The Dispersal of Land Mollusca.The only other group of animals we need now refer to is that of the air-breathing mollusca, commonly called land-shells. These are almost as ubiquitous as insects, though far less numerous; and their wide distribution is by no means so easy to explain. The genera have usually a very wide, and often a cosmopolitan range, while the species are rather restricted, and sometimes wonderfully so. Not only do single islands, however small, often possess peculiar species of land-shells, but sometimes single mountains or valleys, or even a particular mountain side, possess species or varieties found nowhere else upon the globe. It is pretty certain that they have no means of passing over the sea but such as are very rare and exceptional. Some which possess an operculum, or which close the mouth of the shell with a diaphragm of secreted mucus, may float across narrow arms of the sea, especially when protected in the crevices of logs of timber; while in the young state when attached to leaves or twigs they may be carried long distances by hurricanes.15 Owing to their exceedingly slow motion, their powers of voluntary dispersal, even on land, are very limited, and this will explain the extreme restriction of their range in many cases.
Great Antiquity of Land-Shells.The clue to the almost universal distribution of the several families and of many genera, is to be found, however, in their immense antiquity. In the Pliocene and Miocene formations most of the land-shells are either identical with living species or closely allied to them, while even in the Eocene almost all are of living genera, and one British Eocene fossil still lives in Texas. Strange to say, no true land-shells have been discovered in the Secondary formations, but they must certainly have abounded, for in the far more ancient Palæozoic coal measures of Nova Scotia two species belonging to the living genera Pupa and Zonites have been found in considerable abundance.
Land-shells have therefore survived all the revolutions the earth has undergone since Palæozoic times. They have been able to spread slowly but surely into every land that has ever been connected with a continent, while the rare chances of transfer across the ocean, to which we have referred as possible, have again and again occurred during the almost unimaginable ages of their existence. The remotest and most solitary of the islands of the mid-ocean have thus become stocked with them, though the variety of species and genera bears a direct relation to the facilities of transfer, and the shell fauna is never very rich and varied, except in countries which have at one time or other been united to some continental land.
Causes Favouring the Abundance of Land-Shells.The abundance and variety of land-shells is also, more than that of any other class of animals, dependent on the nature of the surface and the absence of enemies, and where these conditions are favourable their forms are wonderfully luxuriant. The first condition is the presence of lime in the soil, and a broken surface of country with much rugged rock offering crevices for concealment and hibernation. The second is a limited bird and mammalian fauna, in which such species as are especially shell-eaters shall be rare or absent. Both these conditions are found in certain large islands, and pre-eminently in the Antilles, which possess more species of land-shells than any single continent. If we take the whole globe, more species of land-shells are found on the islands than on the continentsa state of things to which no approach is made in any other group of animals whatever, but which is perhaps explained by the considerations now suggested.
The Dispersal of Plants.The ways in which plants are dispersed over the earth, and the special facilities they often possess for migration have been pointed out by eminent botanists, and a considerable space might be occupied in giving a summary of what has been written on the subject. In the present work, however, it is only in two or three chapters that I discuss the origin of insular floras in any detail; and it will therefore be advisable to adduce any special facts when they are required to support the argument in particular cases. A few general remarks only will therefore be made here.
Special Adaptability of Seeds for Dispersal.Plants possess many great advantages over animals as regards the power of dispersal, since they are all propagated by seeds or spores, which are hardier than the eggs of even insects, and retain their vitality for a much longer time. Seeds may lie dormant for many years and then vegetate, while they endure extremes of heat, of cold, of drought, or of moisture which would almost always be fatal to animal germs. Among the causes of the dispersal of seeds De Candolle enumerates the wind, rivers, ocean currents, icebergs, birds and other animals, and human agency. Great numbers of seeds are specially adapted for transport by one or other of these agencies. Many are very light, and have winged appendages, pappus, or down, which enable them to be carried enormous distances. It is true, as De Candolle remarks, that we have no actual proofs of their being so carried; but this is not surprising when we consider how small and inconspicuous most seeds are. Supposing every year a million seeds were brought by the wind to the British Isles from the Continent, this would be only ten to a square mile, and the observation of a life-time might never detect one; yet a hundredth part of this number would serve in a few centuries to stock an island like Britain with a great variety of continental plants.
When, however, we consider the enormous quantity of seeds produced by plants, that great numbers of these are more or less adapted to be carried by the wind, and that winds of great violence and long duration occur in most parts of the world, we are as sure that seeds must be carried to great distances as if we had seen them so carried. Such storms carry leaves, hay, dust, and many small objects to a great height in the air, while many insects have been conveyed by them for hundreds of miles out to sea and far beyond what their unaided powers of flight could have effected.
Birds as Agents in the Dispersal of Plants.Birds are undoubtedly important agents in the dispersal of plants over wide spaces of ocean, either by swallowing fruits and rejecting the seeds in a state fit for germination, or by the seeds becoming attached to the plumage of ground-nesting birds, or to the feet of aquatic birds embedded in small quantities of mud or earth. Illustrations of these various modes of transport will be found in Chapter XII. when discussing the origin of the flora of the Azores and Bermuda.
Ocean-currents as Agents in Plant-dispersal.Ocean-currents are undoubtedly more important agents in conveying seeds of plants than they are in the case of any other organisms, and a considerable body of facts and experiments have been collected proving that seeds may sometimes be carried in this way many thousand miles and afterwards germinate. Mr. Darwin made a series of interesting experiments on this subject, some of which will be given in the chapter above referred to.
Dispersal along Mountain Chains.These various modes of transport are, as will be shown when discussing special cases, amply sufficient to account for the vegetation found on oceanic islands, which almost always bears a close relation to that of the nearest continent; but there are other phenomena presented by the dispersal of species and genera of plants over very wide areas, especially when they occur in widely separated portions of the northern and southern hemispheres, that are not easily explained by such causes alone. It is here that transmission along mountain chains has probably been effective; and the exact mode in which this has occurred is discussed in Chapter XXIII., where a considerable body of facts is given, showing that extensive migrations may be effected by a succession of moderate steps, owing to the frequent exposure of fresh surfaces of soil or débris on mountain sides and summits, offering stations on which foreign plants can temporarily establish themselves.
When, however, we consider the enormous quantity of seeds produced by plants, that great numbers of these are more or less adapted to be carried by the wind, and that winds of great violence and long duration occur in most parts of the world, we are as sure that seeds must be carried to great distances as if we had seen them so carried. Such storms carry leaves, hay, dust, and many small objects to a great height in the air, while many insects have been conveyed by them for hundreds of miles out to sea and far beyond what their unaided powers of flight could have effected.
Birds as Agents in the Dispersal of Plants.Birds are undoubtedly important agents in the dispersal of plants over wide spaces of ocean, either by swallowing fruits and rejecting the seeds in a state fit for germination, or by the seeds becoming attached to the plumage of ground-nesting birds, or to the feet of aquatic birds embedded in small quantities of mud or earth. Illustrations of these various modes of transport will be found in Chapter XII. when discussing the origin of the flora of the Azores and Bermuda.
Ocean-currents as Agents in Plant-dispersal.Ocean-currents are undoubtedly more important agents in conveying seeds of plants than they are in the case of any other organisms, and a considerable body of facts and experiments have been collected proving that seeds may sometimes be carried in this way many thousand miles and afterwards germinate. Mr. Darwin made a series of interesting experiments on this subject, some of which will be given in the chapter above referred to.
Dispersal along Mountain Chains.These various modes of transport are, as will be shown when discussing special cases, amply sufficient to account for the vegetation found on oceanic islands, which almost always bears a close relation to that of the nearest continent; but there are other phenomena presented by the dispersal of species and genera of plants over very wide areas, especially when they occur in widely separated portions of the northern and southern hemispheres, that are not easily explained by such causes alone. It is here that transmission along mountain chains has probably been effective; and the exact mode in which this has occurred is discussed in Chapter XXIII., where a considerable body of facts is given, showing that extensive migrations may be effected by a succession of moderate steps, owing to the frequent exposure of fresh surfaces of soil or débris on mountain sides and summits, offering stations on which foreign plants can temporarily establish themselves.
Antiquity of Plants as affecting their Distribution.We have already referred to the importance of great antiquity in enabling us to account for the wide dispersal of some genera and species of insects and land-shells, and recent discoveries in fossil botany show that this cause has also had great influence in the case of plants. Rich floras have been discovered in the Miocene, the Eocene, and the Upper Cretaceous formations, and these consist almost wholly of living genera, and many of them of species very closely allied to existing forms. We have therefore every reason to believe that a large number of our plant-species have survived great geological, geographical, and climatal changes; and this fact, combined with the varied and wonderful powers of dispersal many of them possess, renders it far less difficult to understand the examples of wide distribution of the genera and species of plants than in the case of similar instances among animals. This subject will be further alluded to when discussing the origin of the New Zealand flora, in Chapter XXII.
CHAPTER VI
GEOGRAPHICAL AND GEOLOGICAL CHANGES: THE PERMANENCE OF CONTINENTS
Changes of Land, and Sea, their Nature and ExtentShore-deposits and Stratified RocksThe Movements of ContinentsSupposed Oceanic Formations; the Origin of ChalkFresh-water and Shore-deposits as proving the Permanence of ContinentsOceanic Islands as indications of the Permanence of Continents and OceansGeneral Stability of Continents with constant Change of FormEffect of Continental Changes on the Distribution of AnimalsChanged Distribution proved by the Extinct Animals of Different EpochsSummary of Evidence for the general Permanence of Continents and Oceans.
The changes of land and sea which have occurred in particular cases will be described when we discuss the origin and relations of the faunas of the different classes of islands. We have here only to consider the general character and extent of such changes, and to correct some erroneous ideas which are prevalent on the subject.
Changes of Land and Sea, their Nature and Extent.It is a very common belief that geological evidence proves a complete change of land and sea to have taken place over and over again. Every foot of dry land has undoubtedly, at one time or other, formed part of a sea-bottom, and we can hardly exclude the surfaces occupied by volcanic and fresh-water deposits, since, in many cases, if not in all, these rest upon a substratum of marine formations. At first sight, therefore, it seems a necessary inference that when the present continents were under water there must have been other continents situated where we now find the oceans, from which the sediments came to form the various deposits we now see. This view was held by so acute and learned a geologist as Sir Charles Lyell, who says:"Continents, therefore, although permanent for whole geological epochs, shift their positions entirely in the course of ages."16 Mr. T. Mellard Reade, late President of the Geological Society of Liverpool, so recently as 1878, says:"While believing that the ocean-depths are of enormous age, it is impossible to resist other evidences that they have once been land. The very continuity of animal and vegetable life on the globe points to it. The molluscous fauna of the eastern coast of North America is very similar to that of Europe, and this could not have happened without littoral continuity, yet there are depths of 1,500 fathoms between these continents."17 It is certainly strange that a geologist should not remember the recent and long-continued warm climates of the Arctic regions, and see that a connection of Northern Europe by Iceland with Greenland and Labrador over a sea far less than a thousand fathoms deep would furnish the "littoral continuity" required. Again, in the same pamphlet Mr. Reade says:"It can be mathematically demonstrated that the whole, or nearly the whole, of the sea-bottom has been at one time or other dry land. If it were not so, and the oscillations, of the level of the land with respect to the sea were confined within limits near the present continents, the results would have been a gradual diminution instead of development of the calcareous rocks. To state the case in common language, the calcareous portion of the rocks would have been washed out during the mutations, the destruction and redeposit of the continental rocks, and eventually deposited in the depths of the immutable sea far from land. Immense beds of limestone would now exist at the bottom of the ocean, while the land would be composed of sandstones and argillaceous shales. The evidence of chemistry thus confirms the inductions drawn from the distribution of animal life upon the globe."
So far from this being a "mathematical demonstration," it appears to me to be a complete misinterpretation of the facts. Animals did not create the lime which they secrete from the sea-water, and therefore we have every reason to believe that the inorganic sources which originally supplied it still keep up that supply, though perhaps in diminished quantity. Again, the great lime-secreterscoralswork in water of moderate depth, that is, near land, while there is no proof whatever that there is any considerable accumulation of limestone at the bottom of the deep ocean. On the contrary, the fact ascertained by the Challenger, that beyond a certain depth the "calcareous" ooze ceases, and is replaced by red and grey clays, although the calcareous organisms still abound in the surface waters of the ocean, shows that the lime is dissolved again by the excess of carbonic acid usually found at great depths, and its accumulation thus prevented. As to the increase of limestones in recent as compared with older formations, it may be readily explained by two considerations: in the first place, the growth and development of the land in longer and more complex shore lines and the increase of sedimentary over volcanic formations may have offered more stations favourable to the growth of coral; while the solubility of limestone in rain-water renders the destruction of such rocks more rapid than that of sandstones and shales, and would thus, by supplying more calcareous matter in solution for secretion by limestone-forming organisms, lead to their comparative abundance in later as compared with earlier formations.