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Charles Robert Darwin (1809–1882).  Origin of Species.
The Harvard Classics.  1909–14.
 
XIII. Geographical Distribution—Continued
 
Fresh-water Productions
 
 
AS lakes and river-systems are separated from each other by barriers of land, it might have been thought that fresh-water productions would not have ranged widely within the same country, and as the sea is apparently a still more formidable barrier, that they would never have extended to distant countries. But the case is exactly the reverse. Not only have many fresh-water species, belonging to different classes, an enormous range, but allied species prevail in a remarkable manner throughout the world. When first collecting in the fresh waters of Brazil, I well remember feeling much surprise at the similarity of the fresh-water insects, shells &c., and at the dissimilarity of the surrounding terrestrial beings, compared with those of Britain.  1
  But the wide ranging power of fresh-water productions can, I think, in most cases be explained by their having become fitted, in a manner highly useful to them, for short and frequent migrations from pond to pond, or from stream to stream, within their own countries; and liability to wide dispersal would follow from this capacity as an almost necessary consequence. We can here consider only a few cases; of these, some of the most difficult to explain are presented by fish. It was formerly believed that the same fresh-water species never existed on two continents distant from each other. But Dr. Günther has lately shown that the Galaxias attenuatus inhabits Tasmania, New Zealand, the Falkland Islands, and the mainland of South America. This is a wonderful case, and probably indicates dispersal from an Antarctic centre during a former warm period. This case, however, is rendered in some degree less surprising by the species of this genus having the power of crossing by some unknown means considerable spaces of open ocean: thus there is one species common to New Zealand and to the Auckland Islands, though separated by a distance of about 230 miles. On the same continent fresh-water fish often range widely, and as if capriciously; for in two adjoining river-systems some of the species may be the same, and some wholly different.  2
  It is probable that they are occasionally transported by what may be called accidental means. Thus fishes still alive are not very rarely dropped at distant points by whirlwinds; and it is known that the ova retain their vitality for a considerable time after removal from the water. Their dispersal may, however, be mainly attributed to changes in the level of the land within the recent period, causing rivers to flow into each other. Instances, also, could be given of this having occurred during floods, without any change of level. The wide difference of the fish on the opposite sides of most mountain-ranges, which are continuous, and which consequently must from an early period have completely prevented the inosculation of the river-systems on the two sides, leads to the same conclusion. Some fresh-water fish belong to very ancient forms, and in such cases there will have been ample time for great geographical changes, and consequently time and means for much migration. Moreover, Dr. Günther has recently been led by several considerations to infer that with fishes the same forms have a long endurance. Salt-water fish can with care be slowly accustomed to live in fresh water; and, according to Valenciennes’ there is hardly a single group of which an the members are confined to fresh water, so that a marine species belonging to a fresh-water group might travel far along the shores of the sea, and could, it is probable, become adapted without much difficulty to the fresh waters of a distant land.  3
  Some species of fresh-water shells have very wide ranges, and allied species which, on our theory, are descended from a common parent, and must have proceeded from a single source, prevail throughout the world. Their distribution at first perplexed me much, as their ova are not likely to be transported by birds; and the ova, as well as the adults, are immediately killed by sea-water. I could not even understand how some naturalised species have spread rapidly throughout the same country. But two facts, which I have observed—and many others no doubt will be discovered—throw some light on this subject. When ducks suddenly emerge from a pond covered with duck-weed, I have twice seen these little plants adhering to their backs; and it has happened to me, in removing a little duck-weed from one aquarium to another, that I have unintentionally stocked the one with fresh-water shells from the other. But another agency is perhaps more effectual: I suspended the feet of a duck in an aquarium, where many ova of fresh-water shells were hatching; and I found that numbers of the extremely minute and just-hatched shells crawled on the feet, and clung to them so firmly that when taken out of the water they could not be jarred off, though at a somewhat more advanced age they would voluntarily drop off. These just-hatched molluscs, though aquatic in their nature, survived on the duck’s feet, in damp air, from twelve to twenty-hours; and in this length of time a duck or heron might fly at least six or seven hundred miles, and if blown across the sea to an oceanic island, or to any other distant point, would be sure to alight on a pool or rivulet. Sir Charles Lyell informs me that a Dytiscus has been caught with an Ancylus (a fresh-water shell like a limpet) firmly adhering to it; and a water-beetle of the same family, a Colymbetes, once flew on board the ‘Beagle’, when forty-five miles distant from the nearest land: how much farther it might have been blown by a favouring gale no one can tell.  4
  With respect to plants, it has long been known what enormous ranges many fresh-water, and even marsh species, have, both over continents and to the most remote oceanic islands. This is strikingly illustrated, according to Alph. de Candolle, in those large groups of terrestrial plants, which have very few aquatic members; for the latter seem immediately to acquire, as if in consequence, a wide range. I think favourable means of dispersal explain this fact. I have before mentioned that earth occasionally adheres in some quantity to the feet and beaks of birds. Wading birds, which frequent the muddy edges of ponds, if suddenly flushed, would be the most likely to have muddy feet. Birds of this order wander more than those of any other; and they are occasionally found on the most remote and barren islands of the open ocean; they would not be likely to alight on the surface of the sea, so that any dirt on their feet would not be washed off; and when gaining the land, they would be sure to fly to their natural fresh-water haunts. I do not believe that botanists are aware how charged the mud of ponds is with seeds; I have tried several little experiments, but will here give only the most striking case: I took in February three tablespoonfuls of mud from three different points, beneath water, on the edge of a little pond: this mud when dried weighed only 63/4 ounces; I kept it covered up in my study for six months, pulling up and counting each plant as it grew; the plants were of many kinds, and were altogether 537 in number; and yet the viscid mud was all contained in a breakfast cup! Considering these facts, I think it would be an inexplicable circumstance if water-birds did not transport the seeds of fresh-water plants to unstocked ponds and streams, situated at very distant points. The  same agency may have come into play with the eggs of some of the smaller fresh-water animals.  5
  Other and unknown agencies probably have also played a part. I have stated that fresh-water fish eat some kinds of seeds, though they reject many other kinds after having swallowed them; even small fish swallow seeds of moderate size, as of the yellow water-lily and Potamogeton. Herons and other birds, century after century, have gone on daily devouring fish; they then take flight and go to other waters, or are blown across the sea; and we have seen that seeds retain their power of germination, when rejected many hours afterwards in pellets or in the excrement. When I saw the great size of the seeds of that fine water-lily, the Nelumbium, and remembered Alph. de Candolle’s remarks on the distribution of this plant, I thought that the means of its dispersal must remain inexplicable; but Audubon states that he found the seeds of the great southern water-lily (probably, according to Dr. Hooker, the Nelumbium luteum) in a heron’s stomach. Now this bird must often have flown with its stomach thus well stocked to distant ponds, and then getting a hearty meal of fish, analogy makes me believe that it would have rejected the seeds in a pellet in a fit state for germination.  6
  In considering these several means of distribution, it should be remembered that when a pond or stream is first formed, for instance, on a rising islet, it will be unoccupied; and a single seed or egg will have a good chance of succeeding. Although there will always be a struggle for life between the inhabitants of the same pond, however few in kind, yet as the number even in a well-stocked pond is small in comparison with the number of species inhabiting an equal area of land, the competition between them will probably be less severe than between terrestrial species; consequently an intruder from the waters of a foreign country would have a better chance of seizing on new place, than in the case of terrestrial colonists. We should also remember that many fresh-water productions are low in the scale of nature, and we have reason to believe that such beings become modified more slowly than the high; and this will give time for the migration of aquatic species. We should not forget the probability of many fresh-water forms laving formerly ranged continuously over immense areas, and then having become extinct at intermediate points. But the wide distribution of fresh-water plants and of the lower animals, whether retaining the same identical form or in some degree modified, apparently depends in main part on the wide dispersal of their seeds and eggs by animals, more especially by fresh-water birds, which have great powers of flight, and naturally travel from one piece of water to another.  7
 

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