Sir Charles Lyell observes that if, by the attacks of seals or other marine foes, salmon are reduced in numbers, the consequence will be that otters, living far inland, will be deprived of food and will then destroy many young birds or quadrupeds, so that the increase of a marine animal may cause the destruction of many land animals hundreds of miles away. Mr. Darwin carefully observed the effects produced by planting a few hundred acres of Scotch fir, in Staffordshire, on part of a very extensive heath which had never been cultivated. After the planted portion was about twenty-five years old he observed that the change in the native vegetation was greater than is often seen in passing from one quite different soil to another. Besides a great change in the proportional numbers of the native heath-plants, twelve species which could not be found on the heath flourished in the plantations. The effect on the insect life must have been still greater, for six insectivorous birds which were very common in the plantations were not to be seen on the heath, which was, however, frequented by two or three different species of insectivorous birds. It would have required continued study for several years to determine all the differences in the organic life of the two areas, but the facts stated by Mr. Darwin are sufficient to show how great a change may be effected by the introduction of a single kind of tree and the keeping out of cattle.
The next case I will give in Mr. Darwin's own words: "In several parts of the world insects determine the existence of cattle. Perhaps Paraguay offers the most curious instance of this; for here neither cattle nor horses nor dogs have ever run wild, though they swarm southward and northward in a feral state; and Azara and Rengger have shown that this is caused by the greater numbers, in Paraguay, of a certain fly which lays its eggs in the navels of these animals when first born. The increase of these flies, numerous as they are, must be habitually checked by some means, probably by other parasitic insects. Hence, if certain insectivorous birds were to decrease in Paraguay, the parasitic insects would probably increase; and this would lessen the number of the navel-frequenting flies—then cattle and horses would become feral, and this would greatly alter (as indeed I have observed in parts of South America) the vegetation: this again would largely affect the insects, and this, as we have just seen in Staffordshire, the insectivorous birds, and so onward in ever-increasing circles of complexity. Not that under nature the relations will ever be as simple as this. Battle within battle must be continually recurring with varying success; and yet in the long run the forces are so nicely balanced, that the face of nature remains for a long time uniform, though assuredly the merest trifle would give the victory to one organic being over another."[7]
Such cases as the above may perhaps be thought exceptional, but there is good reason to believe that they are by no means rare, but are illustrations of what is going on in every part of the world, only it is very difficult for us to trace out the complex reactions that are everywhere occurring. The general impression of the ordinary observer seems to be that wild animals and plants live peaceful lives and have few troubles, each being exactly suited to its place and surroundings, and therefore having no difficulty in maintaining itself. Before showing that this view is, everywhere and always, demonstrably untrue, we will consider one other case of the complex relations of distinct organisms adduced by Mr. Darwin, and often quoted for its striking and almost eccentric character. It is now well known that many flowers require to be fertilised by insects in order to produce seed, and this fertilisation can, in some cases, only be effected by one particular species of insect to which the flower has become specially adapted. Two of our common plants, the wild heart's-ease (Viola tricolor) and the red clover (Trifolium pratense), are thus fertilised by humble-bees almost exclusively, and if these insects are prevented from visiting the flowers, they produce either no seed at all or exceedingly few. Now it is known that field-mice destroy the combs and nests of humble-bees, and Colonel Newman, who has paid great attention to these insects, believes that more than two-thirds of all the humble-bees' nests in England are thus destroyed. But the number of mice depends a good deal on the number of cats; and the same observer says that near villages and towns he has found the nests of humble-bees more numerous than elsewhere, which he attributes to the number of cats that destroy the mice. Hence it follows, that the abundance of red clover and wild heart's-ease in a district will depend on a good supply of cats to kill the mice, which would otherwise destroy and keep down the humble-bees and prevent them from fertilising the flowers. A chain of connection has thus been found between such totally distinct organisms as flesh-eating mammalia and sweet-smelling flowers, the abundance or scarcity of the one closely corresponding to that of the other!
The following account of the struggle between trees in the forests of Denmark, from the researches of M. Hansten-Blangsted, strikingly illustrates our subject.[8] The chief combatants are the beech and the birch, the former being everywhere successful in its invasions. Forests composed wholly of birch are now only found in sterile, sandy tracts; everywhere else the trees are mixed, and wherever the soil is favourable the beech rapidly drives out the birch. The latter loses its branches at the touch of the beech, and devotes all its strength to the upper part where it towers above the beech. It may live long in this way, but it succumbs ultimately in the fight—of old age if of nothing else, for the life of the birch in Denmark is shorter than that of the beech. The writer believes that light (or rather shade) is the cause of the superiority of the latter, for it has a greater development of its branches than the birch, which is more open and thus allows the rays of the sun to pass through to the soil below, while the tufted, bushy top of the beech preserves a deep shade at its base. Hardly any young plants can grow under the beech except its own shoots; and while the beech can nourish under the shade of the birch, the latter dies immediately under the beech. The birch has only been saved from total extermination by the facts that it had possession of the Danish forests long before the beech ever reached the country, and that certain districts are unfavourable to the growth of the latter. But wherever the soil has been enriched by the decomposition of the leaves of the birch the battle begins. The birch still flourishes on the borders of lakes and other marshy places, where its enemy cannot exist. In the same way, in the forests of Zeeland, the fir forests are disappearing before the beech. Left to themselves, the firs are soon displaced by the beech. The struggle between the latter and the oak is longer and more stubborn, for the branches and foliage of the oak are thicker, and offer much resistance to the passage of light. The oak, also, has greater longevity; but, sooner or later, it too succumbs, because it cannot develop in the shadow of the beech. The earliest forests of Denmark were mainly composed of aspens, with which the birch was apparently associated; gradually the soil was raised, and the climate grew milder; then the fir came and formed large forests. This tree ruled for centuries, and then ceded the first place to the holm-oak, which is now giving way to the beech. Aspen, birch, fir, oak, and beech appear to be the steps in the struggle for the survival of the fittest among the forest-trees of Denmark.
It may be added that in the time of the Romans the beech was the principal forest-tree of Denmark as it is now, while in the much earlier bronze age, represented by the later remains found in the peat bogs, there were no beech-trees, or very few, the oak being the prevailing tree, while in the still earlier stone period the fir was the most abundant. The beech is a tree essentially of the temperate zone, having its northern limit considerably southward of the oak, fir, birch, or aspen, and its entrance into Denmark was no doubt due to the amelioration of the climate after the glacial epoch had entirely passed away. We thus see how changes of climate, which are continually occurring owing either to cosmical or geographical causes, may initiate a struggle among plants which may continue for thousands of years, and which must profoundly modify the relations of the animal world, since the very existence of innumerable insects, and even of many birds and mammals, is dependent more or less completely on certain species of plants.
The Struggle for Existence on the Pampas.
Another illustration of the struggle for existence,in which both plants and animals are implicated, is afforded by the pampas of the southern part of South America. The absence of trees from these vast plains has been imputed by Mr. Darwin to the supposed inability of the tropical and sub-tropical forms of South America to thrive on them, and there being no other source from which they could obtain a supply; and that explanation was adopted by such eminent botanists as Mr. Ball and Professor Asa Gray. This explanation has always seemed to me unsatisfactory, because there are ample forests both in the temperate regions of the Andes and on the whole west coast down to Terra del Fuego; and it is inconsistent with what we know of the rapid variation and adaptation of species to new conditions. What seems a more satisfactory explanation has been given by Mr. Edwin Clark, a civil engineer, who resided nearly two years in the country and paid much attention to its natural history. He says: "The peculiar characteristics of these vast level plains which descend from the Andes to the great river basin in unbroken monotony, are the absence of rivers or water-storage, and the periodical occurrence of droughts, or 'siccos,' in the summer months. These conditions determine the singular character both of its flora and fauna.
"The soil is naturally fertile and favourable for the growth of trees, and they grow luxuriantly wherever they are protected. The eucalyptus is covering large tracts wherever it is enclosed, and willows, poplars, and the fig surround every estancia when fenced in.
"The open plains are covered with droves of horses and cattle, and overrun by numberless wild rodents, the original tenants of the pampas. During the long periods of drought, which are so great a scourge to the country, these animals are starved by thousands, destroying, in their efforts to live, every vestige of vegetation. In one of these 'siccos,' at the time of my visit, no less than 50,000 head of oxen and sheep and horses perished from starvation and thirst, after tearing deep out of the soil every trace of vegetation, including the wiry roots of the pampas-grass. Under such circumstances the existence of an unprotected tree is impossible. The only plants that hold their own, in addition to the indestructible thistles, grasses, and clover, are a little herbaceous oxalis, producing viviparous buds of extraordinary vitality, a few poisonous species, such as the hemlock, and a few tough, thorny dwarf-acacias and wiry rushes, which even a starving rat refuses.
"Although the cattle are a modern introduction, the numberless indigenous rodents must always have effectually prevented the introduction of any other species of plants; large tracts are still honeycombed by the ubiquitous biscacho, a gigantic rabbit; and numerous other rodents still exist, including rats and mice, pampas-hares, and the great nutria and carpincho (capybara) on the river banks."[9]