Part 14 (1/2)

Sir C. Lyell and other authors have ably treated this subject. I can give here only the briefest abstract of the more important facts. Change of climate must have had a powerful influence on migration: a region when its climate was different may have been a high road for migration, but now be impa.s.sable; I shall, however, presently have to discuss this branch of the subject in some detail. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two marine faunas; submerge it, or let it formerly have been submerged, and the two faunas will now blend or may formerly have blended: where the sea now extends, land may at a former period have connected islands or possibly even continents together, and thus have allowed terrestrial productions to pa.s.s from one to the other. No geologist will dispute that great mutations of level have occurred within the period of existing organisms. Edward Forbes insisted that all the islands in the Atlantic must recently have been connected with Europe or Africa, and Europe likewise with America. Other authors have thus hypothetically bridged over every ocean, and have united almost every island to some mainland. If indeed the arguments used by Forbes are to be trusted, it must be admitted that scarcely a single island exists which has not recently been united to some continent. This view cuts the Gordian knot of the dispersal of the same species to the most distant points, and removes many a difficulty: but to the best of my judgment we are not authorized in admitting such enormous geographical changes within the period of existing species. It seems to me that we have abundant evidence of great oscillations of level in our continents; but not of such vast changes in their position and extension, as to have united them within the recent period to each other and to the several intervening oceanic islands. I freely admit the former existence of many islands, now buried beneath the sea, which may have served as halting places for plants and for many animals during their migration. In the coral-producing oceans such sunken islands are now marked, as I believe, by rings of coral or atolls standing over them. Whenever it is fully admitted, as I believe it will some day be, that each species has proceeded from a single birthplace, and when in the course of time we know something definite about the means of distribution, we shall be enabled to speculate with security on the former extension of the land.

But I do not believe that it will ever be proved that within the recent period continents which are now quite separate, have been continuously, or almost continuously, united with each other, and with the many existing oceanic islands. Several facts in distribution,--such as the great difference in the marine faunas on the opposite sides of almost every continent,--the close relation of the tertiary inhabitants of several lands and even seas to their present inhabitants,--a certain degree of relation (as we shall hereafter see) between the distribution of mammals and the depth of the sea,--these and other such facts seem to me opposed to the admission of such prodigious geographical revolutions within the recent period, as are necessitated on the view advanced by Forbes and admitted by his many followers. The nature and relative proportions of the inhabitants of oceanic islands likewise seem to me opposed to the belief of their former continuity with continents. Nor does their almost universally volcanic composition favour the admission that they are the wrecks of sunken continents;--if they had originally existed as mountain-ranges on the land, some at least of the islands would have been formed, like other mountain-summits, of granite, metamorphic schists, old fossiliferous or other such rocks, instead of consisting of mere piles of volcanic matter.

I must now say a few words on what are called accidental means, but which more properly might be called occasional means of distribution.

I shall here confine myself to plants. In botanical works, this or that plant is stated to be ill adapted for wide dissemination; but for transport across the sea, the greater or less facilities may be said to be almost wholly unknown. Until I tried, with Mr. Berkeley's aid, a few experiments, it was not even known how far seeds could resist the injurious action of sea-water. To my surprise I found that out of 87 kinds, 64 germinated after an immersion of 28 days, and a few survived an immersion of 137 days. For convenience sake I chiefly tried small seeds, without the capsule or fruit; and as all of these sank in a few days, they could not be floated across wide s.p.a.ces of the sea, whether or not they were injured by the salt-water. Afterwards I tried some larger fruits, capsules, etc., and some of these floated for a long time. It is well known what a difference there is in the buoyancy of green and seasoned timber; and it occurred to me that floods might wash down plants or branches, and that these might be dried on the banks, and then by a fresh rise in the stream be washed into the sea. Hence I was led to dry stems and branches of 94 plants with ripe fruit, and to place them on sea water. The majority sank quickly, but some which whilst green floated for a very short time, when dried floated much longer; for instance, ripe hazel-nuts sank immediately, but when dried, they floated for 90 days and afterwards when planted they germinated; an asparagus plant with ripe berries floated for 23 days, when dried it floated for 85 days, and the seeds afterwards germinated: the ripe seeds of Helosciadium sank in two days, when dried they floated for above 90 days, and afterwards germinated. Altogether out of the 94 dried plants, 18 floated for above 28 days, and some of the 18 floated for a very much longer period. So that as 64/87 seeds germinated after an immersion of 28 days; and as 18/94 plants with ripe fruit (but not all the same species as in the foregoing experiment) floated, after being dried, for above 28 days, as far as we may infer anything from these scanty facts, we may conclude that the seeds of 14/100 plants of any country might be floated by sea-currents during 28 days, and would retain their power of germination. In Johnston's Physical Atlas, the average rate of the several Atlantic currents is 33 miles per diem (some currents running at the rate of 60 miles per diem); on this average, the seeds of 14/100 plants belonging to one country might be floated across 924 miles of sea to another country; and when stranded, if blown to a favourable spot by an inland gale, they would germinate.

Subsequently to my experiments, M. Martens tried similar ones, but in a much better manner, for he placed the seeds in a box in the actual sea, so that they were alternately wet and exposed to the air like really floating plants. He tried 98 seeds, mostly different from mine; but he chose many large fruits and likewise seeds from plants which live near the sea; and this would have favoured the average length of their flotation and of their resistance to the injurious action of the salt-water. On the other hand he did not previously dry the plants or branches with the fruit; and this, as we have seen, would have caused some of them to have floated much longer. The result was that 18/98 of his seeds floated for 42 days, and were then capable of germination. But I do not doubt that plants exposed to the waves would float for a less time than those protected from violent movement as in our experiments.

Therefore it would perhaps be safer to a.s.sume that the seeds of about 10/100 plants of a flora, after having been dried, could be floated across a s.p.a.ce of sea 900 miles in width, and would then germinate.

The fact of the larger fruits often floating longer than the small, is interesting; as plants with large seeds or fruit could hardly be transported by any other means; and Alph. de Candolle has shown that such plants generally have restricted ranges.

But seeds may be occasionally transported in another manner. Drift timber is thrown up on most islands, even on those in the midst of the widest oceans; and the natives of the coral-islands in the Pacific, procure stones for their tools, solely from the roots of drifted trees, these stones being a valuable royal tax. I find on examination, that when irregularly shaped stones are embedded in the roots of trees, small parcels of earth are very frequently enclosed in their interstices and behind them,--so perfectly that not a particle could be washed away in the longest transport: out of one small portion of earth thus COMPLETELY enclosed by wood in an oak about 50 years old, three dicotyledonous plants germinated: I am certain of the accuracy of this observation.

Again, I can show that the carca.s.ses of birds, when floating on the sea, sometimes escape being immediately devoured; and seeds of many kinds in the crops of floating birds long retain their vitality: peas and vetches, for instance, are killed by even a few days' immersion in sea-water; but some taken out of the crop of a pigeon, which had floated on artificial salt-water for 30 days, to my surprise nearly all germinated.

Living birds can hardly fail to be highly effective agents in the transportation of seeds. I could give many facts showing how frequently birds of many kinds are blown by gales to vast distances across the ocean. We may I think safely a.s.sume that under such circ.u.mstances their rate of flight would often be 35 miles an hour; and some authors have given a far higher estimate. I have never seen an instance of nutritious seeds pa.s.sing through the intestines of a bird; but hard seeds of fruit will pa.s.s uninjured through even the digestive organs of a turkey. In the course of two months, I picked up in my garden 12 kinds of seeds, out of the excrement of small birds, and these seemed perfect, and some of them, which I tried, germinated. But the following fact is more important: the crops of birds do not secrete gastric juice, and do not in the least injure, as I know by trial, the germination of seeds; now after a bird has found and devoured a large supply of food, it is positively a.s.serted that all the grains do not pa.s.s into the gizzard for 12 or even 18 hours. A bird in this interval might easily be blown to the distance of 500 miles, and hawks are known to look out for tired birds, and the contents of their torn crops might thus readily get scattered. Mr. Brent informs me that a friend of his had to give up flying carrier-pigeons from France to England, as the hawks on the English coast destroyed so many on their arrival. Some hawks and owls bolt their prey whole, and after an interval of from twelve to twenty hours, disgorge pellets, which, as I know from experiments made in the Zoological Gardens, include seeds capable of germination. Some seeds of the oat, wheat, millet, canary, hemp, clover, and beet germinated after having been from twelve to twenty-one hours in the stomachs of different birds of prey; and two seeds of beet grew after having been thus retained for two days and fourteen hours. Freshwater fish, I find, eat seeds of many land and water plants: fish are frequently devoured by birds, and thus the seeds might be transported from place to place. I forced many kinds of seeds into the stomachs of dead fish, and then gave their bodies to fis.h.i.+ng-eagles, storks, and pelicans; these birds after an interval of many hours, either rejected the seeds in pellets or pa.s.sed them in their excrement; and several of these seeds retained their power of germination. Certain seeds, however, were always killed by this process.

Although the beaks and feet of birds are generally quite clean, I can show that earth sometimes adheres to them: in one instance I removed twenty-two grains of dry argillaceous earth from one foot of a partridge, and in this earth there was a pebble quite as large as the seed of a vetch. Thus seeds might occasionally be transported to great distances; for many facts could be given showing that soil almost everywhere is charged with seeds. Reflect for a moment on the millions of quails which annually cross the Mediterranean; and can we doubt that the earth adhering to their feet would sometimes include a few minute seeds? But I shall presently have to recur to this subject.

As icebergs are known to be sometimes loaded with earth and stones, and have even carried brushwood, bones, and the nest of a land-bird, I can hardly doubt that they must occasionally have transported seeds from one part to another of the arctic and antarctic regions, as suggested by Lyell; and during the Glacial period from one part of the now temperate regions to another. In the Azores, from the large number of the species of plants common to Europe, in comparison with the plants of other oceanic islands nearer to the mainland, and (as remarked by Mr. H. C.

Watson) from the somewhat northern character of the flora in comparison with the lat.i.tude, I suspected that these islands had been partly stocked by ice-borne seeds, during the Glacial epoch. At my request Sir C. Lyell wrote to M. Hartung to inquire whether he had observed erratic boulders on these islands, and he answered that he had found large fragments of granite and other rocks, which do not occur in the archipelago. Hence we may safely infer that icebergs formerly landed their rocky burthens on the sh.o.r.es of these mid-ocean islands, and it is at least possible that they may have brought thither the seeds of northern plants.

Considering that the several above means of transport, and that several other means, which without doubt remain to be discovered, have been in action year after year, for centuries and tens of thousands of years, it would I think be a marvellous fact if many plants had not thus become widely transported. These means of transport are sometimes called accidental, but this is not strictly correct: the currents of the sea are not accidental, nor is the direction of prevalent gales of wind.

It should be observed that scarcely any means of transport would carry seeds for very great distances; for seeds do not retain their vitality when exposed for a great length of time to the action of seawater; nor could they be long carried in the crops or intestines of birds. These means, however, would suffice for occasional transport across tracts of sea some hundred miles in breadth, or from island to island, or from a continent to a neighbouring island, but not from one distant continent to another. The floras of distant continents would not by such means become mingled in any great degree; but would remain as distinct as we now see them to be. The currents, from their course, would never bring seeds from North America to Britain, though they might and do bring seeds from the West Indies to our western sh.o.r.es, where, if not killed by so long an immersion in salt-water, they could not endure our climate. Almost every year, one or two land-birds are blown across the whole Atlantic Ocean, from North America to the western sh.o.r.es of Ireland and England; but seeds could be transported by these wanderers only by one means, namely, in dirt sticking to their feet, which is in itself a rare accident. Even in this case, how small would the chance be of a seed falling on favourable soil, and coming to maturity! But it would be a great error to argue that because a well-stocked island, like Great Britain, has not, as far as is known (and it would be very difficult to prove this), received within the last few centuries, through occasional means of transport, immigrants from Europe or any other continent, that a poorly-stocked island, though standing more remote from the mainland, would not receive colonists by similar means.

I do not doubt that out of twenty seeds or animals transported to an island, even if far less well-stocked than Britain, scarcely more than one would be so well fitted to its new home, as to become naturalised.

But this, as it seems to me, is no valid argument against what would be effected by occasional means of transport, during the long lapse of geological time, whilst an island was being upheaved and formed, and before it had become fully stocked with inhabitants. On almost bare land, with few or no destructive insects or birds living there, nearly every seed, which chanced to arrive, would be sure to germinate and survive.

DISPERSAL DURING THE GLACIAL PERIOD.

The ident.i.ty of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where the Alpine species could not possibly exist, is one of the most striking cases known of the same species living at distant points, without the apparent possibility of their having migrated from one to the other. It is indeed a remarkable fact to see so many of the same plants living on the snowy regions of the Alps or Pyrenees, and in the extreme northern parts of Europe; but it is far more remarkable, that the plants on the White Mountains, in the United States of America, are all the same with those of Labrador, and nearly all the same, as we hear from Asa Gray, with those on the loftiest mountains of Europe. Even as long ago as 1747, such facts led Gmelin to conclude that the same species must have been independently created at several distinct points; and we might have remained in this same belief, had not Aga.s.siz and others called vivid attention to the Glacial period, which, as we shall immediately see, affords a simple explanation of these facts. We have evidence of almost every conceivable kind, organic and inorganic, that within a very recent geological period, central Europe and North America suffered under an Arctic climate. The ruins of a house burnt by fire do not tell their tale more plainly, than do the mountains of Scotland and Wales, with their scored flanks, polished surfaces, and perched boulders, of the icy streams with which their valleys were lately filled. So greatly has the climate of Europe changed, that in Northern Italy, gigantic moraines, left by old glaciers, are now clothed by the vine and maize. Throughout a large part of the United States, erratic boulders, and rocks scored by drifted icebergs and coast-ice, plainly reveal a former cold period.

The former influence of the glacial climate on the distribution of the inhabitants of Europe, as explained with remarkable clearness by Edward Forbes, is substantially as follows. But we shall follow the changes more readily, by supposing a new glacial period to come slowly on, and then pa.s.s away, as formerly occurred. As the cold came on, and as each more southern zone became fitted for arctic beings and ill-fitted for their former more temperate inhabitants, the latter would be supplanted and arctic productions would take their places. The inhabitants of the more temperate regions would at the same time travel southward, unless they were stopped by barriers, in which case they would perish. The mountains would become covered with snow and ice, and their former Alpine inhabitants would descend to the plains. By the time that the cold had reached its maximum, we should have a uniform arctic fauna and flora, covering the central parts of Europe, as far south as the Alps and Pyrenees, and even stretching into Spain. The now temperate regions of the United States would likewise be covered by arctic plants and animals, and these would be nearly the same with those of Europe; for the present circ.u.mpolar inhabitants, which we suppose to have everywhere travelled southward, are remarkably uniform round the world. We may suppose that the Glacial period came on a little earlier or later in North America than in Europe, so will the southern migration there have been a little earlier or later; but this will make no difference in the final result.

As the warmth returned, the arctic forms would retreat northward, closely followed up in their retreat by the productions of the more temperate regions. And as the snow melted from the bases of the mountains, the arctic forms would seize on the cleared and thawed ground, always ascending higher and higher, as the warmth increased, whilst their brethren were pursuing their northern journey. Hence, when the warmth had fully returned, the same arctic species, which had lately lived in a body together on the lowlands of the Old and New Worlds, would be left isolated on distant mountain-summits (having been exterminated on all lesser heights) and in the arctic regions of both hemispheres.

Thus we can understand the ident.i.ty of many plants at points so immensely remote as on the mountains of the United States and of Europe.

We can thus also understand the fact that the Alpine plants of each mountain-range are more especially related to the arctic forms living due north or nearly due north of them: for the migration as the cold came on, and the re-migration on the returning warmth, will generally have been due south and north. The Alpine plants, for example, of Scotland, as remarked by Mr. H. C. Watson, and those of the Pyrenees, as remarked by Ramond, are more especially allied to the plants of northern Scandinavia; those of the United States to Labrador; those of the mountains of Siberia to the arctic regions of that country. These views, grounded as they are on the perfectly well-ascertained occurrence of a former Glacial period, seem to me to explain in so satisfactory a manner the present distribution of the Alpine and Arctic productions of Europe and America, that when in other regions we find the same species on distant mountain-summits, we may almost conclude without other evidence, that a colder climate permitted their former migration across the low intervening tracts, since become too warm for their existence.

If the climate, since the Glacial period, has ever been in any degree warmer than at present (as some geologists in the United States believe to have been the case, chiefly from the distribution of the fossil Gnathodon), then the arctic and temperate productions will at a very late period have marched a little further north, and subsequently have retreated to their present homes; but I have met with no satisfactory evidence with respect to this intercalated slightly warmer period, since the Glacial period.

The arctic forms, during their long southern migration and re-migration northward, will have been exposed to nearly the same climate, and, as is especially to be noticed, they will have kept in a body together; consequently their mutual relations will not have been much disturbed, and, in accordance with the principles inculcated in this volume, they will not have been liable to much modification. But with our Alpine productions, left isolated from the moment of the returning warmth, first at the bases and ultimately on the summits of the mountains, the case will have been somewhat different; for it is not likely that all the same arctic species will have been left on mountain ranges distant from each other, and have survived there ever since; they will, also, in all probability have become mingled with ancient Alpine species, which must have existed on the mountains before the commencement of the Glacial epoch, and which during its coldest period will have been temporarily driven down to the plains; they will, also, have been exposed to somewhat different climatal influences. Their mutual relations will thus have been in some degree disturbed; consequently they will have been liable to modification; and this we find has been the case; for if we compare the present Alpine plants and animals of the several great European mountain-ranges, though very many of the species are identically the same, some present varieties, some are ranked as doubtful forms, and some few are distinct yet closely allied or representative species.

In ill.u.s.trating what, as I believe, actually took place during the Glacial period, I a.s.sumed that at its commencement the arctic productions were as uniform round the polar regions as they are at the present day. But the foregoing remarks on distribution apply not only to strictly arctic forms, but also to many sub-arctic and to some few northern temperate forms, for some of these are the same on the lower mountains and on the plains of North America and Europe; and it may be reasonably asked how I account for the necessary degree of uniformity of the sub-arctic and northern temperate forms round the world, at the commencement of the Glacial period. At the present day, the sub-arctic and northern temperate productions of the Old and New Worlds are separated from each other by the Atlantic Ocean and by the extreme northern part of the Pacific. During the Glacial period, when the inhabitants of the Old and New Worlds lived further southwards than at present, they must have been still more completely separated by wider s.p.a.ces of ocean. I believe the above difficulty may be surmounted by looking to still earlier changes of climate of an opposite nature.

We have good reason to believe that during the newer Pliocene period, before the Glacial epoch, and whilst the majority of the inhabitants of the world were specifically the same as now, the climate was warmer than at the present day. Hence we may suppose that the organisms now living under the climate of lat.i.tude 60 deg, during the Pliocene period lived further north under the Polar Circle, in lat.i.tude 66 deg-67 deg; and that the strictly arctic productions then lived on the broken land still nearer to the pole. Now if we look at a globe, we shall see that under the Polar Circle there is almost continuous land from western Europe, through Siberia, to eastern America. And to this continuity of the circ.u.mpolar land, and to the consequent freedom for intermigration under a more favourable climate, I attribute the necessary amount of uniformity in the sub-arctic and northern temperate productions of the Old and New Worlds, at a period anterior to the Glacial epoch.

Believing, from reasons before alluded to, that our continents have long remained in nearly the same relative position, though subjected to large, but partial oscillations of level, I am strongly inclined to extend the above view, and to infer that during some earlier and still warmer period, such as the older Pliocene period, a large number of the same plants and animals inhabited the almost continuous circ.u.mpolar land; and that these plants and animals, both in the Old and New Worlds, began slowly to migrate southwards as the climate became less warm, long before the commencement of the Glacial period. We now see, as I believe, their descendants, mostly in a modified condition, in the central parts of Europe and the United States. On this view we can understand the relations.h.i.+p, with very little ident.i.ty, between the productions of North America and Europe,--a relations.h.i.+p which is most remarkable, considering the distance of the two areas, and their separation by the Atlantic Ocean. We can further understand the singular fact remarked on by several observers, that the productions of Europe and America during the later tertiary stages were more closely related to each other than they are at the present time; for during these warmer periods the northern parts of the Old and New Worlds will have been almost continuously united by land, serving as a bridge, since rendered impa.s.sable by cold, for the inter-migration of their inhabitants.

During the slowly decreasing warmth of the Pliocene period, as soon as the species in common, which inhabited the New and Old Worlds, migrated south of the Polar Circle, they must have been completely cut off from each other. This separation, as far as the more temperate productions are concerned, took place long ages ago. And as the plants and animals migrated southward, they will have become mingled in the one great region with the native American productions, and have had to compete with them; and in the other great region, with those of the Old World. Consequently we have here everything favourable for much modification,--for far more modification than with the Alpine productions, left isolated, within a much more recent period, on the several mountain-ranges and on the arctic lands of the two Worlds. Hence it has come, that when we compare the now living productions of the temperate regions of the New and Old Worlds, we find very few identical species (though Asa Gray has lately shown that more plants are identical than was formerly supposed), but we find in every great cla.s.s many forms, which some naturalists rank as geographical races, and others as distinct species; and a host of closely allied or representative forms which are ranked by all naturalists as specifically distinct.

As on the land, so in the waters of the sea, a slow southern migration of a marine fauna, which during the Pliocene or even a somewhat earlier period, was nearly uniform along the continuous sh.o.r.es of the Polar Circle, will account, on the theory of modification, for many closely allied forms now living in areas completely sundered. Thus, I think, we can understand the presence of many existing and tertiary representative forms on the eastern and western sh.o.r.es of temperate North America; and the still more striking case of many closely allied crustaceans (as described in Dana's admirable work), of some fish and other marine animals, in the Mediterranean and in the seas of j.a.pan,--areas now separated by a continent and by nearly a hemisphere of equatorial ocean.