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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
06 - Difficutiles in Theory 06-09 - Cases of Difficulty 10 Although we must be extremely cautious in concluding that any organ could not have been produced by successive, small, transitional gradations, yet undoubtedly serious cases of difficulty occur.

One of the most serious is that of neuter insects, which are often differently constructed from either the males or fertile females; but this case will be treated of in the next chapter.
Ant Castes
Ant Castes


The electric organs of fishes offer another case of special difficulty; for it is impossible to conceive by, what steps these wondrous organs have been produced.
Electric Eel
Electric Eel


But this is not surprising, for we do not even know of what use they are.

In the Gymnotus and torpedo they no doubt serve as powerful means of defence, and perhaps for securing prey; yet in the ray, as observed by Matteucci, an analogous organ in the tail manifests but little electricity, even when the animal is greatly irritated; so little, that it can hardly be of any use for the above purposes.
gymnotus
gymnotus

torpedo
torpedo

Electric  Ray
Electric Ray


Moreover, in the ray, besides the organ just referred to, there is, as Dr. R. McDonnell has shown, another organ near the head, not known to be electrical, but which appears to be the real homologue of the electric battery in the torpedo.

It is generally admitted that there exists between these organs and ordinary muscle a close analogy, in intimate structure, in the distribution of the nerves, and in the manner in which they are acted on by various reagents.

It should, also, be especially observed that muscular contraction is accompanied by an electrical discharge; and, as Dr. Radcliffe insists, "in the electrical apparatus of the torpedo during rest, there would seem be a charge in every respect like that which is met with in muscle and nerve during rest, and the discharge of the torpedo, instead of being peculiar, may be only another form of the discharge which depends upon the action of muscle and motor nerve."

Beyond this we cannot at present go in the way of explanation; but as we know so little about the uses of these organs, and as we know nothing about the habits and structure of the progenitors of the existing electric fishes, it would be extremely bold to maintain that no serviceable transitions are possible by which these organs might have been gradually developed.
11 - Geographical Distribution 11-03 - Affinity of the productions of the same continent 10 A third great fact, partly included in the foregoing statements, is the affinity of the productions of the same continent or sea, though the species themselves are distinct at different points and stations.

It is a law of the widest generality, and every continent offers innumerable instances.

Nevertheless the naturalist in travelling, for instance, from north to south never fails to be struck by the manner in which successive groups of beings, specifically distinct, yet clearly related, replace each other.

He hears from closely allied, yet distinct kinds of birds, notes nearly similar, and sees their nests similarly constructed, but not quite alike, with eggs coloured in nearly the same manner.

bird
bird

egg
egg


The plains near the Straits of Magellan are inhabited by one species of Rhea (American ostrich), and northward the plains of La Plata by another species of the same genus; and not by a true ostrich or emeu, like those found in Africa and Australia under the same latitude.

Straits of Magellan
Straits of Magellan

rhea
rhea

La Plata
La Plata

ostrich
ostrich

emeu
emeu

Africa
Africa

Australia
Australia


On these same plains of La Plata, we see the agouti and bizcacha, animals having nearly the same habits as our hares and rabbits and belonging to the same order of Rodents, but they plainly display an American type of structure.

La Plata
La Plata

agouti
agouti

bizcacha
bizcacha

hare
hare

rabbit
rabbit

America
America


We ascend the lofty peaks of the Cordillera and we find an alpine species of bizcacha; we look to the waters, and we do not find the beaver or musk-rat, but the coypu and capybara, rodents of the American type.

Cordillera
Cordillera

beaver
beaver

Musk Rat
Musk Rat

coypu
coypu

capybara
capybara


Innumerable other instances could be given.

If we look to the islands off the American shore, however much they may differ in geological structure, the inhabitants, though they may be all peculiar species, are essentially American.

We may look back to past ages, as shown in the last chapter, and we find American types then prevalent on the American continent and in the American seas.

We see in these facts some deep organic bond, prevailing throughout space and time, over the same areas of land and water, and independent of their physical conditions.

The naturalist must feel little curiosity, who is not led to inquire what this bond is.

This bond, on my theory, is simply inheritance, that cause which alone, as far as we positively know, produces organisms quite like, or, as we see in the case of varieties nearly like each other.

The dissimilarity of the inhabitants of different regions may be attributed to modification through natural selection, and in a quite subordinate degree to the direct influence of different physical conditions.

The degree of dissimilarity will depend on the migration of the more dominant forms of life from one region into another having been effected with more or less ease, at periods more or less remote; on the nature and number of the former immigrants; -- and on their action and reaction, in their mutual struggles for life; the relation of organism to organism being, as I have already often remarked, the most important of all relations.

Thus the high importance of barriers comes into play by checking migration; as does time for the slow process of modification through natural selection.

Widely-ranging species, abounding in individuals, which have already triumphed over many competitors in their own widely-extended homes will have the best chance of seizing on new places, when they spread into new countries.

In their new homes they will be exposed to new conditions, and will frequently undergo further modification and improvement; and thus they will become still further victorious, and will produce groups of modified descendants.

On this principle of inheritance with modification, we can understand how it is that sections of genera, whole genera, and even families are confined to the same areas, as is so commonly and notoriously the case.

I believe, as was remarked in the last chapter, in no law of necessary development.

As the variability of each species is an independent property, and will be taken advantage of by natural selection, only so far as it profits the individual in its complex struggle for life, so the degree of modification in different species will be no uniform quantity.

If, for instance, a number of species, which stand in direct competition with each other, migrate in a body into a new and afterwards isolated country, they will be little liable to modification; for neither migration nor isolation in themselves can do anything.

These principles come into play only by bringing organisms into new relations with each other, and in a lesser degree with the surrounding physical conditions.

As we have seen in the last chapter that some forms have retained nearly the same character from an enormously remote geological period, so certain species have migrated over vast spaces, and have not become greatly modified.
03 - Struggle for Existence 03-03 - Geometrical Ratio of Increase 10 A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase.

Every being, which during its natural lifetime produces several eggs or seeds, must suffer destruction during some period of its life, and during some season or occasional year, otherwise, on the principle of geometrical increase, its numbers would quickly become so inordinately great that no country could support the product.

egg
egg

seeds
seeds


Hence, as more individuals are produced than can possibly survive, there must in every case be a struggle for existence, either one individual with another of the same species, or with the individuals of distinct species, or with the physical conditions of life.

It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms; for in this case there can be no artificial increase of food, and no prudential restraint from marriage.

Although some species may be now increasing, more or less rapidly, in numbers, all cannot do so, for the world would not hold them.