Kitabı oku: «Island Life; Or, The Phenomena and Causes of Insular Faunas and Floras», sayfa 31

If New Zealand has really gone through such a series of changes as here suggested, some proofs of it might perhaps be obtained in the outlying islands which were once, presumably, joined with it. And this gives great importance to the statement of the aborigines of the Chatham Islands, that the Apteryx formerly lived there but was exterminated about 1835. It is to be hoped that some search will be made here and also in Norfolk Island, in both of which it is not improbable remains either of Apteryx or Dinornis might be discovered.

So far we find nothing to object to in the speculations of Captain Hutton, with which, on the contrary, we almost wholly concur; but we cannot follow him when he goes on to suggest an Antarctic continent uniting New Zealand and Australia with South America, and probably also with South Africa, in order to explain the existing distribution of struthious birds. Our best anatomists, as we have seen, agree that both Dinornis and Apteryx are more nearly allied to the cassowaries and emus than to the ostriches and rheas; and we see that the form of the sea-bottom suggests a former connection with North Australia and New Guinea—the very region where these types most abound, and where in all probability they originated. The suggestion that all the struthious birds of the world sprang from a common ancestor at no very remote period, and that their existing distribution is due to direct land communication between the countries they now inhabit, is one utterly opposed to all sound principles of reasoning in questions of geographical distribution. For it depends upon two assumptions, both of which are at least doubtful, if not certainly false—the first, that their distribution over the globe has never in past ages been very different from what it is now; and the second, that the ancestral forms of these birds never had the power of flight. As to the first assumption, we have found in almost every case that groups now scattered over two or more continents formerly lived in intervening areas of existing land. Thus the marsupials of South America and Australia are connected by forms which lived in North America and Europe; the camels of Asia and the llamas of the Andes had many extinct common ancestors in North America; the lemurs of Africa and Asia had their ancestors in Europe, as had the trogons of South America, Africa, and tropical Asia. But besides this general evidence we have direct proof that the struthious birds had a wider range in past times than now. Remains of extinct rheas have been found in Central Brazil, and those of ostriches in North India; while remains, believed to be of struthious birds, are found in the Eocene deposits of England; and the Cretaceous rocks of North America have yielded the extraordinary toothed bird, Hesperornis, which Professor O. Marsh declares to have been "a carnivorous swimming ostrich."

As to the second point, we have the remarkable fact that all known birds of this group have not only the rudiments of wing-bones, but also the rudiments of wings, that is, an external limb bearing rigid quills or largely-developed plumes. In the cassowary these wing-feathers are reduced to long spines like porcupine-quills, while even in the Apteryx, the minute external wing bears a series of nearly twenty stiff quill-like feathers.¹⁷⁵ These facts render it almost certain that the struthious birds do not owe their imperfect wings to a direct evolution from a reptilian type, but to a retrograde development from some low form of winged birds, analogous to that which has produced the dodo and the solitaire from the more highly-developed pigeon-type. Professor Marsh has proved, that so far back as the Cretaceous period, the two great forms of birds—those with a keeled sternum and fairly-developed wings, and those with a convex keel-less sternum and rudimentary wings—already existed side by side; while in the still earlier Archæopteryx of the Jurassic period we have a bird with well-developed wings, and therefore probably with a keeled sternum. We are evidently, therefore, very far from a knowledge of the earliest stages of bird life, and our acquaintance with the various forms that have existed is scanty in the extreme; but we may be sure that birds acquired wings, and feathers, and some power of flight, before they developed a keeled sternum, since we see that bats with no such keel fly very well. Since, therefore, the struthious birds all have perfect feathers, and all have rudimentary wings, which are anatomically those of true birds, not the rudimentary fore-legs of reptiles, and since we know that in many higher groups of birds—as the pigeons and the rails—the wings have become more or less aborted, and the keel of the sternum greatly reduced in size by disuse, it seems probable that the very remote ancestors of the rhea, the cassowary, and the apteryx, were true flying birds, although not perhaps provided with a keeled sternum, or possessing very great powers of flight. But in addition to the possible ancestral power of flight, we have the undoubted fact that the rhea and the emu both swim freely, the former having been seen swimming from island to island off the coast of Patagonia. This, taken in connection with the wonderful aquatic ostrich of the Cretaceous period discovered by Professor Marsh, opens up fresh possibilities of migration; while the immense antiquity thus given to the group and their universal distribution in past time, renders all suggestions of special modes of communication between the parts of the globe in which their scattered remnants now happen to exist, altogether superfluous and misleading.

The bearing of this argument on our present subject is, that so far as accounting for the presence of wingless birds in New Zealand is concerned, we have nothing whatever to do with any possible connection, by way of a southern continent or antarctic islands, with South America and South Africa, because the nearest allies of its moas and kiwis are the cassowaries and emus, and we have distinct indications of a former land extension towards North Australia and New Guinea, which is exactly what we require for the original entrance of the struthious type into the New Zealand area.

Winged Birds and Lower Vertebrates of New Zealand.—Having given a pretty full account of the New Zealand fauna elsewhere¹⁷⁶ I need only here point out its bearing on the hypothesis now advanced, of the former land-connection having been with North Australia, New Guinea, and the Western Pacific Islands, rather than with the temperate regions of Australia.

Of the Australian genera of birds, which are found also in New Zealand, almost every one ranges also into New Guinea or the Pacific Islands, while the few that do not extend beyond Australia are found in its northern districts. As regards the peculiar New Zealand genera, all whose affinities can be traced are allied to birds which belong to the tropical parts of the Australian region; while the starling family, to which four of the most remarkable New Zealand birds belong (the genera Creadion, Heterolocha, and Callæas), is totally wanting in temperate Australia and is comparatively scarce in the entire Australian region, but is abundant in the Oriental region, with which New Guinea and the Moluccas are in easy communication. It is certainly a most suggestive fact that there are more than sixty genera of birds peculiar to the Australian continent (with Tasmania), many of them almost or quite confined to its temperate portions, and that no single one of these should be represented in temperate New Zealand.¹⁷⁷ The affinities of the living and more highly organised, no less than those of the extinct and wingless birds, strikingly accord with the line of communication indicated by the deep submarine bank connecting these temperate islands with the tropical parts of the Australian region.

The reptiles, so far as they go, are quite in accordance with the birds. The lizards belong to two genera, Lygosoma, which has a wide range in all the tropics as well as in Australia; and Naultinus, a genus peculiar to New Zealand, but belonging to a family—Geckonidæ—spread over the whole of the warmer parts of the world. Australia, with New Guinea, on the other hand, has a peculiar family, and no less than twenty-one peculiar genera of lizards, many of which are confined to its temperate regions, but no one of them extends to temperate New Zealand.¹⁷⁸ The extraordinary lizard-like Hatteria punctata of New Zealand forms of itself a distinct order of reptiles, in some respects intermediate between lizards and crocodiles, and having therefore no affinity with any living animal.

The only representative of the Amphibia in New Zealand is a solitary frog of a peculiar genus (Liopelma hochstetteri); but it has no affinity for any of the Australian frogs, which are numerous, and belong to eleven different families; while the Liopelma belongs to a very distinct family (Discoglossidæ), confined to the Palæarctic region.

Of the fresh-water fishes we need only say here, that none belong to peculiar Australian types, but are related to those of temperate South America or of Asia.

The Invertebrate classes are comparatively little known, and their modes of dispersal are so varied and exceptional that the facts presented by their distribution can add little weight to those already adduced. We will, therefore, now proceed to the conclusions which can fairly be drawn from the general facts of New Zealand natural history already known to us.

Deductions from the Peculiarities of the New Zealand Fauna.—The total absence (or extreme scarcity) of mammals in New Zealand obliges us to place its union with North Australia and New Guinea at a very remote epoch. We must either go back to a time when Australia itself had not yet received the ancestral forms of its present marsupials and monotremes, or we must suppose that the portion of Australia with which New Zealand was connected was then itself isolated from the mainland, and was thus without a mammalian population. We shall see in our next chapter that there are certain facts in the distribution of plants, no less than in the geological structure of the country, which favour the latter view. But we must on any supposition place the union very far back, to account for the total want of identity between the winged birds of New Zealand and those peculiar to Australia, and a similar want of accordance in the lizards, the fresh-water fishes, and the more important insect-groups of the two countries. From what we know of the long geological duration of the generic types of these groups we must certainly go back to the earlier portion of the Tertiary period at least, in order that there should be such a complete disseverance as exists between the characteristic animals of the two countries; and we must further suppose that, since their separation, there has been no subsequent union or sufficiently near approach to allow of any important intermigration, even of winged birds, between them. It seems probable, therefore, that the Bampton shoal west of New Caledonia, and Lord Howe's Island further south, formed the western limits of that extensive land in which the great wingless birds and other isolated members of the New Zealand fauna were developed. Whether this early land extended eastward to the Chatham Islands and southward to the Macquaries we have no means of ascertaining, but as the intervening sea appears to be not more than about 1,500 fathoms deep it is quite possible that such an amount of subsidence may have occurred. It is possible, too, that there may have been an extension northward to the Kermadec Islands, and even further to the Tonga and Fiji Islands, though this is hardly probable, or we should find more community between their productions and those of New Zealand.

A southern extension towards the Antarctic continent at a somewhat later period seems more probable, as affording an easy passage for the numerous species of South American and Antarctic plants, and also for the identical and closely allied fresh-water fishes of these countries.

The subsequent breaking up of this extensive land into a number of separate islands in which the distinct species of moa and kiwi were developed—their union at a later period, and the final submergence of all but the existing islands, is a pure hypothesis, which seems necessary to explain the occurrence of so many species of these birds in a small area but of which we have no independent proof. There are, however, some other facts which would be explained by it, as the presence of three peculiar but allied genera of starlings, the three species of parrots of the genus Nestor, and the six distinct rails of the genus Ocydromus, as well as the numerous species in some of the peculiar New Zealand genera of plants, which seem less likely to have been developed in a single area than when isolated, and thus preserved from the counteracting influence of intercrossing.

In the present state of our knowledge these seem all the conclusions we can arrive at from a study of the New Zealand fauna; but as we fortunately possess a tolerably full and accurate knowledge of the flora of New Zealand, as well as of that of Australia and the south temperate lands generally, it will be well to see how far these conclusions are supported by the facts of plant distribution, and what further indications they afford us of the early history of these most interesting countries. This inquiry is of sufficient importance to occupy a separate chapter.

CHAPTER XXII
THE FLORA OF NEW ZEALAND: ITS AFFINITIES AND PROBABLE ORIGIN

Relations of the New Zealand Flora to that of Australia—General Features of the Australian Flora—The Floras of South-eastern and South-western Australia—Geological Explanation of the Differences of these two Floras—The Origin of the Australian Element in the New Zealand Flora—Tropical Character of the New Zealand Flora Explained—Species Common to New Zealand and Australia mostly Temperate Forms—Why Easily Dispersed Plants have often Restricted Ranges—Summary and Conclusion on the New Zealand Flora.

Although plants have means of dispersal far exceeding those possessed by animals, yet as a matter of fact comparatively few species are carried for very great distances, and the flora of a country taken as a whole usually affords trustworthy indications of its past history. Plants, too, are more numerous in species than the higher animals, and are almost always better known; their affinities have been more systematically studied; and it may be safely affirmed that no explanation of the origin of the fauna of a country can be sound, which does not also explain, or at least harmonise with, the distribution and relations of its flora. The distribution of the two may be very different, but both should be explicable by the same series of geographical changes.

The relations of the flora of New Zealand to that of Australia have long formed an insoluble enigma for botanists. Sir Joseph Hooker, in his most instructive and masterly essay on the flora of Australia, says:—"Under whatever aspect I regard the flora of Australia and of New Zealand, I find all attempts to theorise on the possible causes of their community of feature frustrated by anomalies in distribution, such as I believe no two other similarly situated countries in the globe present. Everywhere else I recognise a parallelism or harmony in the main common features of contiguous floras, which conveys the impression of their generic affinity, at least, being affected by migration from centres of dispersion in one of them, or in some adjacent country. In this case it is widely different. Regarding the question from the Australian point of view, it is impossible in the present state of science to reconcile the fact of Acacia, Eucalyptus, Casuarina, Callitris, &c., being absent in New Zealand, with any theory of transoceanic migration that may be adopted to explain the presence of other Australian plants in New Zealand; and it is very difficult to conceive of a time or of conditions that could explain these anomalies, except by going back to epochs when the prevalent botanical as well as geographical features of each were widely different from what they are now. On the other hand, if I regard the question from the New Zealand point of view, I find such broad features of resemblance, and so many connecting links that afford irresistible evidence of a close botanical connection, that I cannot abandon the conviction that these great differences will present the least difficulties to whatever theory may explain the whole case." I will now state, as briefly as possible, what are the facts above referred to as being of so anomalous a character, and there is little difficulty in doing so, as we have them fully set forth, with admirable clearness, in the essay above alluded to, and in the same writer's Introduction to the Flora of New Zealand, only requiring some slight modifications, owing to the later discoveries which are given in the Handbook of the New Zealand Flora.

Confining ourselves always to flowering plants, we find that the flora of New Zealand is a very poor one, considering the extent of surface, and the favourable conditions of soil and climate. It consists of 1,085 species (our own islands possessing about 1,500), but a very large proportion of these are peculiar, there being no less than 800 endemic species, and thirty-two endemic genera.

Out of the 285 species not peculiar to New Zealand, no less than 215 are Australian, but a considerable number of these are also Antarctic, South American, or European; so that there are only about 100 species absolutely confined to New Zealand and Australia, and, what is important as indicating a somewhat recent immigration, only some half-dozen of these belong to genera which are peculiar to the two countries, and hardly any to the larger and more important Australian genera. Many, too, are rare species in both countries and are often alpines.

Far more important are the relations of the genera and families of the two countries. All the Natural Orders of New Zealand are found in Australia except three—Coriariæ, a widely-scattered group found in South Europe, the Himalayas, and the Andes; Escallonieæ, a widely distributed group; and Chloranthaceæ, found in Tropical Asia, Japan, Polynesia, and South America. Out of a total of 310 New Zealand genera, no less than 248 are Australian, and sixty of these are almost peculiar to the two countries, only thirty-two however being absolutely confined to them.¹⁷⁹ In the three large orders—Compositæ, Orchideæ, and Gramineæ, the genera are almost identical in the two countries, while the species—in the two former especially—are mostly distinct.

Here then we have apparently a wonderful resemblance between the New Zealand flora and that of Australia, indicated by more than two-thirds of the non-peculiar species, and more than nine-tenths of the non-peculiar genera (255) being Australian. But now let us look at the other side of the question.

There are in Australia seven great genera of plants, each containing more than 100 species, all widely spread over the country, and all highly characteristic Australian forms,—Acacia, Eucalyptus, Melaleuca, Leucopogon, Stylidium, Grevillea, and Hakea. These are entirely absent from New Zealand, except one species of Leucopogon, a genus which also has representatives in the Malayan and Pacific Islands. Sixteen more Australian genera have over fifty species each, and of these eight are totally absent from New Zealand, five are represented by one or two species, and only two are fairly represented; but these two—Drosera and Helichrysum—are very widespread genera, and might have reached New Zealand from other countries than Australia.

But this by no means exhausts the differences between New Zealand and Australia. No less than seven Australian Natural Orders—Dilleniaceæ, Buettneriaceæ, Polygaleæ, Tremandreæ, Casuarineæ, Hæmodoraceæ, and Xyrideæ are entirely wanting in New Zealand, and several others which are excessively abundant and highly characteristic of the former country are very poorly represented in the latter. Thus, Leguminosæ are extremely abundant in Australia, where there are over 1,000 species belonging to about 100 genera, many of them altogether peculiar to the country; yet in New Zealand this great order is most scantily represented, there being only five genera and thirteen species; and only two of these genera, Swainsonia and Clianthus, are Australian, and as the latter consists of but two species it may as well have passed from New Zealand to Australia as the other way, or more probably from some third country to them both.¹⁸⁰ Goodeniaceæ with ten genera and 220 species Australian, has but two species in New Zealand—and one of these is a salt-marsh plant found also in Tasmania and in Chile; and four other large Australian orders—Rhamneæ, Myoporineæ, Proteaceæ and Santalaceæ, have very few representatives in New Zealand.

We find, then, that the great fact we have to explain and account for is, the undoubted affinity of the New Zealand flora to that of Australia, but an affinity almost exclusively confined to the least predominant and least peculiar portion of that flora, leaving the most predominant, most characteristic, and most widely distributed portion absolutely unrepresented. We must however be careful not to exaggerate the amount of affinity with Australia, apparently implied by the fact that nearly six-sevenths of the New Zealand genera are also Australian, for, as we have already stated, a very large number of these are European, Antarctic, South American or Polynesian genera, whose presence in the two contiguous areas only indicates a common origin. About one-eighth, only, are absolutely confined to Australia and New Zealand (thirty-two genera), and even of these several are better represented in New Zealand than in Australia, and may therefore have passed from the former to the latter. No less than 174 of the New Zealand genera are temperate South American, many being also Antarctic or European; while others again are especially tropical or Polynesian; yet undoubtedly a larger proportion of the Natural Orders and genera are common to Australia than to any other country, so that we may say that the basis of the flora is Australian with a large intermixture of northern and southern temperate forms and others which have remote world-wide affinities.

General Features of the Australian Flora and its Probable Origin.—Before proceeding to point out how the peculiarities of the New Zealand flora may be best accounted for, it is necessary to consider briefly what are the main peculiarities of Australian vegetation, from which so important a part of that of New Zealand has evidently been derived.

The actual Australian flora consists of two great divisions—a temperate and a tropical, the temperate being again divisible into an eastern and a western portion. All that is most characteristic of the Australian flora belongs to the temperate division (though these often overspread the whole continent), in which are found almost all the remarkable Australian types of vegetation and the numerous genera peculiar to this part of the world. Contrary to what occurs in most other countries, the tropical appears to be less rich in species and genera than the temperate region, and what is still more remarkable it contains fewer peculiar species, and very few peculiar genera. Although the area of tropical Australia is about equal to that of the temperate portions, and it has now been pretty well explored botanically, it has probably not more than half as many species.¹⁸¹ Nearly 500 of its species are identical with Indian or Malayan plants, or are very close representatives of them; while there are more than 200 Indian genera, confined for the most part to the tropical portion of Australia. The remainder of the tropical flora consists of a few species and many genera of temperate Australia which range over the whole continent, but these form only a small portion of the peculiarly Australian genera.

These remarkable facts clearly point to one conclusion—that the flora of tropical Australia is, comparatively, recent and derivative. If we imagine the greater part of North Australia to have been submerged beneath the ocean, from which it rose in the middle or latter part of the Tertiary period, offering an extensive area ready to be covered by such suitable forms of vegetation as could first reach it, something like the present condition of things would inevitably arise. From the north, widespread Indian and Malay plants would quickly enter, while from the south the most dominant forms of warm-temperate Australia, and such as were best adapted to the tropical climate and arid soil, would intermingle with them. Even if numerous islands had occupied the area of Northern Australia for long periods anterior to the final elevation, very much the same state of things would result.

The existence in North and North-east Australia of enormous areas covered with Cretaceous and other Secondary deposits, as well as extensive Tertiary formations, lends support to the view, that during very long epochs temperate Australia was cut off from all close connection with the tropical and northern lands by a wide extent of sea; and this isolation is exactly what was required, in order to bring about the wonderful amount of specialisation and the high development manifested by the typical Australian flora. Before proceeding further, however, let us examine this flora itself, so far as regards its component parts and probable past history.

The Floras of South-eastern and South-western Australia.—The peculiarities presented by the south-eastern and south-western subdivisions of the flora of temperate Australia are most interesting and suggestive, and are, perhaps, unparalleled in any other part of the world. South-west Australia is far less extensive than the south-eastern division—less varied in soil and climate, with no lofty mountains, and much sandy desert; yet, strange to say, it contains an equally rich flora and a far greater proportion of peculiar species and genera of plants. As Sir Joseph Hooker remarks:—"What differences there are in conditions would, judging from analogy with other countries, favour the idea that South-eastern Australia, from its far greater area, many large rivers, extensive tracts of mountainous country and humid forests, would present much the most extensive flora, of which only the drier types could extend into South-western Australia. But such is not the case; for though the far greater area is much the best explored, presents more varied conditions, and is tenanted by a larger number of Natural Orders and genera, these contain fewer species by several hundreds."¹⁸²

The fewer genera of South-western Australia are due almost wholly to the absence of the numerous European, Antarctic, and South-American types found in the south-eastern region, while in purely Australian types it is far the richer, for while it contains most of those found in the east it has a large number altogether peculiar to it; and Sir Joseph Hooker states that "there are about 180 genera, out of 600 in South-western Australia, that are either not found at all in South-eastern, or that are represented there by a very few species only, and these 180 genera include nearly 1,100 species."

Geological Explanation of the Differences of these Two Floras.—These facts again clearly point to the conclusion that South-western Australia is the remnant of the more extensive and more isolated portion of the continent in which the peculiar Australian flora was principally developed. The existence there of a very large area of granite—800 miles in length by nearly 500 in maximum width with detached masses 200 miles to the north and 500 miles to the east—indicates such an extension; for these granitic masses were certainly once buried under piles of stratified rock, since denuded, and then formed the nucleus of the old Western Australian continent. If we take the 1000-fathom line around the southern part of Australia to represent the probable extension of this old land we shall see that it would give a wide additional area south of the Great Australian Bight, and form a continent which, even if the greater part of tropical Australia were submerged, would be sufficient for the development of a peculiar and abundant flora. We must also remember that an elevation of 6000 feet, added to the vast amount which has been taken away by denudation, would change the whole country, including what are now the deserts of the interior, into a mountainous and well-watered region.

But while this rich and peculiar flora was in process of formation, the eastern portion of the continent must either have been widely separated from the western or had perhaps not yet risen from the ocean. The whole of this part of the country consists of Palæozoic and Secondary formations with granite and metamorphic rocks, the Secondary deposits being largely developed on both sides of the central range, extending the whole length of the continent from Tasmania to Cape York, and constituting the greater part of the plateau of the Blue Mountains and other lofty ranges. During some portion of the Secondary and Tertiary periods therefore, this side of Australia must have been almost wholly submerged beneath the ocean; and if we suppose that during this time the western part of the continent was at nearly its maximum extent and elevation, we shall have a sufficient explanation of the great difference between the flora of Western and Eastern Australia, since the latter would only have been able to receive immigrants from the former, at a later period, and in a more or less fragmentary manner.

If we examine the geological map of Australia (given in Stanford's Compendium of Geography and Travel, volume Australasia), we shall see good reason to conclude that the eastern and the western divisions of the country first existed as separate islands, and only became united at a comparatively recent epoch. This is indicated by an enormous stretch of Cretaceous and Tertiary formations extending from the Gulf of Carpentaria completely across the continent to the mouth of the Murray River. During the Cretaceous period, therefore, and probably throughout a considerable portion of the Tertiary epoch,¹⁸³ there must have been a wide arm of the sea occupying this area, dividing the great mass of land on the west—the true seat and origin of the typical Australian flora—from a long but narrow belt of land on the east, indicated by the continuous mass of Secondary and Palæozoic formations already referred to which extend uninterruptedly from Tasmania to Cape York. Whether this formed one continuous land, or was broken up into islands, cannot be positively determined; but the fact that no marine Tertiary beds occur in the whole of this area, renders it probable that it was almost, if not quite, continuous, and that it not improbably extended across to what is now New Guinea. At this epoch, then (as shown in the accompanying map), Australia may, not improbably, have consisted of a very large and fertile western island, almost or quite extratropical, and extending from the Silurian rocks of the Flinders range in South Australia, to about 150 miles west of the present west coast, and southward to about 350 miles south of the Great Australian Bight. To the east of this, at a distance of from 250 to 400 miles, extended in a north and south direction a long but comparatively narrow island, stretching from far south of Tasmania to New Guinea; while the crystalline and Secondary formations of central North Australia probably indicate the existence of one or more large islands in that direction.