Island Life/XXIII

CHAPTER XXIII

ON THE ARCTIC ELEMENT IN SOUTH TEMPERATE FLORAS


 * European Species and Genera of Plants in the Southern Hemisphere—Aggressive Power of the Scandinavian Flora—Means by which Plants have Migrated from North to South—Newly moved Soil as Affording Temporary Stations to Migrating Plants—Elevation and Depression of the Snow-line as Aiding the Migration of Plants—Changes of Climate Favourable to Migration—The Migration from North to South has been long going on—Geological Changes as Aiding Migration—Proofs of Migration by way of the Andes—Proofs of Migration by way of the Himalayas and Southern Asia—Proofs of Migration by way of the African Highlands—Supposed Connection of South Africa and Australia—The Endemic Genera of Plants in New Zealand—The Absence of Southern Types from the Northern Hemisphere—Concluding Remarks on the New Zealand and South Temperate Floras.

We have now to deal with another portion of the New Zealand flora which presents perhaps equal difficulties—that which appears to have been derived from remote parts of the north and south temperate zones; and this will lead us to inquire into the origin of the northern or Arctic element in all the south temperate floras.

More than one-third of the entire number of New Zealand genera (115) are found also in Europe, and even fifty-eight species are identical in these remote parts of the world. Temperate South America has seventy-four genera in common with New Zealand, and there are even eleven species identical in the two countries, as well as thirty-two which are close allies or representative species. A considerable number of these northern or Antarctic plants and many more which are representative species, are found also in Tasmania and in the mountains of temperate Australia; and Sir Joseph Hooker gives a list of thirty-eight species very characteristic of Europe and Northern Asia, but almost or quite unknown in the warmer regions, which yet reappear in temperate Australia. Other genera seem altogether Antarctic—that is, confined to the extreme southern lands and islands; and these often have representative species in Southern America, Tasmania, and New Zealand, while others occur only in one or two of these areas. Many north temperate genera also occur in the mountains of South Africa. On the other hand, few if any of the peculiar Australian or Antarctic types have spread northwards, except some of the former which have reached the mountains of Borneo, and a few of the latter which spread along the Andes to Mexico.

On these remarkable facts, of which I have given but the barest outline, Sir Joseph Hooker makes the following suggestive observations:—

"When I take a comprehensive view of the vegetation of the Old World, I am struck with the appearance it presents of there being a continuous current of vegetation (if I may so fancifully express myself) from Scandinavia to Tasmania; along, in short, the whole extent of that arc of the terrestrial sphere which presents the greatest continuity of land. In the first place Scandinavian genera, and even species, reappear everywhere from Lapland and Iceland to the tops of the Tasmanian Alps, in rapidly diminishing numbers it is true, but in vigorous development throughout. They abound on the Alps and Pyrenees, pass on to the Caucasus and Himalayas, thence they extend along the Khasia Mountains, and those of the peninsulas of India to those of Ceylon and the Malayan Archipelago (Java and Borneo), and after a hiatus of 30° they appear on the Alps of New South Wales, Victoria, and Tasmania, and beyond these again on those of New Zealand and the Antarctic Islands, many of the species remaining unchanged throughout! It matters not what the vegetation of the bases and flanks of these mountains may be; the northern species may be associated with alpine forms of Germanic, Siberian, Oriental, Chinese, American, Malayan, and finally Australian, and Antarctic types; but whereas these are all, more or less, local assemblages, the Scandinavian asserts his prerogative of ubiquity from Britain to beyond its antipodes."

It is impossible to place the main facts more forcibly before the reader than in the above striking passage. It shows clearly that this portion of the New Zealand flora is due to wide-spread causes which have acted with even greater effect in other south temperate lands, and that in order to explain its origin we must grapple with the entire problem of the transfer of the north temperate flora to the southern hemisphere. Taking, therefore, the facts as given by Sir Joseph Hooker in the works already referred to, I shall discuss the whole question broadly, and shall endeavour to point out the general laws and subordinate causes that, in my opinion, have been at work in bringing about the anomalous phenomena of distribution he has done so much to make known and to elucidate.

Aggressive Power of the Scandinavian Flora.—The first important fact bearing upon this question is the wonderful aggressive and colonising power of the Scandinavian flora, as shown by the way in which it establishes itself in any temperate country to which it may gain access. About 150 species have thus established themselves in New Zealand, often taking possession of large tracts of country; about the same number are found in Australia, and nearly as many in the Atlantic states of America, where they form the commonest weeds. Whether or not we accept Mr. Darwin's explanation of this power as due to development in the most extensive land area of the globe where competition has been most severe and long-continued, the fact of the existence of this power remains, and we can see how important an agent it must be in the formation of the floras of any lands to which these aggressive plants have been able to gain access.

But not only are these plants pre-eminently capable of holding their own in any temperate country in the world, but they also have exceptional powers of migration and dispersal over seas and oceans. This is especially well shown by the case of the Azores, where no less than 400 out of a total of 478 flowering plants are identical with European species. These islands are more than 800 miles from Europe, and, as we have already seen in Chapter XII., there is no reason for supposing that they have ever been more nearly connected with it than they are now, since an extension of the European coast to the 1,000-fathom line would very little reduce the distance. Now it is a most interesting and suggestive fact that more than half the European genera which occur in the Australian flora occur also in the Azores, and in several cases even the species are identical in both. The importance of such a case as this cannot be exaggerated, because it affords a demonstration of the power of the very plants in question to pass over wide areas of sea, some no doubt wholly through the air, carried by storms in the same way as the European birds and insects which annually reach the Azores, others by floating on the waters, or by a combination of the two methods; while some may have been carried by aquatic birds, to whose feathers many seeds have the power of attaching themselves, and some even in the stomachs of fruit or seed eating birds. We have in such facts as these a complete disproof of the necessity for those great changes of sea and land which are continually appealed to by those who think land-connection the only efficient means of accounting for the migration of animals or plants; but at the same time we do not neglect to make the fullest use of such moderate changes as all the evidence at our command leads us to believe have actually occurred, and especially of the former existence of intermediate islands, so often indicated by shoals in the midst of the deepest oceans.

Means by which Plants have migrated from North to South.—But if plants can thus pass in considerable numbers and variety over wide seas and oceans, it must be yet more easy for them to traverse continuous areas of land, whereever mountain-chains offer suitable stations at moderate intervals on which they might temporarily establish themselves. The facilities afforded for the transmission of plants by mountains has hardly received sufficient attention. The numerous land-slips, the fresh surfaces of broken rock and precipice, the debris of torrents, and the moraines deposited by glaciers, afford numerous unoccupied stations on which wind-borne seeds have a good chance of germinating. It is a well-known fact that fresh surfaces of soil or rock, such as are presented by railway cuttings and embankments, often produce plants strange to the locality, which survive for a few years, and then disappear as the normal vegetation gains strength and permanence. But such a surface will, in the meantime, have acted as a fresh centre of dispersal; and thus a plant might pass on step by step, by means of stations temporarily occupied, till it reached a district  where, the general conditions being more favourable, it was able to establish itself as a permanent member of the flora. Such, generally speaking, was probably the process by which the Scandinavian flora has made its way to the southern hemisphere; but it could hardly have done so to any important extent without the aid of those powerful causes explained in our eighth chapter—causes which acted as a constantly recurrent motive-power to produce that "continuous current of vegetation" from north to south across the whole width of the tropics referred to by Sir Joseph Hooker. Those causes were, the repeated changes of climate which, during all geological time, appear to have occurred in both hemispheres, culminating at rare intervals in glacial epochs, and which have been shown to depend upon changes of excentricity of the earth's orbit and the occurrence of summer or winter in aphelion, in conjunction with the slower and more irregular changes of geographical conditions; these combined causes acting chiefly through the agency of heat-bearing oceanic currents, and of snow- and ice-collecting highlands. Let us now briefly consider how such changes would act in favouring the dispersal of plants.

Elevation and Depression of the Snow Line as Aiding the Migration of Plants.—We have endeavoured to show (in an earlier portion of this volume) that wherever geographical or physical conditions were such as to produce any considerable amount of perpetual snow, this would be increased whenever a high degree of excentricity concurred with winter in aphelion, and diminished during the opposite phase. On all mountain ranges, therefore, which reached above the snow-line, there would be a periodical increase and decrease of snow, and when there were extensive areas of plateau at about the same level, the lowering of the snow-line might cause such an increased accumulation of snow as to produce great glaciers and ice-fields, such as we have seen occurred in South Africa during the last period of high excentricity. But along with such depression of the line of perpetual snow there would be a corresponding depression of the alpine and sub-alpine zones suitable for the growth of an arctic and temperate vegetation, and, what is perhaps more important, the depression would necessarily produce a great extension of the area of these zones on all high mountains, because as we descend the average slopes become less abrupt,—thus affording a number of new stations suitable for such temperate plants as might first reach them. But just above and below the snow-line is the area of most powerful disintegration and denudation, from the alternate action of frost and sun, of ice and water; and thus the more extended area would be subject to the constant occurrence of land-slips, berg-falls, and floods, with their accompanying accumulations of débris and of alluvial soil, affording innumerable stations in which solitary wind-borne seeds might germinate and temporarily establish themselves.

This lowering and rising of the snow-line each 10,500 years during periods of high excentricity, would occur in the northern and southern hemispheres alternately; and where there were high mountains within the tropics the two would probably overlap each other, so that the northern depression would make itself felt in a slight degree even across the equator some way into the southern hemisphere, and vice versâ; and even if the difference of the height of perpetual snow at the two extremes did not average more than a few hundred feet, this would be amply sufficient to supply the new and unoccupied stations needful to facilitate the migration of plants. It is well known that all great mountain ranges have undergone such fluctuations, as proved by ice-marks below the present level of snow and ice.

But the differences of temperature in the two hemispheres caused by the sun being in perihelion in the winter of the one while it was in aphelion during the same season in the other, would necessarily lead to increased aërial and marine currents, as already explained; and whenever geographical conditions were such as to favour the production of glaciation in any area these effects would become more powerful, and would further aid in the dispersal of the seeds of plants.

Changes of Climate Favourable to Migration.—It is clear then, that during periods when no glacial epochs were produced in the northern hemisphere, and even when a mild climate extended over the whole polar area, alternate changes of climate favouring the dispersal of plants would occur on all high mountains, and with particular force on such as rise above the snow-line. But during that long-continued, though comparatively recent, phase of high excentricity which produced an extensive glaciation in the northern hemisphere and local glaciations in the southern, these risings and lowerings of the snow-line on all mountain ranges would have been at a maximum, and would have been increased by the depression of the ocean which must have arisen from such a vast bulk of water being locked up in land-ice, and which depression would have produced the same effect as a general elevation of all the continents. At this time, too, aërial currents would have attained their maximum of force in both hemispheres; and this would greatly facilitate the dispersal of all wind-borne seeds as well as of those carried in the plumage or in the stomachs of birds, since we have seen, by the cases of the Azores and Bermuda, how vastly the migratory powers of birds are increased by a stormy atmosphere.

Migration from North to South has been long going on.—Now, if each phase of colder and warmer mountain-climate—each alternate depression and elevation of the snow-line, only helped on the migration of a few species some stages of the long route from the north to the south temperate regions, yet, during the long course of the Tertiary period there might well have arisen that representation of the northern flora in the southern hemisphere which is now so conspicuous. For it is very important to remark that it is not the existing flora alone that is represented, such as might have been conveyed during the last glacial epoch only; but we find a whole series of northern types evidently of varying degrees of antiquity, while even some genera characteristic of the southern hemisphere appear to have been originally derived from Europe. Thus Eucalyptus and Metrosideros have been determined by Dr. Ettingshausen from their fruits in the Eocene beds of Sheppey, while Pimelea, Leptomeria and four genera of Proteaceæ have been recognised by Professor Heer in the Miocene of Switzerland; and the former writer has detected fifty-five Australian forms in the Eocene plant beds of Häring (? Belgium). Then we have such peculiar genera as Pachychladon and Notothlaspi of New Zealand said to have affinities with Arctic plants, while Stilbocarpa—another peculiar New Zealand genus—has its nearest allies in the Himalayan and Chinese Aralias. Following these are a whole host of very distinct species of northern genera which may date back to any part of the Tertiary period, and which occur in every south temperate land. Then we have closely allied representative species of European or Arctic plants; and, lastly, a number of identical species,—and these two classes are probably due entirely to the action of the last great glacial epoch, whose long continuance, and the repeated fluctuations of climate with which it commenced and terminated, rendered it an agent of sufficient power to have brought about this result.

Here, then, we have that constant or constantly recurrent process of dispersal acting throughout long periods with varying power—that "continuous current of vegetation" as it has been termed, which the facts demand; and the extraordinary phenomenon of the species and genera of European and even of Arctic plants being represented abundantly in South America, Australia, and New Zealand, thus adds another to the long series of phenomena which are rendered intelligible by frequent alternations of warmer and colder climates in either hemisphere, culminating, at long intervals and in favourable situations, in actual glacial epochs.

Geological Changes as Aiding Migration.—It will be well also to notice here, that there is another aid to dispersion dependent upon the changes effected by denudation during the long periods included in the duration of the species and genera of plants. A considerable number of the plants of the Miocene period of Europe were so much like existing species that although they have generally received fresh names they may well have been identical; and a large proportion of the vegetation during the whole Tertiary period consisted of genera which are still living. But from what is now known of the rate of sub-aërial denudation, we are sure, that during each division of this period many mountain chains must have been considerably lowered, while we know that some of the existing ranges have been greatly elevated. Ancient volcanoes, too, have been destroyed by denudation, and new ones have been built up, so that we may be quite sure that ample means for the transmission of temperate plants across the tropics, may have existed in countries where they are now no longer to be found. The great mountain masses of Guiana and Brazil, for example, must have been far more lofty before the sedimentary covering was denuded from their granitic bosses and metamorphic peaks, and may have aided the southern migration of plants before the final elevation of the Andes. And if Africa presents us with an example of a continent of vast antiquity, we may be sure that its great central plateaux once bore far loftier mountain ranges before they were reduced to their present condition by long ages of denudation.

Proofs of Migration by Way of the Andes.—We are now prepared to apply the principles above laid down to the explanation of the character and affinities of the various portions of the north temperate flora in the southern hemisphere, and especially in Australia and New Zealand.

At the present time the only unbroken chain of highlands and mountains connecting the Arctic and north temperate with the Antarctic lands is to be found in the American continent, the only break of importance being the comparatively low Isthmus of Panama, where there is a distance of about 300 miles occupied by rugged forest-clad hills, between the lofty peaks of Veragua and the northern extremity of the Andes of New Grenada. Such distances are, as we have already seen, no barrier to the diffusion of plants; and we should accordingly expect that this great continuous mountain-chain has formed the most effective agent in aiding the southward migration of the Arctic and north temperate vegetation. We do find, in fact, not only that a large number of northern genera and many species are scattered all along this line of route, but that at the end of the long journey, in Southern Chile and Fuegia, they have established themselves in such numbers as to form an important part of the flora of those countries. From the lists given in the works already referred to, it appears that there are between sixty and seventy northern genera in Fuegia and Southern Chile, while about forty of the species are absolutely identical with those of Europe and the Arctic regions. Considering how comparatively little the mountains of South Temperate America are yet known, this is a very remarkable result, and it proves that the transmission of species must have gone on up to comparatively recent times. Yet, as only a few of these species are now found along the line of migration, we see that they only occupied such stations temporarily; and we may connect their disappearance with the passing away of the last glacial period which, by raising the snow-line, reduced the area on which alone they could exist, and exposed them to the competition of indigenous plants from the belt of country immediately below them.

Now, just as these numerous species and genera have undoubtedly passed along the great American range of mountains, although only now found at its two extremes, so others have doubtless passed on further; and have found more suitable stations or less severe competition in the Antarctic continent and islands, in New Zealand, in Tasmania, and even in Australia itself. The route by which they may have reached these countries is easily marked out. Immediately south of Cape Horn, at a distance of only 500 miles, are the South Shetland Islands and Graham's Land, whence the Antarctic continent or a group of large islands probably extends across or around the south polar area to Victoria Land and thence to Adélie Land. The outlying Young Island, 12,000 feet high, is about 750 miles south of the Macquarie Islands, which may be considered a southern outlier of the New Zealand group; and the Macquarie Islands are about the same distance from the 1,000-fathom line at a point marking the probable southern extension of Tasmania. Other islands may have existed at intermediate points; but, even as it is, these distances are not greater than we know are traversed by plants both by flotation and by aërial currents, especially in such a stormy atmosphere as that of the Antarctic regions. Now, we may further assume, that what we know occurred within the Arctic circle also took place in the Antarctic—that is, that there have been alternations of climate during which some portion of what are now ice-clad lands became able to support a considerable amount of vegetation. During such periods there would be a steady migration of plants from all southern circumpolar countries to people the comparatively unoccupied continent, and the southern extremity of America being considerably the nearest, and also being the best stocked with those northern types which have such great powers of migration and colonisation, such plants would form the bulk of the Antarctic vegetation, and during the continuance of the milder southern climate would occupy the whole area.

When the cold returned and the land again became ice-clad, these plants would be crowded towards the outer margins of the Antarctic land and its islands, and some of them would find their way across the sea to such countries as offered on their mountain summits suitable cool stations; and as this process of alternately receiving plants from Chile and Fuegia and transmitting them in all directions from the central Antarctic land may have been repeated several times during the Tertiary period, we have no difficulty in understanding the general community between the European and Antarctic plants found in all south temperate lands. Kerguelen's Land and The Crozets are within about the same distance from the Antarctic continent as New Zealand and Tasmania, and we need not therefore be surprised at finding in each of these islands some Fuegian species which have not reached the others. Of course, there will remain difficulties of detail, as there always must remain, so long as our knowledge of the past changes of the earth's surface and the history of the particular plants concerned is so imperfect. Sir Joseph Hooker notes, for example, the curious fact that several Compositæ common to three such remote localities as the Auckland Islands, Fuegia, and Kerguelen's Land, have no pappus or seed-down, while such as have pappus are in no case common even to two of these islands. Without knowing the exact history and distribution of the genera to which these plants belong it would be useless to offer any conjecture, except that they are ancient forms which may have survived great geographical changes, or may have some peculiar and exceptional means of dispersion.

Proofs of Migration by way of the Himalayas and Southern Asia.—But although we may thus explain the presence of a considerable portion of the European element in the floras of New Zealand and Australia, we cannot account for the whole of it by this means, because Australia itself contains a host of European and Asiatic genera of which we find no trace in New Zealand or South America, or any other Antarctic land. We find, in fact, in Australia two distinct sets of European plants. First we have a number of species identical with those of Northern Europe or Asia (of the most characteristic of which—thirty-eight in number—Sir Joseph Hooker gives a list); and in the second place a series of European genera usually of a somewhat more southern character, mostly represented by very distinct species, and all absent from New Zealand; such as Clematis, Papaver, Cleome, Polygala, Lavatera, Ajuga, &c. Now of the first set—the North European species—about three-fourths occur in some parts of America, and about half in South Temperate America or New Zealand; whence we may conclude that most of these, as well as some others, have reached Australia by the route already indicated. The second set of Australo-European genera, however, and many others characteristic of the South European or the Himalayan flora, have probably reached Australia by way of the mountains of Southern Asia, Borneo, the Moluccas, and New Guinea, at a somewhat remote period when loftier ranges and some intermediate peaks may have existed, sufficient to carry on the migration by the aid of the alternate climatal changes which are known to have occurred. The long belt of Secondary and Palæozoic formations in East Australia from Tasmania to Cape York continued by the lofty ranges of New Guinea, indicates the route of this immigration, and sufficiently explains how it is that these northern types are almost wholly confined to this part of the Australian continent. Some of the earlier immigrants of this class no doubt passed over to New Zealand and now form a portion of the peculiar genera confined to these two countries; but most of them are of later date, and have thus remained in Australia only.

Proofs of Migration by way of the African Highlands.—It is owing to this twofold current of vegetation flowing into Australia by widely different routes that we have in this distant land a better representation of the European flora, both as regards species and genera, than in any other part of the southern hemisphere; and, so far as I can judge of the facts, there is no general phenomenon—that is, nothing in the distribution of genera and other groups of plants as opposed to cases of individual species—that is not fairly accounted for by such an origin. It further receives support from the case of South Africa, which also contains a large and important representation of the northern flora. But here we see no indications (or very slight ones) of that southern influx which has given Australia such a community of vegetation with the Antarctic lands. There are no less than sixty genera of strictly north temperate plants in South Africa, none of which occur in Australia; while very few of the species, so characteristic of Australia, New Zealand, and Fuegia, are found there. It is clear, therefore, that South Africa has received its European plants by the direct route through the Abyssinian highlands and the lofty equatorial mountains, and mostly at a distant period when the conditions for migration were somewhat more favourable than they are now. The much greater directness of the route from Northern Europe to South Africa than to Australia; and the existence even now of lofty mountains and extensive highlands for a large portion of the distance, will explain (what Sir Joseph Hooker notes as "a very curious fact") why South Africa has more very northern European genera than Australia, while Australia has more identical species and a better representation on the whole of the European flora—this being clearly due to the large influx of species it has received from the Antarctic Islands, in addition to those which have entered it by way of Asia. The greater distance of South Africa even now from any of these islands, and the much deeper sea to the south of the African continent, than in the case of Tasmania and New Zealand, indicating a smaller recent extension southward, is all quite in harmony with the facts of distribution of the northern flora above referred to.

Supposed Connection of South Africa and Australia.—There remains, however, the small amount of direct affinity between the vegetation of South Africa and that of Australia, New Zealand, and Temperate South America, consisting in all of fifteen genera, five of which are confined to Australia and South Africa, while several natural orders are better represented in these two countries than in any other part of the world. This resemblance has been supposed to imply some former land-connection of all the great southern lands, but it appears to me that any such supposition is wholly unnecessary. The differences between the faunas and floras of these countries are too great and too radical to render it possible that any such connection should have existed except at a very remote period. But if we have to go back so far for an explanation, a much simpler one presents itself, and one more in accordance with what we have learnt of the general permanence of deep oceans and the great changes that have taken place in the distribution of all forms of life. Just as we explain the presence of marsupials in Australia and America and of Centetidæ in Madagascar and the Antilles, by the preservation in these localities of remnants of once wide-spread types, so we should prefer to consider the few genera common to Australia and South Africa as remnants of an ancient vegetation, once spread over the northern hemisphere, driven southward by the pressure of more specialised types, and now finding a refuge in these two widely separated southern lands. It is suggestive of such an explanation that these genera are either of very ancient groups—as Conifers and Cycads—or plants of low organisation as the Restiaceæ—or of world-wide distribution, as Melanthaceæ.

The Endemic Genera of Plants in New Zealand.—Returning now to the New Zealand flora, with which we are more especially concerned, there only remains to be considered the peculiar or endemic genera which characterise it. These are thirty-two in number, and are mostly very isolated. A few have affinities with Arctic groups, others with Himalayan, or Australian genera; several are tropical forms, but the majority appear to be altogether peculiar types of world-wide groups—as Leguminosæ, Saxifrageæ, Compositæ, Orchideæ, &c. We must evidently trace back these peculiar forms to the earliest immigrants, either from the north or from the south; and the great antiquity we are obliged to give to New Zealand—an antiquity supported by every feature in its fauna and flora, no less than by its geological structure, and its extinct forms of life—affords ample time for the changes in the general distribution of plants, and for those due to isolation and modification under the influence of changed conditions, which are manifested by the extreme peculiarity of many of these interesting endemic forms.

The Absence of Southern Types from the Northern Hemisphere.—We have now only to notice the singular want of reciprocity in the migrations of northern and southern types of vegetation. In return for the vast number of European plants which have reached Australia, not one single Australian plant has entered any part of the north temperate zone, and the same may be said of the typical southern vegetation in general, whether developed in the Antarctic lands, New Zealand, South America, or South Africa. The furthest northern outliers of the southern flora are a few genera of Antarctic type on the Bornean Alps; the genus Acæna which has a species in California; two representatives of the Australian flora—Casuarina and Stylidium, in the peninsula of India; while China and the Philippines have two strictly Australian genera of Orchideæ—Microtis and Thelymitra, as well as a Restiaceous genus. Several distinct causes appear to have combined to produce this curious inability of the southern flora to make its way into the northern hemisphere. The primary cause is, no doubt, the totally different distribution of land in the two hemispheres, so that in the south there is the minimum of land in the colder parts of the temperate zone and in the north the maximum. This is well shown by the fact that on the parallel of Lat. 50° N. we pass over 240° of land or shallow sea, while on the same parallel of south latitude we have only 4°, where we cross the southern part of Patagonia. Again the three most important south temperate land-areas—South Temperate America, South Africa, and Australia—are widely separated from each other, and have in all probability always been so; whereas the whole of the north temperate lands are practically continuous. It follows that, instead of the enormous northern area, in which highly organised and dominant groups of plants have been developed gifted with great colonising and aggressive powers, we have in the south three comparatively small and detached areas, in which rich floras have been developed with special adaptations to soil, climate, and organic environment, but comparatively impotent and inferior beyond their own domain.

Another circumstance which makes the contest between the northern and southern forms still more unequal, is the much greater hardiness of the former, from having been developed in a colder region, and one where alpine and arctic conditions extensively prevail; whereas the southern floras have been mainly developed in mild regions to which they have been altogether confined. While the northern plants have been driven north or south by each succeeding change of climate, the southern species have undergone comparatively slight changes of this nature, owing to the areas they occupy being unconnected with the ice-bearing Antarctic continent. It follows, that whereas the northern plants find in all these southern lands a milder and more equable climate than that to which they have been accustomed, and are thus often able to grow and flourish even more vigorously than in their native land, the southern plants would find in almost every part of Europe, North America or Northern Asia, a more severe and less equable climate, with winters that usually prove fatal to them even under cultivation. These causes, taken separately, are very powerful, but when combined they must, I think, be held to be amply sufficient to explain why examples of the typical southern vegetation are almost unknown in the north temperate zone, while a very few of them have extended so far as the northern tropic.

Concluding Remarks on the Last Two Chapters.—Our inquiry into the external relations and probable origin of the fauna and flora of New Zealand, has thus led us on to a general theory as to the cause of the peculiar biological relations between the northern and the southern hemispheres; and no better or more typical example could be found of the wide range and great interest of the study of the geographical distribution of animals and plants.

The solution which has here been given of one of the most difficult of this class of problems, has been rendered possible solely by the knowledge very recently obtained of the form of the sea-bottom in the southern ocean, and of the geological structure of the great Australian continent. Without this knowledge we should have nothing but a series of guesses or probabilities on which to found our hypothetical explanation, which we have now been able to build up on a solid foundation of fact. The complete separation of East from West Australia during a portion of the Cretaceous and Tertiary periods, could never have been guessed till it was established by the laborious explorations of the Australian geologists; while the hypothesis of a comparatively shallow sea, uniting New Zealand by a long route with tropical Australia, while a profoundly deep ocean always separated it from temperate Australia, would have been rejected as too improbable a supposition for the foundation of even the most enticing theory. Yet it is mainly by means of these two facts, that we are enabled to give an adequate explanation of the strange anomalies in the flora of Australia and its relation to that of New Zealand.

In the more general explanation of the relations of the various northern and southern floras, I have shown what an important aid to any such explanation is the theory of repeated changes of climate, not necessarily of great amount, given in Chapters VIII. and IX.; while the whole discussion justifies the importance attached to the theory of the general permanence of continents and oceans, as demonstrated in Chapter VI., since any rational explanation based upon facts (as opposed to mere unsupported conjecture) must take such general permanence as a starting-point. The whole inquiry into the phenomena presented by islands, which forms the main subject of the present volume has, I think, shown that this theory does afford a firm foundation for the discussion of questions of distribution and dispersal; and that by its aid, combined with a clear perception of the wonderful powers of dispersion and modification in the organic world when long periods are considered, the most difficult problems connected with this subject cease to be insoluble.

189 Introductory Essay On the Flora of Australia, p. 130.

190 Hooker, On the Flora of Australia, p. 95.—H. C. Watson, in Godman's Azores, pp. 278-286.

191 As this is a point of great interest in its bearing on the dispersal of plants by means of mountain ranges, I have endeavoured to obtain a few illustrative facts:—

1. Mr. William Mitten, of Hurstpierpoint, Sussex, informs me that when the London and Brighton railway was in progress in his neighbourhood, Melilotus vulgaris made its appearance on the banks, remained for several years, and then altogether disappeared. Another case is that of Diplotaxis muralis, which formerly occurred only near the sea-coast of Sussex, and at Lewes; but since the railway was made has spread along it, and still maintains itself abundantly on the railway banks though rarely found anywhere else.

2. A correspondent in Tasmania informs me that whenever the virgin forest is cleared in that island there invariably comes up a thick crop of a plant locally known as fire-weed—a species of Senecio, probably S. Australis. It never grows except where the fire has gone over the ground, and is unknown except in such places. My correspondent adds:—"This autumn I went back about thirty-five miles through a dense forest, along a track marked by some prospectors the year before, and in one spot where they had camped, and the fire had burnt the fallen logs, &c., there was a fine crop of 'fire-weed.' All around for many miles was a forest of the largest trees and dense scrub." Here we have a case in which burnt soil and ashes favour the germination of a particular plant, whose seeds are easily carried by the wind, and it is not difficult to see how this peculiarity might favour the dispersal of the species for enormous distances, by enabling it temporarily to grow and produce seeds on burnt spots.

3. In answer to an inquiry on this subject, Mr. H. C. Watson has been kind enough to send me a detailed account of the progress of vegetation on the railway banks and cuttings about Thames Ditton. This account is written from memory, but as Mr. Watson states that he took a great interest in watching the process year by year, there can be no reason to doubt the accuracy of his memory. I give a few extracts which bear especially on the subject we are discussing.

"One rather remarkable biennial plant appeared early (the second year, as I recollect) and renewed itself either two or three years, namely, Isatis tinctoria—a species usually supposed, to be one of our introduced, but pretty well naturalised, plants. The nearest stations then or since known to me for this Isatis are on chalk about Guildford, twenty miles distant. There were two or three plants of it at first, never more than half a dozen. Once since I saw a plant of Isatis on the railway bank near Vauxhall.

"Close by Ditton Station three species appeared which may be called interlopers. The biennial Barbarea precox, one of these, is the least remarkable, because it might have come as seed in the earth from some garden, or possibly in the Thames gravel (used as ballast). At first it increased to several plants, then became less numerous, and will soon, in all probability, become extinct, crowded out by other plants. The biennial Petroselinum segetum was at first one very luxuriant plant on the slope of the embankment. It increased by seed into a dozen or a score, and is now nearly if not quite extinct. The third species is Linaria purpurea, not strictly a British plant, but one established in some places on old walls. A single root of it appeared on the chalk facing of the embankment by Ditton Station. It has remained there several years and grown into a vigorous specimen. Two or three smaller examples are now seen by it, doubtless sprung from some of the hundreds or thousands of seeds shed by the original one plant. The species is not included in Salmon and Brewer's Flora of Surrey.

"The main line of the railway has introduced into Ditton parish the perennial Arabis hirsuta, likely to become a permanent inhabitant. The species is found on the chalk and greensand miles away from Thames Ditton; but neither in this parish nor in any adjacent parish, so far as known to myself or to the authors of the flora of the county, does it occur. Some years after the railway was made a single root of this Arabis was observed in the brickwork of an arch by which the railway is carried over a public road. A year or two afterwards there were three or four plants. In some later year I laid some of the ripened seed-pods between the bricks in places where the mortar had partly crumbled out. Now there are several scores of specimens in the brickwork of the arch. It is presumable that the first seed may have been brought from Guildford. But how could it get on to the perpendicular face of the brickwork?

"The Bee Orchis (Ophrys apifera), plentiful on some of the chalk lands in Surrey, is not a species of Thames Ditton, or (as I presume) of any adjacent parish. Thus, I was greatly surprised some years back to see about a hundred examples of it in flower in one clayey field either on the outskirts of Thames Ditton or just within the limits of the adjoining parish of Cobham. I had crossed this same field in a former year without observing the Ophrys there. And on finding it in the one field I closely searched the surrounding fields and copses, without finding it anywhere else. Gradually the plants became fewer and fewer in that one field, and some six or eight years after its first discovery there the species had quite disappeared again. I guessed it had been introduced with chalk, but could obtain no evidence to show this."

4. Mr. A. Bennett, of Croydon, has kindly furnished me with some information on the temporary vegetation of the banks and cuttings on the railway from Yarmouth to Caistor in Norfolk, where it passes over extensive sandy Denes with a sparse vegetation. The first year after the railway was made the banks produced abundance of Œnothera odorata and Delphinium Ajacis (the latter only known thirty miles off in cornfields in Cambridgeshire), with Atriplex patula and A. deltoidea. Gradually the native sand plants—Carices, Grasses, Galium verum, &c., established themselves, and year by year covered more ground till the new introductions almost completely disappeared. The same phenomenon was observed in Cambridgeshire between Chesterton and Newmarket, where, the soil being different, Stellaria media and other annuals appeared in large patches; but these soon gave way to a permanent vegetation of grasses, composites, &c., so that in the third year no Stellaria was to be seen.

5. Mr. T. Kirk (writing in 1878) states that—"in Auckland, where a dense sward of grass is soon formed, single specimens of the European milk Thistle (Carduus marianus) have been known for the past fifteen years; but although they seeded freely, the seeds had no opportunity of germinating, so that the thistle did not spread. A remarkable exception to this rule occurred during the formation of the Onehunga railway, where a few seeds fell on disturbed soil, grew up and flowered. The railway works being suspended, the plant increased rapidly, and spread wherever it could find disturbed soil."

Again:—"The fiddle-dock (Rumex pulcher) occurs in great abundance on the formation of new streets, &c., but soon becomes comparatively rare. It seems probable that it was one of the earliest plants naturalised here, but that it partially died out, its buried seeds retaining their vitality."

Medicago sativa and Apium graveolens, are also noted as escapes from cultivation which maintain themselves for a time but soon die out. (Transactions of the New Zealand Institute, Vol. X. p. 367.)

The preceding examples of the temporary establishment of plants on newly exposed soil, often at considerable distances from the localities they usually inhabit, might, no doubt, by further inquiry be greatly multiplied; but, unfortunately, the phenomenon has received little attention, and is not even referred to in the elaborate work of De Candolle (Géographie Botanique Raisonnée) in which almost every other aspect of the dispersion and distribution of plants is fully discussed. Enough has been advanced, however, to show that it is of constant occurrence, and from the point of view here advocated it becomes of great importance in explaining the almost world-wide distribution of many common plants of the north temperate zone.

192 Sir Joseph Hooker informs me that he considers these identifications worthless, and Mr. Bentham has also written very strongly against the value of similar identifications by Heer and Unger. Giving due weight to the opinions of these eminent botanists we must admit that Australian genera have not yet been demonstrated to have existed in Europe during the Tertiary period; but, on the other hand, the evidence that they did so appears to have some weight, on account of the improbability that the numerous resemblances to Australian plants which have been noticed by different observers should all be illusory; while the well established fact of the former wide distribution of many tropical or now restricted types of plants and animals, so frequently illustrated in the present volume, removes the antecedent improbability which is supposed to attach to such identifications. I am myself the more inclined to accept them, because, according to the views here advocated, such migrations must have taken place at remote as well as at recent epochs; and the preservation of some of these types in Australia while they have become extinct in Europe, is exactly paralleled by numerous facts in the distribution of animals which have been already referred to in Chapter XIX., and elsewhere in this volume, and also repeatedly in my larger work.

193 Out of forty-two genera from the Eocene of Sheppey enumerated by Dr. Ettingshausen in the Geological Magazine for January 1880, only two or three appear to be extinct, while there is a most extraordinary intermixture of tropical and temperate forms—Musa, Nipa, and Victoria, with Corylus, Prunus, Acer, &c. The rich Miocene flora of Switzerland, described by Professor Heer, presents a still larger proportion of living genera.

194 The recent discovery by Lieutenant Jensen of a rich flora on rocky peaks rising out of the continental ice of Greenland, as well as the abundant vegetation of the highest northern latitudes, renders it possible that even now the Antarctic continent may not be wholly destitute of vegetation, although its climate and physical condition are far less favourable than those of the Arctic lands. (See Nature, Vol. XXI. p. 345.)

195 Dr. Hector notes the occurrence of the genus Dammara in Triassic deposits, while in the Jurassic period New Zealand possessed the genera Palæozamia, Oleandrium, Alethopteris, Camptopteris, Cycadites, Echinostrobus, &c., all Indian forms of the same age. Neocomian beds contain a true dicotyledonous leaf with Dammara and Araucaria. The Cretaceous deposits have produced a rich flora of dicotyledonous plants, many of which are of the same genera as the existing flora; while the Miocene and other Tertiary deposits produce plants almost identical with those now inhabiting the country, together with many North Temperate genera which have since become extinct. (See p. 499, footnote, and Trans. New Zealand Inst., Vol. XI. 1879, p. 536.)

196 The fact stated in the last edition of the Origin of Species (p. 340) on the authority of Sir Joseph Hooker, that Australian plants are rapidly sowing themselves and becoming naturalised on the Neilgherrie mountains in the southern part of the Indian Peninsula, though an exception to the rule of the inability of Australian plants to become naturalised in the Northern Hemisphere, is yet quite in harmony with the hypothesis here advocated. For not only is the climate of the Neilgherries more favourable to Australian plants than any part of the North Temperate zone, but the entire Indian Peninsula has existed for unknown ages as an island and thus possesses the "insular" characteristic of a comparatively poor and less developed flora and fauna as compared with the truly "continental" Malayan and Himalayan regions. Australian plants are thus enabled to compete with those of the Indian Peninsula highlands with a fair chance of success.