# The Geographical Distribution of Animals/Chapter 2

CHAPTER II.
THE MEANS OF DISPERSAL AND THE MIGRATIONS OF ANIMALS.
All animals are capable of multiplying so rapidly, that if a single pair were placed in a continent with abundance of food and no enemies, they might fully stock it in a very short time. Thus, a bird which produces ten pairs of young during its lifetime (and this is far below the fertility of many birds) will, if we take its life at five years, increase to a hundred millions in about forty years, a number sufficient to stock a large country. Many fishes and insects are capable of multiplying several thousandfold each year, so that in a few years they would reach billions and trillions. Even large and slow breeding mammals, which have only one at a birth but continue to breed from eight to ten successive years, may increase from a single pair to ten millions in less than forty years.
But as animals rarely have an unoccupied country to breed in, and as the food in any one district is strictly limited, their natural tendency is to roam in every direction in search of fresh pastures, or new hunting grounds. In doing so, however, they meet with many obstacles. Rocks and mountains have to be climbed, rivers or marshes to be crossed, deserts or forests to be traversed; while narrow straits or wider arms of the sea separate islands from the main land or continents from each other. We have now to inquire what facilities the different classes of animals have for overcoming these obstacles, and what kind of barriers are most effectual in checking their progress.
Means of Dispersal of Mammalia.—Many of the largest mammalia are able to roam over whole continents and are hardly ​stopped by any physical obstacles. The elephant is almost equally at home on plains and mountains, and it even climbs to the highest summit of Adam's Peak in Ceylon, which is so steep and rocky as to be very difficult of ascent for man. It traverses rivers with great ease and forces its way through the densest jungle. There seems therefore to be no limit to its powers of wandering, but the necessity of procuring food and its capacity of enduring changes of climate. The tiger is another animal with great powers of dispersal. It crosses rivers and sometimes even swims over narrow straits of the sea, and it can endure the severe cold of North China and Tartary as well as the heats of the plains of Bengal. The rhinoceros, the lion, and many of the ruminants have equal powers of dispersal; so that wherever there is land and sufficient food, there are no limits to their possible range. Other groups of animals are more limited in their migrations. The apes, lemurs, and many monkeys are so strictly adapted to an arboreal life that they can never roam far beyond the limits of the forest vegetation. The same may be said of the squirrels, the opossums, the arboreal cats, and the sloths, with many other groups of less importance. Deserts or open country are equally essential to the existence of others. The camel, the hare, the zebra, the giraffe and many of the antelopes could not exist in a forest country any more than could the jerboas or the prairie marmots.
There are other animals which are confined to mountains, and could not extend their range into lowlands or forests. The goats and the sheep are the most striking group of this kind, inhabiting many of the highest mountains of the globe; of which the European ibex and mouflon are striking examples. Rivers are equally necessary to the existence of others, as the beaver, otter, water-vole and capybara; and to such animals high mountain-ranges or deserts must form an absolutely impassable barrier.
Climate as a Limit to the Range of Mammals.—Climate appears to limit the range of many animals, though there is some reason to believe that in many cases it is not the climate itself so much as the change of vegetation consequent on climate which produces the effect. The quadrumana appear to be limited by climate, ​since they inhabit almost all the tropical regions but do not range more than about 10° beyond the southern and 12° beyond the northern tropic, while the great bulk of the species are found only within an equatorial belt about 30° wide. But as these animals are almost exclusively fruit-eaters, their distribution depends as much on vegetation as on temperature; and this is strikingly shown by the fact that the Semnopithecus schistaceus inhabits the Himalayan mountains to a height of 11,000 feet, where it has been seen leaping among fir-trees loaded with snow-wreaths! Some northern animals are bounded by the isothermal of 32°. Such are the polar bear and the walrus, which cannot live in a state of nature far beyond the limits of the frozen ocean; but as they live in confinement in temperate countries, their range is probably limited by other conditions than temperature.
We must not therefore be too hasty in concluding, that animals which we now see confined to a very hot or a very cold climate are incapable of living in any other. The tiger was once considered a purely tropical animal, but it inhabits permanently the cold plains of Manchuria and the Amoor, a country of an almost arctic winter climate. Few animals seem to us more truly inhabitants of hot countries than the elephants and rhinoceroses; yet in Post-tertiary times they roamed over the whole of the northern continents to within the arctic circle; and we know that the climate was then as cold as it is now, from their entire bodies being preserved in ice. Some change must recently have occurred either in the climate, soil, or vegetation of Northern Asia which led to the extinction of these forerunners of existing tropical species; and we must always bear in mind that similar changes may have acted upon other species which we now find restricted within narrow limits, but which may once have roamed over a wide and varied territory.
Valleys and Rivers as Barriers to Mammals.—To animals which thrive best in dry and hilly regions, a broad level and marshy valley must often prove an effectual barrier. The difference of vegetation and of insect life, together with an unhealthy atmosphere, no doubt often checks migration if it is attempted. Thus ​many animals are restricted to the slopes of the Himalayas or to the mountains of Central India, the flat valley of the Ganges forming a limit to their range. In other cases, however, it is the river rather than the valley which is the barrier. In the great Amazonian plains many species of monkeys, birds, and even insects are found up to the river banks on one side but do not cross to the other. Thus in the lower part of the Rio Negro two monkeys, the Jacchus bicolor and the Brachiurus couxiou, are found on the north bank of the river but never on the south, where a red-whiskered Pithecia is alone found. Higher up Ateles paniscus extends to the north bank of the river while Lagothrix humboldtii comes down to the south bank; the former being a native of Guiana, the latter of Ecuador. The range of the birds of the genus Psophia or trumpeters, is also limited by the rivers Amazon, Madeira, Rio Negro and some others; so that in these cases we are able to define the limits of distribution with an unusual degree of accuracy, and there is little doubt the same barriers also limit a large number of other species.
Arms of the Sea as Barriers to Mammals.—Very few mammals can swim over any considerable extent of sea, although many can swim well for short distances. The jaguar traverses the widest streams in South America, and the bear and bison cross the Mississippi; and there can be no doubt that they could swim over equal widths of salt water, and if accidentally carried out to sea might sometimes succeed in reaching islands many miles distant. Contrary to the common notion pigs can swim remarkably well. Sir Charles Lyell tells us in his "Principles of Geology" that during the floods in Scotland in 1829, some pigs only six months old that were carried out to sea, swam five miles and got on shore again. He also states, on the authority of the late Edward Forbes, that a pig jumped overboard to escape from a terrier in the Grecian Archipelago, and swam safely to shore many miles distant. These facts render it probable that wild pigs, from their greater strength and activity, might under favourable circumstances cross arms of the sea twenty or thirty miles wide; and there are facts in the distribution of this tribe of animals which seem to indicate that they have sometimes done so. Deer ​take boldly to the water and can swim considerable distances, but we have no evidence to show how long they could live at sea or how many miles they could traverse. Squirrels, rats, and lemmings often migrate from northern countries in bands of thousands and hundreds of thousands, and pass over rivers, lakes and even arms of the sea, but they generally perish in the saltwater. Admitting, however, the powers of most mammals to swim considerable distances, we have no reason to believe that any of them could traverse without help straits of upwards of twenty miles in width, while in most cases a channel of half that distance would prove an effectual barrier.
Ice-floes and Driftwood as Aiding the Dispersal of Mammals.—In the arctic regions icebergs originate in glaciers which descend into the sea, and often bear masses of gravel, earth, and even some vegetation on their surfaces; and extensive level ice-fields break away and float southwards. These might often carry with them such arctic quadrupeds as frequent the ice, or even on rare occasions true land-animals, which might sometimes be stranded on distant continents or islands. But a more effectual because a more wide-spread agent, is to be found in the uprooted trees and rafts of driftwood often floated down great rivers and carried out to sea. Such rafts or islands are sometimes seen drifting a hundred miles from the mouth of the Ganges with living trees erect upon them; and the Amazon, the Orinoco, Mississippi, Congo, and most great rivers produce similar rafts. Spix and Martius declare that they saw at different times on the Amazon, monkeys, tiger-cats, and squirrels, being thus carried down the stream. On the Parana, pumas, squirrels, and many other quadrupeds have been seen on these rafts; and Admiral W. H. Smyth informed Sir C. Lyell that among the Philippine islands after a hurricane, he met with floating masses of wood with trees growing upon them, so that they were at first mistaken for islands till it was found that they were rapidly drifting along. Here therefore, we have ample means for carrying all the smaller and especially the arboreal mammals out to sea; and although in most cases they would perish there, yet in some favourable instances strong winds or ​unusual tidal currents might carry them safely to shores perhaps several hundred miles from their native country. The fact of green trees so often having been seen erect on these rafts is most important; for they would act as a sail by which the raft might he propelled in one direction for several days in succession, and thus at last reach a shore to which a current alone would never have carried it.
There are two groups of mammals which have quite exceptional means of dispersal—the bats which fly, and the cetacea, seals, &c., which swim. The former are capable of traversing considerable spaces of sea, since two North American species either regularly or occasionally visit the Bermudas, a distance of 600 miles from the mainland. The oceanic mammals (whales and porpoises) seem to have no barrier but temperature; the polar species being unable to cross the equator, while the tropical forms are equally unfitted for the cold polar waters. The shore-feeding manatees, however, can only live where they find food; and a long expanse of rocky coast would probably be as complete a barrier to them as a few hundred miles of open ocean. The amphibious seals and walruses seem many of them to be capable of making long sea journeys, some of the species being found on islands a thousand miles apart, but none of the arctic are identical with the antarctic species.
The otters with one exception are freshwater animals, and we have no reason to believe they could or would traverse any great distances of salt water. In fact, they would be less liable to dispersal across arms of the sea than purely terrestrial species, since their powers of swimming would enable them to regain the shore if accidentally carried out to sea by a sudden flood.
Means of Dispersal of Birds.—It would seem at first sight that no barriers could limit the range of birds, and that they ought to be the most ubiquitous of living things, and little fitted therefore to throw any light on the laws or causes of the geographical distribution of animals. This, however, is far from being the case; many groups of birds are almost as strictly limited by barriers as the mammalia; and from their larger numbers and the avidity with which they have been collected, they furnish ​materials of the greatest value for our present study. The different groups of birds offer remarkable contrasts in the extent of their range, some being the most cosmopolite of the higher animals, while others are absolutely confined to single spots on the earth's surface. The petrels (Procellariidæ) and the gulls (Laridæ) are among the greatest wanderers; but most of the species are confined to one or other of the great oceans, or to the arctic or antarctic seas, a few only being found with scarcely any variation over almost the whole globe. The sandpipers and plovers wander along the shores as far as do the petrels over the ocean. Great numbers of them breed in the arctic regions and migrate as far as India and Australia, or down to Chili and Brazil; the species of the old and new worlds, however, being generally distinct. In striking contrast to these wide ranges we find many of the smaller perching birds, with some of the parrots and pigeons, confined to small islands of a few square miles in extent, or to single valleys or mountains on the mainland.
Dispersal of Birds by Winds.—Those groups of birds which possess no powers of flight, such as the ostrich, cassowary, and apteryx, are in exactly the same position as mammalia as regards their means of dispersal, or are perhaps even inferior to them; since, although they are able to cross rivers by swimming, it is doubtful if they could remain so long in the water as most land quadrupeds. A very large number of short-winged birds, such as toucans, pittas, and wrens, are perhaps worse off; for they can fly very few miles at a time, and on falling into the water would soon be drowned. It is only the strong-flying species that can venture to cross any great width of sea; and even these rarely do so unless compelled by necessity to migrate in search of food, or to a more genial climate. Small and weak birds are, however, often carried accidentally across great widths of ocean by violent gales. This is well exemplified by the large numbers of stragglers from North America, which annually reach the Bermudas. No less than sixty-nine species of American birds have occurred in Europe, most of them in Britain and Heligoland. They consist chiefly of migratory birds which in autumn ​return along the eastern coasts of the United States, and often fly from point to point across bays and inlets. They are then liable to be blown out to sea by storms, which are prevalent at this season; and it is almost always at this time of year that their occurrence has been noted on the shores of Europe. It may, however, be doubted whether this is not an altogether modern phenomenon, dependent on the number of vessels constantly on the Atlantic which afford resting-places to the wanderers; as it is hardly conceivable that such birds as titlarks, cuckoos, wrens, warblers, and rails, could remain on the wing without food or rest, the time requisite to pass over 2,000 miles of ocean. It is somewhat remarkable that no European birds reach the American coast but a few which pass by way of Iceland and Greenland; whereas a considerable number do reach the Azores, fully half way across; so that their absence can hardly be due to the prevailing winds being westerly. The case of the Azores is, however, an argument for the unassisted passage of birds for that distance; since two of the finches are peculiar 'species,' but closely allied to European forms, so that their progenitors must, probably, have reached the islands before the Atlantic was a commercial highway.
Barriers to the Dispersal of Birds.—We have seen that, as a rule, wide oceans are an almost absolute barrier to the passage of most birds from one continent to another; but much narrower seas and straits are also very effectual barriers where the habits of the birds are such as to preserve them from being carried away by storms. All birds which frequent thickets and forests, and which feed near or on the ground, are secure from such accidents; and they are also restricted in their range by the extent of the forests they inhabit. In South America a large number of the birds have their ranges determined by the extent of the forest country, while others are equally limited to the open plains. Such species are also bounded by mountain ranges whenever these rise above the woody region. Great rivers, such as the Amazon, also limit the range of many birds, even when there would seem to be no difficulty in their crossing them. The supply of food, and the kind of vegetation, soil, and climate ​best suited to a bird's habits, are probably the causes which mark out the exact limits of the range of each species; to which must be added the prevalence of enemies of either the parent birds, the eggs, or the young. In the Malay Archipelago pigeons abound most where monkeys do not occur; and in South America the same birds are comparatively scarce in the forest plains where monkeys are very abundant, while they are plentiful on the open plains and campos, and on the mountain plateaux, where these nest-hunting quadrupeds are rarely found. Some birds are confined to swamps, others to mountains; some can only live on rocky streams, others on deserts or grassy plains.
The Phenomena of Migration.—The term "migration" is often applied to the periodical or irregular movements of all animals; but it may be questioned whether there are any regular migrants but birds and fishes. The annual or periodical movements of mammalia are of a different class. Monkeys ascend the Himalayas in summer to a height of 10,000 to 12,000 feet, and descend again in winter. Wolves everywhere descend from the mountains to the lowlands in severe weather. In dry seasons great herds of antelopes move southwards towards the Cape of Good Hope. The well-known lemmings, in severe winters, at long intervals, move down from the mountains of Scandinavia in immense numbers, crossing lakes and rivers, eating their way through haystacks, and surmounting every obstacle till they reach the sea, whence very few return. The alpine hare, the arctic fox, and many other animals, exhibit similar phenomena on a smaller scale; and generally it may be said, that whenever a favourable succession of seasons has led to a great multiplication of any species, it must on the pressure of hunger seek food in fresh localities. For such movements as these we have no special term. The summer and winter movements best correspond to true migration, but they are always on a small scale, and of limited extent; the other movements are rather temporary incursions than true migrations.
The annual movements of many fishes are more strictly analogous to the migration of birds, since they take place in large bodies and often to considerable distances, and are ​immediately connected with the process of reproduction. Some, as the salmon, enter rivers; others, as the herring and mackerel, approach the coast in the breeding season; but the exact course of their migrations is unknown, and owing to our complete ignorance of the area each species occupies in the ocean, and the absence of such barriers and of such physical diversities as occur on the land, they are of far less interest and less connected with our present study than the movements of birds, to which we shall now confine ourselves.
Migrations of Birds.—In all the temperate parts of the globe there are a considerable number of birds which reside only a part of the year, regularly arriving and leaving at tolerably fixed epochs. In our own country many northern birds visit us in winter, such as the fieldfare, redwing, snow-bunting, turnstone, and numerous ducks and waders; with a few, like the black redstart, and (according to Rev. C. A. Johns) some of the woodcocks from the south. In the summer a host of birds appear—the cuckoo, the swifts and swallows, and numerous warblers, being the most familiar,—which stay to build their nests and rear their young, and then leave us again. These are true migrants; but a number of other birds visit us occasionally, like the waxwing, the oriole, and the beefeater, and can only be classed as stragglers, which, perhaps from too rapid multiplication one year and want of food the next, are driven to extend their ordinary range of migration to an unusual degree. We will now endeavour to sketch the chief phenomena of migration in different countries.
Europe.—It is well ascertained that most of the birds that spend their spring and summer in the temperate parts of Europe pass the winter in North Africa and Western Asia. The winter visitants, on the other hand, pass the summer in the extreme north of Europe and Asia, many of them having been found to breed in Lapland. The arrival of migratory birds from the south is very constant as to date, seldom varying more than a week or two, without any regard to the weather at the time; but the departure is less constant, and more dependent on the weather. Thus the swallow always comes to us about the middle ​of April, however cold it may be, while its departure may take place from the end of September to late in October, and is said by Forster to occur on the first N. or N.E. wind after the 20th of September.
Almost all the migratory birds of Europe go southward to the Mediterranean, move along its coasts east or west, and cross over in three places only; either from the south of Spain, in the neighbourhood of Gibraltar, from Sicily over Malta, or to the east by Greece and Cyprus. They are thus always in sight of land. The passage of most small birds (and many of the larger ones too) takes place at night; and they only cross the Mediterranean when the wind is steady from near the east or west, and when there is moonlight.
It is a curious fact, but one that seems to be well authenticated, that the males often leave before the females, and both before the young birds, which in considerable numbers migrate later and alone. These latter, however, seldom go so far as the old ones; and numbers of young birds do not cross the Mediterranean, but stay in the south of Europe. The same rule applies to the northward migration; the young birds stopping short of the extreme arctic regions, to which the old birds migrate. When old and young go together, however, the old birds take the lead. In the south of Europe few of the migratory birds stay to breed, but pass on to more temperate zones; thus, in the south of France, out of 350 species only 60 breed there. The same species is often sedentary in one part of Europe and migratory in another; thus, the chaffinch is a constant resident in England, Germany, and the middle of France; but a migrant in the south of France and in Holland: the rook visits the south of France in winter only: the Falco tinnunculus is both a resident and a migrant in the south of France, according to M. Marcel de Serres, there being two regular passages every year, while a certain number always remain.
​We see, then, that migration is governed by certain intelligible laws; and that it varies in many of its details, even in the same species, according to changed conditions. It may be looked upon as an exaggeration of a habit common to all locomotive animals, of moving about in search of food. This habit is greatly restricted in quadrupeds by their inability to cross the sea or even to pass through the highly-cultivated valleys of such countries as Europe; but the power of flight in birds enables them to cross every kind of country, and even moderate widths of sea; and as they mostly travel at night and high in the air, their movements are difficult to observe, and are supposed to be more mysterious than they perhaps are. In the tropics birds move about to different districts according as certain fruits become ripe, certain insects abundant, or as flooded tracts dry up. On the borders of the tropics and the temperate zone extends a belt of country of a more or less arid character, and liable to be parched at the summer solstice. In winter and early spring its northern margin is verdant, but it soon becomes burnt up, and most of its birds necessarily migrate to the more fertile regions to the north of them. They thus follow the spring or summer as it advances from the south towards the pole, feeding on the young flower buds, the abundance of juicy larvæ, and on the ripening fruits; and as soon as these become scarce they retrace their steps homewards to pass the winter. Others whose home is nearer the pole are driven south by cold, hunger, and darkness, to more hospitable climes, returning northward in the early summer. As a typical example of a migratory bird, let us take the nightingale. During the winter this bird inhabits almost all North Africa, Asia Minor, and the Jordan Valley. Early in April it passes into Europe by the three routes already mentioned, and spreads over France, Britain, Denmark, and the south of Sweden, which it reaches by the beginning of May. It does not enter Brittany, the Channel Islands, or the western part of England, never visiting Wales, except the extreme south of Glamorganshire, and rarely extending farther north than Yorkshire. It spreads over Central Europe, through Austria and Hungary to Southern Russia and the warmer parts of Siberia, ​but it nevertheless breeds in the Jordan Valley, so that in some places it is only the surplus population that migrates. In August and September, all who can return to their winter quarters.
Migrations of this type probably date back from at least the period when there was continuous land along the route passed over; and it is a suggestive fact that this land connection is known to have existed in recent geological times. Britain was connected with the Continent during, and probably before, the glacial epoch; and Gibraltar, as well as Sicily and Malta, were also recently united with Africa, as is proved by the fossil elephants and other large mammalia found in their caverns, by the comparatively shallow water still existing in this part of the Mediterranean while the remainder is of oceanic profundity, and by the large amount of identity in the species of land animals still inhabiting the opposite shores of the Mediterranean. The submersion of these two tracts of land (which were perhaps of considerable extent) would be a slow process, and from year to year the change might be hardly perceptible. It is easy to see how the migration that had once taken place over continuous land would be kept up, first over lagoons and marshes, then over a narrow channel, and subsequently over a considerable sea, no one generation of birds ever perceiving any difference in the route.
There is, however, no doubt that the sea-passage is now very dangerous to many birds. Quails cross in immense flocks, and great numbers are drowned at sea whenever the weather is unfavourable. Some individuals always stay through the winter in the south of Europe, and a few even in England and Ireland; and were the sea to become a little wider the migration would cease, and the quail, like some other birds, would remain divided between south Europe and north Africa. Aquatic birds are observed to follow the routes of great rivers and lakes, and the shores of the sea. One great body reaches central Europe by way of the Danube from the shores of the Black Sea; another ascends the Rhone Valley from the Gulf of Lyons.
​India and China.—In the peninsula of India and in China great numbers of northern birds arrive during September and October, and leave from March to May. Among the smaller birds are wagtails, pipits, larks, stonechats, warblers, thrushes, buntings, shrikes, starlings, hoopoes, and quails. Some species of cranes and storks, many ducks, and great numbers of Scolopacidæ also visit India in winter; and to prey upon these come a band of rapacious birds—the peregrine falcon, the hobby, kestrel, common sparrowhawk, harrier, and the short-eared owl. These birds are almost all natives of Europe and Western Asia; they spread over all northern and central India, mingling with the sedentary birds of the oriental fauna, and give to the ornithology of Hindostan at this season quite a European aspect. The peculiar species of the higher Himalayas do not as a rule descend to the plains in winter, but merely come lower down the mountains; and in southern India and Ceylon comparatively few of these migratory birds appear.
In China the migratory birds follow generally the coast line, coming southwards in winter from eastern Siberia and northern Japan; while a few purely tropical forms travel northwards in summer to Japan, and on the mainland as far as the valley of the Amoor.
North America.—The migrations of birds in North America have been carefully studied by resident naturalists, and present some interesting features. The birds of the eastern parts of North America are pre-eminently migratory, a much smaller proportion being permanent residents than in corresponding latitudes in Europe. Thus, in Massachusetts there are only about 30 species of birds which are resident all the year, while the regular summer visitors are 106. Comparing with this our own country, though considerably further north, the proportions are reversed; there being 140 residents and 63 summer visitors. This difference is clearly due to the much greater length and severity of the winter, and the greater heat of summer, in America than with us. The number of permanent residents increases pretty regularly as we go southward; but the number of birds at any locality during the breeding season seems to increase as we go ​northward as far as Canada, where, according to Mr. Allen, more species breed than in the warm Southern States. Even in the extreme north, beyond the limit of forests, there are no less than 60 species which breed; in Canada about 160; while in Carolina there are only 135, and in Louisiana, 130. The extent of the migration varies greatly, some species only going a few degrees north and south, while others migrate annually from the tropics to the extreme north of the continent; and every gradation occurs between these extremes. Among those which migrate furthest are the species of Dendrœca, and other American flycatching warblers (Mniotiltidæ), many of which breed on the shores of Hudson's Bay, and spend the winter in Mexico or the West Indian islands.
The great migratory movement of American birds is almost wholly confined to the east coast; the birds of the high central plains and of California being for the most part sedentary, or only migrating for short distances. All the species which reach South America, and most of those which winter in Mexico and Guatemala, are exclusively eastern species; though a few Rocky Mountain birds range southward along the plateaux of Mexico and Guatemala, but probably not as regular annual migrants.
In America as in Europe birds appear in spring with great regularity, while the time of the autumnal return is less constant. More curious is the fact, also observed in both hemispheres, that they do not all return by the same route followed in going northwards, some species being constant visitors to certain localities in spring but not in autumn, others in autumn but not in spring.
Some interesting cases have been observed in America of a gradual alteration in the extent of the migration of certain birds. A Mexican swallow (Hirundo lunifrons) first appeared in Ohio in 1815. Year by year it increased the extent of its range till by 1845 it had reached Maine and Canada; and it is now quoted by American writers as extending its annual migrations to Hudson's Bay. An American wren (Troglodytes ludovicianus) is another bird which has spread considerably northwards since ​the time of the ornithologist Wilson; and the rice-bird, or "Bob-o'-link," of the Americans, continually widens its range as rice and wheat are more extensively cultivated. This bird winters in Cuba and other West Indian Islands, and probably also in Mexico. In April it enters the Southern States and passes northward, till in June it reaches Canada and extends west to the Saskatchewan River in 54° north latitude.
South Temperate America.—The migratory birds of this part of the world have been observed by Mr. Hudson at Buenos Ayres. As in Europe and North America, there are winter and summer visitors, from Patagonia and the tropics respectively. Species of Pyrocephalus, Milvulus, swallows, and a hummingbird, are among the most regular of the summer visitors. They are all insectivorous birds. From Patagonia species of Tænioptera, Cinclodes, and Centrites, come in winter, with two gulls, two geese, and six snipes and plovers. Five species of swallows appear at Buenos Ayres in spring, some staying to breed, others passing on to more temperate regions farther south. As a rule the birds which come late and leave early are the most regular. Some are very irregular in their movements, the Molothrus bonariensis, for example, sometimes leaves early in autumn, sometimes remains all the winter. Some resident birds also move in winter to districts where they are never seen in summer.
General Remarks on Migration.—The preceding summary of the main facts of migration (which might have been almost indefinitely extended, owing to the great mass of detailed information that exists on the subject) appears to accord with the view already suggested, that the "instinct" of migration has arisen from the habit of wandering in search of food common to all animals, but greatly exaggerated in the case of birds by their powers of flight and by the necessity for procuring a large amount of soft insect food for their unfledged young. Migration in its simple form may be best studied in North America, where it takes place over a continuous land surface with a considerable change of climate from south to north. We have here (as probably in Europe and elsewhere) every grade of migration, from species which merely shift the northern and southern ​limits of their range a few hundred miles, so that in the central parts of the area the species is a permanent resident, to others which move completely over 1,000 miles of latitude, so that in all the intervening districts they are only known as birds of passage. Now, just as the rice-bird and the Mexican swallow have extended their migrations, owing to favourable conditions induced by human agency; so we may presume that large numbers of species would extend their range where favourable conditions arose through natural causes. If we go back only as far as the height of the glacial epoch, there is reason to believe that all North America, as far south as about 40° north latitude, was covered with an almost continuous and perennial ice-sheet. At this time the migratory birds would extend up to this barrier (which would probably terminate in the midst of luxuriant vegetation, just as the glaciers of Switzerland now often terminate amid forests and corn-fields), and as the cold decreased and the ice retired almost imperceptibly year by year, would follow it up farther and farther according as the peculiarities of vegetation and insect-food were more or less suited to their several constitutions. It is an ascertained fact that many individual birds return year after year to build their nests in the same spot. This shows a strong local attachment, and is, in fact, the faculty or feeling on which their very existence probably depends. For were they to wander at random each year, they would almost certainly not meet with places so well suited to them, and might even get into districts where they or their young would inevitably perish. It is also a curious fact that in so many cases the old birds migrate first, leaving the young ones behind, who follow some short time later, but do not go so far as their parents. This is very strongly opposed to the notion of an imperative instinct. The old birds have been before, the young have not; and it is only when the old ones have all or nearly all gone that the young go too, probably following some of the latest stragglers. They wander, however, almost at random, and the majority are destroyed before the next spring. This is proved by the fact that the birds which return in spring are as a rule not more numerous than those which came the ​preceding spring, whereas those which went away in autumn were two or three times as numerous. Those young birds that do get back, however, have learnt by experience, and the next year they take care to go with the old ones. The most striking fact in favour of the "instinct" of migration is the "agitation," or excitement, of confined birds at the time when their wild companions are migrating. It seems probable, however, that this is what may be called a social excitement, due to the anxious cries of the migrating birds; a view supported by the fact stated by Marcel de Serres, that the black swan of Australia, when domesticated in Europe, sometimes joins wild swans in their northward migration. We must remember too that migration at the proper time is in many cases absolutely essential to the existence of the species; and it is therefore not improbable that some strong social emotion should have been gradually developed in the race, by the circumstance that all who for want of such emotion did not join their fellows inevitably perished.
The mode by which a passage originally overland has been converted into one over the sea offers no insuperable difficulties, as has already been pointed out. The long flights of some birds without apparently stopping on the way is thought to be inexplicable, as well as their finding their nesting-place of the previous year from a distance of many hundreds or even a thousand miles. But the observant powers of animals are very great; and birds flying high in the air may be guided by the physical features of the country spread out beneath them in a way that would be impracticable to purely terrestrial animals.
It is assumed by some writers that the breeding-place of a species is to be considered as its true home rather than that to which it retires in winter; but this can hardly be accepted as a rule of universal application. A bird can only breed successfully where it can find sufficient food for its young; and the reason probably why so many of the smaller birds leave the warm southern regions to breed in temperate or even cold latitudes, is because caterpillars and other soft insect larvæ are there abundant at the proper time, while in their winter home the ​larvæ have all changed into winged insects. But this favourable breeding district will change its position with change of climate; and as the last great change has been one of increased warmth in all the temperate zones, it is probable that many of the migratory birds are comparatively recent visitors. Other changes may however have taken place, affecting the vegetation and consequently the insects of a district; and we have seldom the means of determining in any particular case in what direction the last extension of range occurred. For the purposes of the study of geographical distribution therefore, we must, except in special cases, consider the true range of a species to comprise all the area which it occupies regularly for any part of the year, while all those districts which it only visits at more or less distant intervals, apparently driven by storms or by hunger, and where it never regularly or permanently settles, should not be included as forming part of its area of distribution.
Means of Dispersal of Reptiles and Amphibia.—If we leave out of consideration the true marine groups—the turtles and sea-snakes—reptiles are scarcely more fitted for traversing seas and oceans than are mammalia. We accordingly find that in those oceanic islands which possess no indigenous mammals, land reptiles are also generally wanting. The several groups of these animals, however, differ considerably both in their means of dispersal and in their power of resisting adverse conditions. Snakes are most dependent on climate, becoming very scarce in temperate and cold climates and entirely ceasing at 62° north latitude, and they do not ascend very lofty mountains, ceasing at 6,000 feet elevation in the Alps. Some inhabit deserts, others swamps and marshes, while many are adapted for a life in forests. They swim rivers easily, but apparently have no means of passing the sea, since they are very rarely found on oceanic islands. Lizards are also essentially tropical, but they go somewhat farther north than snakes, and ascend higher on the mountains, reaching 10,000 feet in the Alps. They possess too some unknown means (probably in the egg-state) of passing over the ocean, since they are found to inhabit many islands where there are neither mammalia nor snakes.
​The amphibia are much less sensitive to cold than are true reptiles, and they accordingly extend much farther north, frogs being found within the arctic circle. Their semi-aquatic life also gives them facilities for dispersal, and their eggs are no doubt sometimes carried by aquatic birds from one pond or stream to another. Salt water is fatal to them as well as to their eggs, and hence it arises that they are seldom found in those oceanic islands from which mammalia are absent. Deserts and oceans would probably form the most effectual barriers to their dispersal; whereas both snakes and lizards abound in deserts, and have some means of occasionally passing the ocean which frogs and salamanders do not seem to possess.
Means of Dispersal of Fishes.—The fact that the same species of freshwater fish often inhabit distinct river systems, proves that they have some means of dispersal over land. The many authentic accounts of fish falling from the atmosphere, indicate one of the means by which they may be transferred from one river basin to another, viz., by hurricanes and whirlwinds, which often carry up considerable quantities of water and with it fishes of small size. In volcanic countries, also, the fishes of subterranean streams may sometimes be thrown up by volcanic explosions, as Humboldt relates happened in South America. Another mode by which fishes may be distributed is by their eggs being occasionally carried away by aquatic birds; and it is stated by Gmelin that geese and ducks during their migrations feed on the eggs of fish, and that some of these pass through their bodies with their vitality unimpaired. Even water-beetles flying from one pond to another might occasionally carry with them some of the smaller eggs of fishes. But it is probable that fresh-water fish are also enabled to migrate by changes of level causing streams to alter their course and carry their waters into adjacent basins. On plateaux the sources of distinct river systems often approach each other, and the same thing occurs with lateral tributaries on the lowlands near their mouths. Such changes, although small in extent, and occurring only at long intervals, would ​act very powerfully in modifying the distribution of fresh-water fish.
Sea fish would seem at first sight to have almost unlimited means of dispersal, but this is far from being the case. Temperature forms a complete barrier to a large number of species, cold water being essential to many, while others can only dwell in the warmth of the tropics. Deep water is another barrier to large numbers of species which are adapted to shores and shallows; and thus the Atlantic is quite as impassable a gulf to most fishes as it is to birds. Many sea fishes migrate to a limited extent for the purpose of depositing their spawn in favourable situations. The herring, an inhabitant of the deep sea, comes in shoals to our coast in the breeding season; while the salmon quits the northern seas and enters our rivers, mounting upwards to the clear cold water near their sources to deposit its eggs. Keeping in mind the essential fact that changes of temperature and of depth are the main barriers to the dispersal of fish, we shall find little difficulty in tracing the causes that have determined their distribution.
Means of Dispersal of Mollusca.—The marine, fresh-water, and land mollusca are three groups whose powers of dispersal and consequent distribution are very different, and must be separately considered. The Pteropoda, the Ianthina, and other groups of floating molluscs, drift about in mid-ocean, and their dispersal is probably limited chiefly by temperature, but perhaps also by the presence of enemies or the scarcity of proper food. The univalve and bivalve mollusca, of which the whelk and the cockle may be taken as types, move so slowly in their adult state, that we should expect them to have an exceedingly limited distribution; but the young of all these are free swimming embryos, and they thus have a powerful means of dispersal, and are carried by tides and currents so as ultimately to spread over every shore and shoal that offers conditions favourable for their development. The fresh water molluscs, which one might at first suppose could not range beyond their own river-basin, are yet very widely distributed in common with almost all other fresh water productions; and Mr. Darwin has shown that this is ​due to the fact, that ponds and marshes are constantly frequented by wading and swimming birds which are pre-eminently wanderers, and which frequently carry away with them the seeds of plants, and the eggs of molluscs and aquatic insects. Fresh water molluscs just hatched were found to attach themselves to a duck's foot suspended in an aquarium; and they would thus be easily carried from one lake or river to another, and by the help of different species of aquatic birds, might soon spread all over the globe. Even a water-beetle has been caught with a small living shell (Ancylus) attached to it; and these fly long distances and are liable to be blown out to sea, one having been caught on board the Beagle when forty-five miles from land. Although fresh water molluscs and their eggs must frequently be carried out to sea, yet this cannot lead to their dispersal, since salt water is almost immediately fatal to them; and we are therefore forced to conclude that the apparently insignificant and uncertain means of dispersal above alluded to are really what have led to their wide distribution. The true land-shells offer a still more difficult case, for they are exceedingly sensitive to the influence of salt water; they are not likely to be carried by aquatic birds, and yet they are more or less abundant all over the globe, inhabiting the most remote oceanic islands. It has been found, however, that land-shells have the power of lying dormant a long time. Some have lived two years and a half shut up in pill boxes; and one Egyptian desert snail came to life after having been glued down to a tablet in the British Museum for four years!
We are indebted to Mr. Darwin for experiments on the power of land shells to resist sea water, and he found that when they had formed a membranous diaphragm over the mouth of the shell they survived many days' immersion (in one case fourteen days); and another experimenter, quoted by Mr. Darwin, found that out of one hundred land shells immersed for a fortnight in the sea, twenty-seven recovered. It is therefore quite possible for them to be carried in the chinks of drift wood for many hundred miles across the sea, and this is probably one of the most effectual modes of their dispersal. Very young shells would also ​sometimes attach themselves to the feet of birds walking or resting on the ground, and as many of the waders often go far inland, this may have been one of the methods of distributing species of land shells; for it must always be remembered that nature can afford to wait, and that if but once in a thousand years a single bird should convey two or three minute snails to a distant island, this is all that is required for us to find that island well stocked with a great and varied population of land shells.
Means of Dispersal of Insects and the Barriers which Limit their Range.—Winged insects, as a whole, have perhaps more varied means of dispersal over the globe than any other highly organised animals. Many of them can fly immense distances, and the more delicate ones are liable to be carried by storms and hurricanes over a wide expanse of ocean. They are often met with far out at sea. Hawk-moths frequently fly on board ships as they approach the shores of tropical countries, and they have sometimes been captured more than 250 miles from the nearest land. Dragon-flies came on board the Adventure frigate when fifty miles off the coast of South America. A southerly wind brought flies in myriads to Admiral Smyth's ship in the Mediterranean when he was 100 miles distant from the coast of Africa. A large Indian beetle (Chrysochroa ocellata) was quite recently caught alive in the Bay of Bengal by Captain Payne of the barque William Mansoon, 273 miles from the nearest land. Darwin caught a locust 370 miles from land; and in 1844 swarms of locusts several miles in extent, and as thick as the flakes in a heavy snowstorm, visited Madeira. These must have come with perfect safety more than 300 miles; and as they continued flying over the island for a long time, they could evidently have travelled to a much greater distance. Numbers of living beetles belonging to seven genera, some aquatic and some terrestrial, were caught by Mr. Darwin in the open sea, seventeen miles from the coast of South America, and they did not seem injured by the salt water. Almost all the accidental causes that lead to the dispersal of the higher animals would be still more favourable for insects. Floating trees could carry hundreds of insects for one bird or mammal; and so many of the larvæ, eggs, ​and pupæ of insects have their abode in solid timber, that they might survive being floated immense distances. Great numbers of tropical insects have been captured in the London docks, where they have been brought in foreign timber; and some have emerged from furniture after remaining torpid for many years. Most insects have the power of existing weeks or months without food, and some are very tenacious of life. Many beetles will survive immersion for hours in strong spirit; and water a few degrees below the boiling point will not always kill them. We can therefore easily understand how, in the course of ages insects may become dispersed by means which would be quite inadequate in the case of the higher animals. The drift-wood and tropical fruits that reach Ireland and the Orkneys; the double cocoa-nuts that cross the Indian ocean from the Seychelle Islands to the coast of Sumatra; the winds that carry volcanic dust and ashes for thousands of miles; the hurricanes that travel in their revolving course over wide oceans; all indicate means by which a few insects may, at rare intervals be carried to remote regions, and become the progenitors of a group of allied forms.
But the dispersal of insects requires to be looked at from another point of view. They are, of all animals, perhaps the most wonderfully adapted for special conditions; and are so often fitted to fill one place in nature and one only, that the barriers against their permanent displacement are almost as numerous and as effective as their means of dispersal. Hundreds of species of lepidoptera, for example, can subsist in the larva state only on one species of plant; so that even if the perfect insects were carried to a new country, the continuance of the race would depend upon the same or a closely allied plant being abundant there. Other insects require succulent vegetable food all the year round, and are therefore confined to tropical regions; some can live only in deserts, others in forests; some are dependent on water-plants, some on mountain-vegetation. Many are so intimately connected with other insects during some part of their existence that they could not live without them; such are the parasitical hymenoptera and diptera, and those mimicking species whose welfare depends upon their being ​mistaken for something else. Then again, insects have enemies in every stage of their existence—the egg, the larva, the pupa, and the perfect form; and the abundance of any one of these enemies may render their survival impossible in a country otherwise well suited to them. Ever bearing in mind these two opposing classes of facts, we shall not be surprised at the enormous range of some groups of insects, and at the extreme localization of others; and shall be able to give a rational account of many phenomena of distribution that would otherwise seem quite unintelligible.
* ↑ Marcel de Serres states this as a general fact for wading and swimming birds. He says that the old birds arrive in the extreme north almost alone, the young remaining on the shores of the Baltic, or on the lakes of Austria, Hungary, and Russia. See his prize essay, Des Causes des Migrations, &c. 2nd. ed., Paris, 1845, p. 121.
* ↑ Quoted in Lyell's Principles of Geology (11th ed. vol. ii. p. 374), from Amœn. Acad. Essay 75.