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12 - Geographical Distribution -- continued 12-10 - Distribution of fresh-water productions 10 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 impassable barrier, that they never would have extended to distant countries.

But the case is exactly the reverse.

Not only have many fresh-water species, belonging to quite different classes, an enormous range, but allied species prevail in a remarkable manner throughout the world.

I well remember, when first collecting in the fresh waters of Brazil, 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.

Brazil
Brazil


But this power in fresh-water productions of ranging widely, though so unexpected, 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; and liability to wide dispersal would follow from this capacity as an almost necessary consequence.

We can here consider only a few cases.

In regard to fish, I believe that the same species never occur in the fresh waters of distant continents.

But on the same continent the species often range widely and almost capriciously; for two river-systems will have some fish in common and some different.

A few facts seem to favour the possibility of their occasional transport by accidental means; like that of the live fish not rarely dropped by whirlwinds in India, and the vitality of their ova when removed from the water.

India
India


But I am inclined to attribute the dispersal of fresh-water fish mainly to slight changes within the recent period in the level of the land, having caused rivers to flow into each other.

Instances, also, could be given of this having occurred during floods, without any change of level.

We have evidence in the loess of the Rhine of considerable changes of level in the land within a very recent geological period, and when the surface was peopled by existing land and fresh-water shells.

Rhine
Rhine


The wide difference of the fish on opposite sides of continuous mountain-ranges, which from an early period must have parted river-systems and completely prevented their inosculation, seems to lead to this same conclusion.

With respect to allied fresh-water fish occurring at very distant points of the world, no doubt there are many cases which cannot at present be explained: but 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.

In the second place, 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 fishes confined exclusively to fresh water, so that we may imagine that a marine member of a fresh-water group might travel far along the shores of the sea, and subsequently become modified and adapted to the fresh waters of a distant land.

Some species of fresh-water shells have a very wide range, and allied species, which, on my theory, are descended from a common parent and must have proceeded from a single source, prevail throughout the world.

Freshwater Shells
Freshwater Shells


Their distribution at first perplexed me much, as their ova are not likely to be transported by birds, and they are immediately killed by sea water, as are the adults.

I could not even understand how some naturalised species have rapidly spread throughout the same country.

But two facts, which I have observed and no doubt many others remain to be observed throw some light on this subject.

When a duck suddenly emerges from a pond covered with duck-weed, I have twice seen these little plants adhering to its back; and it has happened to me, in removing a little duck-weed from one aquarium to another, that I have quite unintentionally stocked the one with fresh-water shells from the other.

duck
duck

duckweed (Lemnaceae)
duckweed (Lemnaceae)

aquarium
aquarium


But another agency is perhaps more effectual: I suspended a duck's feet, which might represent those of a bird sleeping in a natural pond, 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 would be sure to alight on a pool or rivulet, if blown across sea to an oceanic island or to any other distant point.

heron
heron


Sir Charles Lyell also informs me that a Dyticus 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 flown with a favouring gale no one can tell.

Sir Charles Lyell
Sir Charles Lyell

dyticus
dyticus

ancylus
ancylus

Colymbetes
Colymbetes

HMS Beagle
HMS Beagle

beagle
beagle


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 shown, as remarked by Alph. de Candolle, in large groups of terrestrial plants, which have only a very few aquatic members; for these latter seem immediately to acquire, as if in consequence, a very wide range.

Alphonse de Candolle
Alphonse de Candolle


I think favourable means of dispersal explain this fact.

I have before mentioned that earth occasionally, though rarely, adheres in some quantity to the feet and beaks of birds.

beak
beak


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 I can show are the greatest wanderers, and are occasionally found on the most remote and barren islands in the open ocean; they would not be likely to alight on the surface of the sea, so that the dirt would not be washed off their feet; when making 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 table-spoonfuls of mud from three different points, beneath water, on the edge of a little pond; this mud when dry weighed only 6 3/4 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 vast distances, and if consequently the range of these plants was not very great.

seeds
seeds

Breakfast Cup
Breakfast Cup


The same agency may have come into play with the eggs of some of the smaller fresh-water animals.

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.

Yellow Water Lily
Yellow Water Lily

potamogeton
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 in pellets or in excrement, many hours afterwards.

heron
heron


When I saw the great size of the seeds of that fine water-lily, the Nelumbium, and remembered Alph. de Candolle's remarks on this plant, I thought that its distribution must remain quite 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; although I do not know the fact, yet analogy makes me believe that a heron flying to another pond and getting a hearty meal of fish, would probably reject from its stomach a pellet containing the seeds of the Nelumbium undigested; or the seeds might be dropped by the bird whilst feeding its young, in the same way as fish are known sometimes to be dropped.

nelumbium (Lotus)
nelumbium (Lotus)

Alphonse de Candolle
Alphonse de Candolle

John James Audubon
John James Audubon

Joseph Dalton Hooker
Joseph Dalton Hooker

Nelumbium Lutea
Nelumbium Lutea

heron
heron

stomach
stomach


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 individuals of the species, however few, already occupying any pond, yet as the number of kinds is small, compared with those on the land, the competition will probably be less severe between aquatic than between terrestrial species; consequently an intruder from the waters of a foreign country, would have a better chance of seizing on a place, than in the case of terrestrial colonists.

We should, also, remember that some, perhaps many, fresh-water productions are low in the scale of nature, and that we have reason to believe that such low beings change or become modified less quickly than the high; and this will give longer time than the average for the migration of the same aquatic species.

We should not forget the probability of many species having formerly ranged as continuously as fresh-water productions ever can range, over immense areas, and having subsequently become extinct in intermediate regions.

But the wide distribution of fresh-water plants and of the lower animals, whether retaining the same identical form or in some degree modified, I believe mainly depends on the wide dispersal of their seeds and eggs by animals, more especially by fresh-water birds, which have large powers of flight, and naturally travel from one to another and often distant piece of water.

Nature, like a careful gardener, thus takes her seeds from a bed of a particular nature, and drops them in another equally well fitted for them.

gardener
gardener
12 - Geographical Distribution -- continued 12-20 - On the inhabitants of oceanic islands 10 We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty, on the view that all the individuals both of the same and of allied species have descended from a single parent; and therefore have all proceeded from a common birthplace, notwithstanding that in the course of time they have come to inhabit distant points of the globe.

I have already stated that I cannot honestly admit Forbes's view on continental extensions, which, if legitimately followed out, would lead to the belief that within the recent period all existing islands have been nearly or quite joined to some continent.

Edward Forbes
Edward Forbes


This view would remove many difficulties, but it would not, I think, explain all the facts in regard to insular productions.

In the following remarks I shall not confine myself to the mere question of dispersal; but shall consider some other facts, which bear on the truth of the two theories of independent creation and of descent with modification.

The species of all kinds which inhabit oceanic islands are few in number compared with those on equal continental areas: Alph. de Candolle admits this for plants, and Wollaston for insects.

Alphonse de Candolle
Alphonse de Candolle


If we look to the large size and varied stations of New Zealand, extending over 780 miles of latitude, and compare its flowering plants, only 750 in number, with those on an equal area at the Cape of Good Hope or in Australia, we must, I think, admit that something quite independently of any difference in physical conditions has caused so great a difference in number.

New Zealand
New Zealand


Even the uniform county of Cambridge has 847 plants, and the little island of Anglesea 764, but a few ferns and a few introduced plants are included in these numbers, and the comparison in some other respects is not quite fair.

Cambridge County
Cambridge County

Anglesea
Anglesea


We have evidence that the barren island of Ascension aboriginally possessed under half-a-dozen flowering plants; yet many have become naturalised on it, as they have on New Zealand and on every other oceanic island which can be named.

Ascension Island
Ascension Island


In St. Helena there is reason to believe that the naturalised plants and animals have nearly or quite exterminated many native productions.

St. Helena
St. Helena


He who admits the doctrine of the creation of each separate species, will have to admit, that a sufficient number of the best adapted plants and animals have not been created on oceanic islands; for man has unintentionally stocked them from various sources far more fully and perfectly than has nature.

Although in oceanic islands the number of kinds of inhabitants is scanty, the proportion of endemic species (i.e. those found nowhere else in the world) is often extremely large.

If we compare, for instance, the number of the endemic land-shells in Madeira, or of the endemic birds in the Galapagos Archipelago, with the number found on any continent, and then compare the area of the islands with that of the continent, we shall see that this is true.

Land Shell
Land Shell

Madeira
Madeira

Galapagos Islands
Galapagos Islands


This fact might have been expected on my theory for, as already explained, species occasionally arriving after long intervals in a new and isolated district, and having to compete with new associates, will be eminently liable to modification, and will often produce groups of modified descendants.

But it by no means follows, that, because in an island nearly all the species of one class are peculiar, those of another class, or of another section of the same class, are peculiar; and this difference seems to depend on the species which do not become modified having immigrated with facility and in a body, so that their mutual relations have not been much disturbed.

Thus in the Galapagos Islands nearly every land-bird, but only two out of the eleven marine birds, are peculiar; and it is obvious that marine birds could arrive at these islands more easily than land-birds.

Bermuda, on the other hand, which lies at about the same distance from North America as the Galapagos Islands do from South America, and which has a very peculiar soil, does not possess one endemic land bird; and we know from Mr. J. M. Jones's admirable account of Bermuda, that very many North American birds, during their great annual migrations, visit either periodically or occasionally this island.

Bermuda
Bermuda


Madeira does not possess one peculiar bird, and many European and African birds are almost every year blown there, as I am informed by Mr. E. V. Harcourt.

Africa
Africa


So that these two islands of Bermuda and Madeira have been stocked by birds, which for long ages have struggled together in their former homes, and have become mutually adapted to each other; and when settled in their new homes, each kind will have been kept by the others to their proper places and habits, and will consequently have been little liable to modification.

Madeira, again, is inhabited by a wonderful number of peculiar land-shells, whereas not one species of sea-shell is confined to its shores: now, though we do not know how seashells are dispersed, yet we can see that their eggs or larvae, perhaps attached to seaweed or floating timber, or to the feet of wading-birds, might be transported far more easily than land-shells, across three or four hundred miles of open sea.

Sea Shell
Sea Shell


The different orders of insects in Madeira apparently present analogous facts.

Oceanic islands are sometimes deficient in certain classes, and their places are apparently occupied by the other inhabitants; in the Galapagos Islands reptiles, and in New Zealand gigantic wingless birds, take the place of mammals.

Extinct Birds of New Zealand
Extinct Birds of New Zealand


In the plants of the Galapagos Islands, Dr. Hooker has shown that the proportional numbers of the different orders are very different from what they are elsewhere.

Joseph Dalton Hooker
Joseph Dalton Hooker


Such cases are generally accounted for by the physical conditions of the islands; but this explanation seems to me not a little doubtful.

Facility of immigration, I believe, has been at least as important as the nature of the conditions.

Many remarkable little facts could be given with respect to the inhabitants of remote islands.

For instance, in certain islands not tenanted by mammals, some of the endemic plants have beautifully hooked seeds; yet few relations are more striking than the adaptation of hooked seeds for transportal by the wool and fur of quadrupeds.

This case presents no difficulty on my view, for a hooked seed might be transported to an island by some other means; and the plant then becoming slightly modified, but still retaining its hooked seeds, would form an endemic species, having as useless an appendage as any rudimentary organ, for instance, as the shrivelled wings under the soldered elytra of many insular beetles.

Hooked Seed
Hooked Seed

beetle
beetle


Again, islands often possess trees or bushes belonging to orders which elsewhere include only herbaceous species; now trees, as Alph. de Candolle has shown, generally have, whatever the cause may be, confined ranges.

Hence trees would be little likely to reach distant oceanic islands; and an herbaceous plant, though it would have no chance of successfully competing in stature with a fully developed tree, when established on an island and having to compete with herbaceous plants alone, might readily gain an advantage by growing taller and taller and overtopping the other plants.

If so, natural selection would often tend to add to the stature of herbaceous plants when growing on an island, to whatever order they belonged, and thus convert them first into bushes and ultimately into trees.

island
island

tree
tree
12 - Geographical Distribution -- continued 12-30 - Absence of Batrachians and of terrestrial Mammals 10 With respect to the absence of whole orders on oceanic islands, Bory St. Vincent long ago remarked that Batrachians (frogs, toads, newts) have never been found on any of the many islands with which the great oceans are studded.

Bory Saint Vincent
Bory Saint Vincent

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frogs

toad
toad

newt
newt
12 - Geographical Distribution -- continued 12-30 - Absence of Batrachians and of terrestrial Mammals 20 I have taken pains to verify this assertion, and I have found it strictly true.

I have, however, been assured that a frog exists on the mountains of the great island of New Zealand; but I suspect that this exception (if the information be correct) may be explained through glacial agency.

New Zealand
New Zealand


This general absence of frogs, toads, and newts on so many oceanic islands cannot be accounted for by their physical conditions; indeed it seems that islands are peculiarly well fitted for these animals; for frogs have been introduced into Madeira, the Azores, and Mauritius, and have multiplied so as to become a nuisance.

island
island

Madeira
Madeira

Azores
Azores

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Mauritius