of remote oceanic islands by the agency of birds alone; because, when blown out to sea land-birds must reach some island or perish, and all which come within sight of an island will struggle to reach it as their only refuge. But, with mountain summits the case is altogether different, because, being surrounded by land instead of by sea, no bird would need to fly, or to be carried by the wind, for several hundred miles at a stretch to another mountain summit, but would find a refuge in the surrounding uplands, ridges, valleys, or plains. As a rule the birds that frequent lofty mountain tops are peculiar species, allied to those of the surrounding district; and there is no indication whatever of the pa.s.sage of birds from one remote mountain to another in any way comparable with the flights of birds which are known to reach the Azores annually, or even with the few regular migrants from Australia to New Zealand. It is almost impossible to conceive that the seeds of the Himalayan primula should have been thus carried to Java; but, by means of gales of wind, and intermediate stations from fifty to a few hundred miles apart, where the seeds might vegetate for a year or two and produce fresh seed to be again carried on in the same manner, the transmission might, after many failures, be at last effected.

A very important consideration is the vastly larger scale on which wind-carriage of seeds must act, as compared with bird-carriage. It can only be a few birds which carry seeds attached to their feathers or feet. A very small proportion of these would carry the seeds of Alpine plants; while an almost infinitesimal fraction of these latter would convey the few seeds attached to them safely to an oceanic island or remote mountain. But winds, in the form of whirlwinds or tornadoes, gales or hurricanes, are perpetually at work over large areas of land and sea. Insects and light particles of matter are often carried up to the tops of high mountains; and, from the very nature and origin of winds, they usually consist of ascending or descending currents, the former capable of suspending such small and light objects as are many seeds long enough for them to be carried enormous distances. For each single seed carried away by external attachment to the feet or feathers of a bird, countless millions are probably carried away by violent winds; and the chance of conveyance to a great distance and in a definite direction must be many times greater by the latter mode than by the former.[182] We have seen that inorganic particles of much greater specific gravity than seeds, and nearly as heavy as the smallest kinds, are carried to great distances through the air, and we can therefore hardly doubt that some seeds are carried as far. The direct agency of the wind, as a supplement to bird-transport, will help to explain the presence in oceanic islands of plants growing in dry or rocky places whose small seeds are not likely to become attached to birds; while it seems to be the only effective agency possible in the dispersal of those species of alpine or sub-alpine plants found on the summits of distant mountains, or still more widely separated in the temperate zones of the northern and southern hemispheres.

_Concluding Remarks._

On the general principles that have been now laid down, it will be found that all the chief facts of the geographical distribution of animals and plants can be sufficiently understood. There will, of course, be many cases of difficulty and some seeming anomalies, but these can usually be seen to depend on our ignorance of some of the essential factors of the problem. Either we do not know the distribution of the group in recent geological times, or we are still ignorant of the special methods by which the organisms are able to cross the sea. The latter difficulty applies especially to the lizard tribe, which are found in almost all the tropical oceanic islands; but the particular mode in which they are able to traverse a wide expanse of ocean, which is a perfect barrier to batrachia and almost so to snakes, has not yet been discovered. Lizards are found in all the larger Pacific Islands as far as Tahiti, while snakes do not extend beyond the Fiji Islands; and the latter are also absent from Mauritius and Bourbon, where lizards of seven or eight species abound. Naturalists resident in the Pacific Islands would make a valuable contribution to our science by studying the life-history of the native lizards, and endeavouring to ascertain the special facilities they possess for crossing over wide s.p.a.ces of ocean.

FOOTNOTES:

[Footnote 163: See A. Aga.s.siz, _Three Cruises of the Blake_ (Cambridge, Ma.s.s., 1888), vol. i. p. 127, footnote.]

[Footnote 164: Even the extremely fine Mississippi mud is nowhere found beyond a hundred miles from the mouths of the river in the Gulf of Mexico (A. Aga.s.siz, _Three Cruises of the Blake_, vol. i. p. 128).]

[Footnote 165: I have given a full summary of the evidence for the permanence of oceanic and continental areas in my _Island Life_, chap.

vi.]

[Footnote 166: For a full account of the peculiarities of the Madagascar fauna, see my _Island Life_, chap. xix.]

[Footnote 167: See _Island Life_, p. 446, and the whole of chaps. xxi.

xxii. More recent soundings have shown that the Map at p. 443, as well as that of the Madagascar group at p. 387, are erroneous, the ocean around Norfolk Island and in the Straits of Mozambique being more than 1000 fathoms deep. The general argument is, however, unaffected.]

[Footnote 168: For some details of these migrations, see the author's _Geographical Distribution of Animals_, vol. i. p. 140; also Heilprin's _Geographical and Geological Distribution of Animals_.]

[Footnote 169: For a full discussion of this question, see _Island Life_, pp. 390-420.]

[Footnote 170: _Geographie Botanique_, p. 798.]

[Footnote 171: _Nature_, 1st April 1886.]

[Footnote 172: Report of the Brit. a.s.soc. Committee on Migration of Birds during 1886.]

[Footnote 173: _Trans. Ent. Soc._, 1871, p. 184.]

[Footnote 174: _Nature_ (1875), vol. xii. pp. 279, 298.]

[Footnote 175: I am indebted to Professor R. Meldola of the Finsbury Technical Inst.i.tute, and Rev. T.D. t.i.tmas of Charterhouse for furnis.h.i.+ng me with the weights required.]

[Footnote 176: See _Nature_, vol. vi. p. 164, for a summary of Kerner's paper.]

[Footnote 177: It seems quite possible that the absence of pappus in this case is a recent adaptation, and that it has been brought about by causes similar to those which have reduced or aborted the wings of insects in oceanic islands. For when a plant has once reached one of the storm-swept islands of the southern ocean, the pappus will be injurious for the same reason that the wings of insects are injurious, since it will lead to the seeds being blown out to sea and destroyed. The seeds which are heaviest and have least pappus will have the best chance of falling on the ground and remaining there to germinate, and this process of selection might rapidly lead to the entire disappearance of the pappus.]