Important geographical changes, such as the joining of lands that before were separate, or the dividing of continuous lands by transgressions and incursions of the sea, must necessarily have a profound effect upon the distribution of land mammals. Separated land-areas, however similar may have been their faunas at the time of separation, will, through the operation of the divergent evolutionary process, grow more unlike in proportion to the length of time that the separation continues. Regions which have been severed within a short time (in the geological sense of a short time) are zoölogically very similar or even identical, while those that have long been isolated are correspondingly peculiar. Attention has already been called, in another connection, to the contrasted cases of such great continental islands as Great Britain, Java, Sumatra, etc., on the one hand, and Australia, on the other. The continental islands, which have but lately been detached from the neighbouring main lands, are hardly more peculiar zoölogically than equal areas of the adjoining continents, while the long-continued isolation of Australia has made it the most peculiar region of the earth. Climatic changes, which, as we saw in [Chapter I], have indubitably taken place many times, have also had a great effect in shifting the distribution of mammals, which in its present form is the outcome of a very long series of geographical and climatic changes, on the one hand, and of adaptive changes in the animals themselves, on the other.
Of almost equal importance as a barrier is climate and especially temperature. Not that similar climates can produce similar forms in separate areas. Regions of almost exactly similar climate in Australia, Africa and South America have totally different faunas, but, within continuous land-areas, the most effective of barriers is temperature. This acts differently in the case of limiting the northward spread of southern forms and the southward spread of northern species. Dr. Merriam’s long study of this problem has led him to the conclusion that southern species are bounded on the north by the temperature of the breeding season, in which the total quantity of heat must reach a certain minimum, while “animals and plants are restricted in southward distribution by the mean temperature of a brief period covering the hottest part of the year.” On the Pacific coast there is a remarkable mingling in the same areas of species which, east of the high mountains, are distributed in sharply separated zones. This is explained by the mild and equable climate of the coastal belt, where the hottest season of the year does not reach the limiting maximum for the northern species, while the total quantity of heat in the breeding season is sufficient to enable southern species to thrive and maintain themselves.
Dr. Merriam thus sums up the effects of climatic factors upon distribution: “Humidity and other secondary causes determine the presence or absence of particular species in particular localities within their appropriate zones, but temperature pre-determines the possibilities of distribution; it fixes the limits beyond which species cannot pass.” “Concurrently with these changes in vegetation from the south northward occur equally marked differences in the mammals, birds, reptiles, and insects. Among mammals the tapirs, monkeys, armadillos, nasuas, peccaries, and opossums of Central America and Mexico are replaced to the northward by wood-rats, marmots, chipmunks, foxes, rabbits, short-tailed field-mice of several genera, shrews, wild-cats, lynxes, short-tailed porcupines, elk, moose, reindeer, sables, fishers, wolverines, lemmings, musk-oxen, and polar bears.”
Dr. J. A. Allen has reached closely similar conclusions. “Of strictly climatic influences, temperature is by far the most important, although moisture plays an influential part. Where a low temperature prevails life, both animal and vegetable, is represented by comparatively few forms; under a high temperature it is characterized by great diversity and luxuriance. Within the Arctic Circle the species of both animals and plants are not only few, but they are widely distributed, being for the most part everywhere the same. Under the tropics they are a hundred fold more numerous and of comparatively restricted distribution.” “The influence of temperature is perhaps most strikingly displayed in the distribution of life upon the slopes of a high mountain, especially if situated near the tropics. While its base may be clothed with palms and luxuriant tropical vegetation, its summit may be snow-capped and barren.... The animal life becomes likewise correspondingly changed, tropical forms of mammals, birds, and insects of the lower slopes gradually giving place to such as are characteristic of arctic latitudes.” “The effect of humidity upon plant life is thus obvious, but it is equally potent, though less evident, upon animal life. Many animals ... are so fitted for a forest life, as regards both food and shelter, that their very existence depends upon such surroundings.... Thus moisture alone may determine the character of life over extensive regions.”
While climate is thus the most important of the barriers which determine distribution in continuous land-areas, the absence of any particular species from a given region is no proof that the climate is unsuitable to that species. This is sufficiently shown by the manner in which animals introduced into a new country often run wild and multiply to an incredible extent, as the rabbits have done in Australia, the Mongoose in Jamaica, horses on our western plains, horses and cattle on the Pampas of Argentina, etc.
Topographical features, such as great mountain-ranges and plateaus, also limit many species, not only by the difficulty of crossing them, but also by the effect which they have upon temperature and moisture. For this reason long ranges of mountains and table-lands may carry a northern fauna very far to the south of its ordinary range, as do the mountain-systems of North America in a very conspicuous manner. The great Mexican plateau is zoölogically a part of North America, while the low coastal lands as far as southeastern Texas have Central American affinities.
A different kind of obstacle to the spread of a species into a new area may be the pre-occupation of that area by another species. The pre-occupier may be one that plays so similar a part in the economy of nature as to leave no opportunity for the newcomer to establish itself. On the other hand, the obstructing form may be an active enemy and of a totally different character from the intruder, as in the case of the Tse-tse Fly in parts of Africa. The bite of the fly is fatal to horses and oxen, so that these mammals are unable to enter the fly-infested regions. Many times in the course of the Tertiary period various mammals reached North America from the south or from the Old World, which were unable to gain a permanent foothold and speedily died out. At this distance of time it is seldom, if ever, possible to explain why a species which succeeded in reaching this continent could not maintain itself, though the most probable assumption is that the forms already in possession of the land were an insuperable obstacle to the intruders.
The rate of dispersal of a species into new areas may be fast or slow, according as the conditions are more or less favourable. Newly introduced insect-pests, like the Gypsy and the Brown-tailed Moths in New England, often spread with portentous rapidity; and introduced mammals have frequently taken possession of vast areas in a surprisingly short time. One of the most remarkable of these cases is cited by Darwin. “In the time of Sarmiento (1580) these Indians had bows and arrows, now long since disused; they then also possessed some horses. This is a very curious fact, showing the extraordinarily rapid multiplication of horses in South America. The horse was first landed at Buenos Ayres in 1537 and the colony being then for a time deserted, the horse ran wild; in 1580, only forty-three years afterwards, we hear of them at the Strait of Magellan!” (“Voyage of a Naturalist,” pp. 232-233.) In this example, something must be allowed for human agency, but even so, it is very surprising.
In the case of lands newly raised above the sea and connecting formerly separated areas, it is necessary that they should first be taken possession of by vegetation, before they can become passable by animals, for the migration of mammals from continent to continent is an entirely distinct phenomenon from the annual migration of birds. The latter, though a fact familiar to every one, is an unexplained mystery, and it is somewhat unfortunate that the same term should be used for the completely different process of the spread of mammals into newly opened land. This spread is purely unconscious and is due to the pressure of increasing numbers upon the means of subsistence, each new generation ranging farther and farther from the original home of the species and continuing so to extend until some insuperable obstacle is encountered. When a sea-barrier is removed by upheaval and the newly formed land rendered habitable for mammals through the invasion of plants, the interrupted process is resumed and an interchange of species between the areas thus connected is brought about. The interchange is, however, always an incomplete one, certain forms not being able so to extend their range, because of climatic differences, pre-occupation or some such barrier.