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<h2> CHAPTER XII. GEOGRAPHICAL DISTRIBUTION. </h2>
<p>Present distribution cannot be accounted for by differences in physical<br/>
conditions—Importance of barriers—Affinity of the productions of the<br/>
same continent—Centres of creation—Means of dispersal by changes of<br/>
climate and of the level of the land, and by occasional means—Dispersal<br/>
during the Glacial period—Alternate Glacial periods in the North and<br/>
South.<br/></p>
<p>In considering the distribution of organic beings over the face of the
globe, the first great fact which strikes us is, that neither the
similarity nor the dissimilarity of the inhabitants of various regions can
be wholly accounted for by climatal and other physical conditions. Of
late, almost every author who has studied the subject has come to this
conclusion. The case of America alone would almost suffice to prove its
truth; for if we exclude the arctic and northern temperate parts, all
authors agree that one of the most fundamental divisions in geographical
distribution is that between the New and Old Worlds; yet if we travel over
the vast American continent, from the central parts of the United States
to its extreme southern point, we meet with the most diversified
conditions; humid districts, arid deserts, lofty mountains, grassy plains,
forests, marshes, lakes and great rivers, under almost every temperature.
There is hardly a climate or condition in the Old World which cannot be
paralleled in the New—at least so closely as the same species
generally require. No doubt small areas can be pointed out in the Old
World hotter than any in the New World; but these are not inhabited by a
fauna different from that of the surrounding districts; for it is rare to
find a group of organisms confined to a small area, of which the
conditions are peculiar in only a slight degree. Notwithstanding this
general parallelism in the conditions of Old and New Worlds, how widely
different are their living productions!</p>
<p>In the southern hemisphere, if we compare large tracts of land in
Australia, South Africa, and western South America, between latitudes 25
and 35 degrees, we shall find parts extremely similar in all their
conditions, yet it would not be possible to point out three faunas and
floras more utterly dissimilar. Or, again, we may compare the productions
of South America south of latitude 35 degrees with those north of 25
degrees, which consequently are separated by a space of ten degrees of
latitude, and are exposed to considerably different conditions; yet they
are incomparably more closely related to each other than they are to the
productions of Australia or Africa under nearly the same climate.
Analogous facts could be given with respect to the inhabitants of the sea.</p>
<p>A second great fact which strikes us in our general review is, that
barriers of any kind, or obstacles to free migration, are related in a
close and important manner to the differences between the productions of
various regions. We see this in the great difference in nearly all the
terrestrial productions of the New and Old Worlds, excepting in the
northern parts, where the land almost joins, and where, under a slightly
different climate, there might have been free migration for the northern
temperate forms, as there now is for the strictly arctic productions. We
see the same fact in the great difference between the inhabitants of
Australia, Africa, and South America under the same latitude; for these
countries are almost as much isolated from each other as is possible. On
each continent, also, we see the same fact; for on the opposite sides of
lofty and continuous mountain-ranges, and of great deserts and even of
large rivers, we find different productions; though as mountain chains,
deserts, etc., are not as impassable, or likely to have endured so long,
as the oceans separating continents, the differences are very inferior in
degree to those characteristic of distinct continents.</p>
<p>Turning to the sea, we find the same law. The marine inhabitants of the
eastern and western shores of South America are very distinct, with
extremely few shells, crustacea, or echinodermata in common; but Dr.
Gunther has recently shown that about thirty per cent of the fishes are
the same on the opposite sides of the isthmus of Panama; and this fact has
led naturalists to believe that the isthmus was formerly open. Westward of
the shores of America, a wide space of open ocean extends, with not an
island as a halting-place for emigrants; here we have a barrier of another
kind, and as soon as this is passed we meet in the eastern islands of the
Pacific with another and totally distinct fauna. So that three marine
faunas range northward and southward in parallel lines not far from each
other, under corresponding climate; but from being separated from each
other by impassable barriers, either of land or open sea, they are almost
wholly distinct. On the other hand, proceeding still further westward from
the eastern islands of the tropical parts of the Pacific, we encounter no
impassable barriers, and we have innumerable islands as halting-places, or
continuous coasts, until, after travelling over a hemisphere, we come to
the shores of Africa; and over this vast space we meet with no
well-defined and distinct marine faunas. Although so few marine animals
are common to the above-named three approximate faunas of Eastern and
Western America and the eastern Pacific islands, yet many fishes range
from the Pacific into the Indian Ocean, and many shells are common to the
eastern islands of the Pacific and the eastern shores of Africa on almost
exactly opposite meridians of longitude.</p>
<p>A third great fact, partly included in the foregoing statement, is the
affinity of the productions of the same continent or of the same 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, though nearly
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. 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 emu,
like those inhabiting Africa and Australia under the same latitude. 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. 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 muskrat, but the coypu and capybara, rodents of the South
American type. 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 are essentially American, though
they may be all peculiar species. We may look back to past ages, as shown
in the last chapter, and we find American types then prevailing on the
American continent and in the American seas. We see in these facts some
deep organic bond, throughout space and time, over the same areas of land
and water, independently of physical conditions. The naturalist must be
dull who is not led to inquire what this bond is.</p>
<p>The bond is simply inheritance, that cause which alone, as far as we
positively know, produces organisms quite like each other, or, as we see
in the case of varieties, nearly alike. The dissimilarity of the
inhabitants of different regions may be attributed to modification through
variation and natural selection, and probably in a subordinate degree to
the definite influence of different physical conditions. The degrees of
dissimilarity will depend on the migration of the more dominant forms of
life from one region into another having been more or less effectually
prevented, at periods more or less remote—on the nature and number
of the former immigrants—and on the action of the inhabitants on
each other in leading to the preservation of different modifications; the
relation of organism to organism in the struggle for life 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 out 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.</p>
<p>There is no evidence, as was remarked in the last chapter, of the
existence of any 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 each individual in its
complex struggle for life, so the amount of modification in different
species will be no uniform quantity. If a number of species, after having
long competed with each other in their old home, were to migrate in a body
into a new and afterwards isolated country, they would be little liable to
modification; for neither migration nor isolation in themselves effect
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 or at all modified.</p>
<p>According to these views, it is obvious that the several species of the
same genus, though inhabiting the most distant quarters of the world, must
originally have proceeded from the same source, as they are descended from
the same progenitor. In the case of those species which have undergone,
during whole geological periods, little modification, there is not much
difficulty in believing that they have migrated from the same region; for
during the vast geographical and climatical changes which have supervened
since ancient times, almost any amount of migration is possible. But in
many other cases, in which we have reason to believe that the species of a
genus have been produced within comparatively recent times, there is great
difficulty on this head. It is also obvious that the individuals of the
same species, though now inhabiting distant and isolated regions, must
have proceeded from one spot, where their parents were first produced:
for, as has been explained, it is incredible that individuals identically
the same should have been produced from parents specifically distinct.</p>
<p>SINGLE CENTRES OF SUPPOSED CREATION.</p>
<p>We are thus brought to the question which has been largely discussed by
naturalists, namely, whether species have been created at one or more
points of the earth's surface. Undoubtedly there are many cases of extreme
difficulty in understanding how the same species could possibly have
migrated from some one point to the several distant and isolated points,
where now found. Nevertheless the simplicity of the view that each species
was first produced within a single region captivates the mind. He who
rejects it, rejects the vera causa of ordinary generation with subsequent
migration, and calls in the agency of a miracle. It is universally
admitted, that in most cases the area inhabited by a species is
continuous; and that when a plant or animal inhabits two points so distant
from each other, or with an interval of such a nature, that the space
could not have been easily passed over by migration, the fact is given as
something remarkable and exceptional. The incapacity of migrating across a
wide sea is more clear in the case of terrestrial mammals than perhaps
with any other organic beings; and, accordingly, we find no inexplicable
instances of the same mammals inhabiting distant points of the world. No
geologist feels any difficulty in Great Britain possessing the same
quadrupeds with the rest of Europe, for they were no doubt once united.
But if the same species can be produced at two separate points, why do we
not find a single mammal common to Europe and Australia or South America?
The conditions of life are nearly the same, so that a multitude of
European animals and plants have become naturalised in America and
Australia; and some of the aboriginal plants are identically the same at
these distant points of the northern and southern hemispheres? The answer,
as I believe, is, that mammals have not been able to migrate, whereas some
plants, from their varied means of dispersal, have migrated across the
wide and broken interspaces. The great and striking influence of barriers
of all kinds, is intelligible only on the view that the great majority of
species have been produced on one side, and have not been able to migrate
to the opposite side. Some few families, many subfamilies, very many
genera, a still greater number of sections of genera, are confined to a
single region; and it has been observed by several naturalists that the
most natural genera, or those genera in which the species are most closely
related to each other, are generally confined to the same country, or if
they have a wide range that their range is continuous. What a strange
anomaly it would be if a directly opposite rule were to prevail when we go
down one step lower in the series, namely to the individuals of the same
species, and these had not been, at least at first, confined to some one
region!</p>
<p>Hence, it seems to me, as it has to many other naturalists, that the view
of each species having been produced in one area alone, and having
subsequently migrated from that area as far as its powers of migration and
subsistence under past and present conditions permitted, is the most
probable. Undoubtedly many cases occur in which we cannot explain how the
same species could have passed from one point to the other. But the
geographical and climatical changes which have certainly occurred within
recent geological times, must have rendered discontinuous the formerly
continuous range of many species. So that we are reduced to consider
whether the exceptions to continuity of range are so numerous, and of so
grave a nature, that we ought to give up the belief, rendered probable by
general considerations, that each species has been produced within one
area, and has migrated thence as far as it could. It would be hopelessly
tedious to discuss all the exceptional cases of the same species, now
living at distant and separated points; nor do I for a moment pretend that
any explanation could be offered of many instances. But, after some
preliminary remarks, I will discuss a few of the most striking classes of
facts, namely, the existence of the same species on the summits of distant
mountain ranges, and at distant points in the Arctic and Antarctic
regions; and secondly (in the following chapter), the wide distribution of
fresh water productions; and thirdly, the occurrence of the same
terrestrial species on islands and on the nearest mainland, though
separated by hundreds of miles of open sea. If the existence of the same
species at distant and isolated points of the earth's surface can in many
instances be explained on the view of each species having migrated from a
single birthplace; then, considering our ignorance with respect to former
climatical and geographical changes, and to the various occasional means
of transport, the belief that a single birthplace is the law seems to me
incomparably the safest.</p>
<p>In discussing this subject we shall be enabled at the same time to
consider a point equally important for us, namely, whether the several
species of a genus which must on our theory all be descended from a common
progenitor, can have migrated, undergoing modification during their
migration from some one area. If, when most of the species inhabiting one
region are different from those of another region, though closely allied
to them, it can be shown that migration from the one region to the other
has probably occurred at some former period, our general view will be much
strengthened; for the explanation is obvious on the principle of descent
with modification. A volcanic island, for instance, upheaved and formed at
the distance of a few hundreds of miles from a continent, would probably
receive from it in the course of time a few colonists, and their
descendants, though modified, would still be related by inheritance to the
inhabitants of that continent. Cases of this nature are common, and are,
as we shall hereafter see, inexplicable on the theory of independent
creation. This view of the relation of the species of one region to those
of another, does not differ much from that advanced by Mr. Wallace, who
concludes that "every species has come into existence coincident both in
space and time with a pre-existing closely allied species." And it is now
well known that he attributes this coincidence to descent with
modification.</p>
<p>The question of single or multiple centres of creation differs from
another though allied question, namely, whether all the individuals of the
same species are descended from a single pair, or single hermaphrodite, or
whether, as some authors suppose, from many individuals simultaneously
created. With organic beings which never intercross, if such exist, each
species, must be descended from a succession of modified varieties, that
have supplanted each other, but have never blended with other individuals
or varieties of the same species, so that, at each successive stage of
modification, all the individuals of the same form will be descended from
a single parent. But in the great majority of cases, namely, with all
organisms which habitually unite for each birth, or which occasionally
intercross, the individuals of the same species inhabiting the same area
will be kept nearly uniform by intercrossing; so that many individuals
will go on simultaneously changing, and the whole amount of modification
at each stage will not be due to descent from a single parent. To
illustrate what I mean: our English race-horses differ from the horses of
every other breed; but they do not owe their difference and superiority to
descent from any single pair, but to continued care in the selecting and
training of many individuals during each generation.</p>
<p>Before discussing the three classes of facts, which I have selected as
presenting the greatest amount of difficulty on the theory of "single
centres of creation," I must say a few words on the means of dispersal.</p>
<p>MEANS OF DISPERSAL.</p>
<p>Sir C. Lyell and other authors have ably treated this subject. I can give
here only the briefest abstract of the more important facts. Change of
climate must have had a powerful influence on migration. A region now
impassable to certain organisms from the nature of its climate, might have
been a high road for migration, when the climate was different. I shall,
however, presently have to discuss this branch of the subject in some
detail. Changes of level in the land must also have been highly
influential: a narrow isthmus now separates two marine faunas; submerge
it, or let it formerly have been submerged, and the two faunas will now
blend together, or may formerly have blended. Where the sea now extends,
land may at a former period have connected islands or possibly even
continents together, and thus have allowed terrestrial productions to pass
from one to the other. No geologist disputes that great mutations of level
have occurred within the period of existing organisms. Edward Forbes
insisted that all the islands in the Atlantic must have been recently
connected with Europe or Africa, and Europe likewise with America. Other
authors have thus hypothetically bridged over every ocean, and united
almost every island with some mainland. If, indeed, the arguments used by
Forbes are to be trusted, it must be admitted that scarcely a single
island exists which has not recently been united to some continent. This
view cuts the Gordian knot of the dispersal of the same species to the
most distant points, and removes many a difficulty; but to the best of my
judgment we are not authorized in admitting such enormous geographical
changes within the period of existing species. It seems to me that we have
abundant evidence of great oscillations in the level of the land or sea;
but not of such vast changes in the position and extension of our
continents, as to have united them within the recent period to each other
and to the several intervening oceanic islands. I freely admit the former
existence of many islands, now buried beneath the sea, which may have
served as halting places for plants and for many animals during their
migration. In the coral-producing oceans such sunken islands are now
marked by rings of coral or atolls standing over them. Whenever it is
fully admitted, as it will some day be, that each species has proceeded
from a single birthplace, and when in the course of time we know something
definite about the means of distribution, we shall be enabled to speculate
with security on the former extension of the land. But I do not believe
that it will ever be proved that within the recent period most of our
continents which now stand quite separate, have been continuously, or
almost continuously united with each other, and with the many existing
oceanic islands. Several facts in distribution—such as the great
difference in the marine faunas on the opposite sides of almost every
continent—the close relation of the tertiary inhabitants of several
lands and even seas to their present inhabitants—the degree of
affinity between the mammals inhabiting islands with those of the nearest
continent, being in part determined (as we shall hereafter see) by the
depth of the intervening ocean—these and other such facts are
opposed to the admission of such prodigious geographical revolutions
within the recent period, as are necessary on the view advanced by Forbes
and admitted by his followers. The nature and relative proportions of the
inhabitants of oceanic islands are likewise opposed to the belief of their
former continuity of continents. Nor does the almost universally volcanic
composition of such islands favour the admission that they are the wrecks
of sunken continents; if they had originally existed as continental
mountain ranges, some at least of the islands would have been formed, like
other mountain summits, of granite, metamorphic schists, old fossiliferous
and other rocks, instead of consisting of mere piles of volcanic matter.</p>
<p>I must now say a few words on what are called accidental means, but which
more properly should be called occasional means of distribution. I shall
here confine myself to plants. In botanical works, this or that plant is
often stated to be ill adapted for wide dissemination; but the greater or
less facilities for transport across the sea may be said to be almost
wholly unknown. Until I tried, with Mr. Berkeley's aid, a few experiments,
it was not even known how far seeds could resist the injurious action of
sea-water. To my surprise I found that out of eighty-seven kinds,
sixty-four germinated after an immersion of twenty-eight days, and a few
survived an immersion of 137 days. It deserves notice that certain orders
were far more injured than others: nine Leguminosae were tried, and, with
one exception, they resisted the salt-water badly; seven species of the
allied orders, Hydrophyllaceae and Polemoniaceae, were all killed by a
month's immersion. For convenience sake I chiefly tried small seeds
without the capsules or fruit; and as all of these sank in a few days,
they could not have been floated across wide spaces of the sea, whether or
not they were injured by salt water. Afterwards I tried some larger
fruits, capsules, etc., and some of these floated for a long time. It is
well known what a difference there is in the buoyancy of green and
seasoned timber; and it occurred to me that floods would often wash into
the sea dried plants or branches with seed-capsules or fruit attached to
them. Hence I was led to dry the stems and branches of ninety-four plants
with ripe fruit, and to place them on sea-water. The majority sank
quickly, but some which, whilst green, floated for a very short time, when
dried floated much longer; for instance, ripe hazel-nuts sank immediately,
but when dried they floated for ninety days, and afterwards when planted
germinated; an asparagus plant with ripe berries floated for twenty-three
days, when dried it floated for eighty-five days, and the seeds afterwards
germinated: the ripe seeds of Helosciadium sank in two days, when dried
they floated for above ninety days, and afterwards germinated. Altogether,
out of the ninety-four dried plants, eighteen floated for above
twenty-eight days; and some of the eighteen floated for a very much longer
period. So that as 64/87 kinds of seeds germinated after an immersion of
twenty-eight days; and as 18/94 distinct species with ripe fruit (but not
all the same species as in the foregoing experiment) floated, after being
dried, for above twenty-eight days, we may conclude, as far as anything
can be inferred from these scanty facts, that the seeds of 14/100 kinds of
plants of any country might be floated by sea-currents during twenty-eight
days, and would retain their power of germination. In Johnston's Physical
Atlas, the average rate of the several Atlantic currents is thirty-three
miles per diem (some currents running at the rate of sixty miles per
diem); on this average, the seeds of 14/100 plants belonging to one
country might be floated across 924 miles of sea to another country; and
when stranded, if blown by an inland gale to a favourable spot, would
germinate.</p>
<p>Subsequently to my experiments, M. Martens tried similar ones, but in a
much better manner, for he placed the seeds in a box in the actual sea, so
that they were alternately wet and exposed to the air like really floating
plants. He tried ninety-eight seeds, mostly different from mine, but he
chose many large fruits, and likewise seeds, from plants which live near
the sea; and this would have favoured both the average length of their
flotation and their resistance to the injurious action of the salt-water.
On the other hand, he did not previously dry the plants or branches with
the fruit; and this, as we have seen, would have caused some of them to
have floated much longer. The result was that 18/98 of his seeds of
different kinds floated for forty-two days, and were then capable of
germination. But I do not doubt that plants exposed to the waves would
float for a less time than those protected from violent movement as in our
experiments. Therefore, it would perhaps be safer to assume that the seeds
of about 10/100 plants of a flora, after having been dried, could be
floated across a space of sea 900 miles in width, and would then
germinate. The fact of the larger fruits often floating longer than the
small, is interesting; as plants with large seeds or fruit which, as Alph.
de Candolle has shown, generally have restricted ranges, could hardly be
transported by any other means.</p>
<p>Seeds may be occasionally transported in another manner. Drift timber is
thrown up on most islands, even on those in the midst of the widest
oceans; and the natives of the coral islands in the Pacific procure stones
for their tools, solely from the roots of drifted trees, these stones
being a valuable royal tax. I find that when irregularly shaped stones are
embedded in the roots of trees, small parcels of earth are very frequently
enclosed in their interstices and behind them, so perfectly that not a
particle could be washed away during the longest transport: out of one
small portion of earth thus COMPLETELY enclosed by the roots of an oak
about fifty years old, three dicotyledonous plants germinated: I am
certain of the accuracy of this observation. Again, I can show that the
carcasses of birds, when floating on the sea, sometimes escape being
immediately devoured; and many kinds of seeds in the crops of floating
birds long retain their vitality: peas and vetches, for instance, are
killed by even a few days' immersion in sea-water; but some taken out of
the crop of a pigeon, which had floated on artificial sea-water for thirty
days, to my surprise nearly all germinated.</p>
<p>Living birds can hardly fail to be highly effective agents in the
transportation of seeds. I could give many facts showing how frequently
birds of many kinds are blown by gales to vast distances across the ocean.
We may safely assume that under such circumstances their rate of flight
would often be thirty-five miles an hour; and some authors have given a
far higher estimate. I have never seen an instance of nutritious seeds
passing through the intestines of a bird; but hard seeds of fruit pass
uninjured through even the digestive organs of a turkey. In the course of
two months, I picked up in my garden twelve kinds of seeds, out of the
excrement of small birds, and these seemed perfect, and some of them,
which were tried, germinated. But the following fact is more important:
the crops of birds do not secrete gastric juice, and do not, as I know by
trial, injure in the least the germination of seeds; now, after a bird has
found and devoured a large supply of food, it is positively asserted that
all the grains do not pass into the gizzard for twelve or even eighteen
hours. A bird in this interval might easily be blown to the distance of
five hundred miles, and hawks are known to look out for tired birds, and
the contents of their torn crops might thus readily get scattered. Some
hawks and owls bolt their prey whole, and after an interval of from twelve
to twenty hours, disgorge pellets, which, as I know from experiments made
in the Zoological Gardens, include seeds capable of germination. Some
seeds of the oat, wheat, millet, canary, hemp, clover, and beet germinated
after having been from twelve to twenty-one hours in the stomachs of
different birds of prey; and two seeds of beet grew after having been thus
retained for two days and fourteen hours. Fresh-water fish, I find, eat
seeds of many land and water plants; fish are frequently devoured by
birds, and thus the seeds might be transported from place to place. I
forced many kinds of seeds into the stomachs of dead fish, and then gave
their bodies to fishing-eagles, storks, and pelicans; these birds, after
an interval of many hours, either rejected the seeds in pellets or passed
them in their excrement; and several of these seeds retained the power of
germination. Certain seeds, however, were always killed by this process.</p>
<p>Locusts are sometimes blown to great distances from the land. I myself
caught one 370 miles from the coast of Africa, and have heard of others
caught at greater distances. The Rev. R.T. Lowe informed Sir C. Lyell that
in November, 1844, swarms of locusts visited the island of Madeira. They
were in countless numbers, as thick as the flakes of snow in the heaviest
snowstorm, and extended upward as far as could be seen with a telescope.
During two or three days they slowly careered round and round in an
immense ellipse, at least five or six miles in diameter, and at night
alighted on the taller trees, which were completely coated with them. They
then disappeared over the sea, as suddenly as they had appeared, and have
not since visited the island. Now, in parts of Natal it is believed by
some farmers, though on insufficient evidence, that injurious seeds are
introduced into their grass-land in the dung left by the great flights of
locusts which often visit that country. In consequence of this belief Mr.
Weale sent me in a letter a small packet of the dried pellets, out of
which I extracted under the microscope several seeds, and raised from them
seven grass plants, belonging to two species, of two genera. Hence a swarm
of locusts, such as that which visited Madeira, might readily be the means
of introducing several kinds of plants into an island lying far from the
mainland.</p>
<p>Although the beaks and feet of birds are generally clean, earth sometimes
adheres to them: in one case I removed sixty-one grains, and in another
case twenty-two grains of dry argillaceous earth from the foot of a
partridge, and in the earth there was a pebble as large as the seed of a
vetch. Here is a better case: the leg of a woodcock was sent to me by a
friend, with a little cake of dry earth attached to the shank, weighing
only nine grains; and this contained a seed of the toad-rush (Juncus
bufonius) which germinated and flowered. Mr. Swaysland, of Brighton, who
during the last forty years has paid close attention to our migratory
birds, informs me that he has often shot wagtails (Motacillae), wheatears,
and whinchats (Saxicolae), on their first arrival on our shores, before
they had alighted; and he has several times noticed little cakes of earth
attached to their feet. Many facts could be given showing how generally
soil is charged with seeds. For instance, Professor Newton sent me the leg
of a red-legged partridge (Caccabis rufa) which had been wounded and could
not fly, with a ball of hard earth adhering to it, and weighing six and a
half ounces. The earth had been kept for three years, but when broken,
watered and placed under a bell glass, no less than eighty-two plants
sprung from it: these consisted of twelve monocotyledons, including the
common oat, and at least one kind of grass, and of seventy dicotyledons,
which consisted, judging from the young leaves, of at least three distinct
species. With such facts before us, can we doubt that the many birds which
are annually blown by gales across great spaces of ocean, and which
annually migrate—for instance, the millions of quails across the
Mediterranean—must occasionally transport a few seeds embedded in
dirt adhering to their feet or beaks? But I shall have to recur to this
subject.</p>
<p>As icebergs are known to be sometimes loaded with earth and stones, and
have even carried brushwood, bones, and the nest of a land-bird, it can
hardly be doubted that they must occasionally, as suggested by Lyell, have
transported seeds from one part to another of the arctic and antarctic
regions; and during the Glacial period from one part of the now temperate
regions to another. In the Azores, from the large number of plants common
to Europe, in comparison with the species on the other islands of the
Atlantic, which stand nearer to the mainland, and (as remarked by Mr. H.C.
Watson) from their somewhat northern character, in comparison with the
latitude, I suspected that these islands had been partly stocked by
ice-borne seeds during the Glacial epoch. At my request Sir C. Lyell wrote
to M. Hartung to inquire whether he had observed erratic boulders on these
islands, and he answered that he had found large fragments of granite and
other rocks, which do not occur in the archipelago. Hence we may safely
infer that icebergs formerly landed their rocky burdens on the shores of
these mid-ocean islands, and it is at least possible that they may have
brought thither the seeds of northern plants.</p>
<p>Considering that these several means of transport, and that other means,
which without doubt remain to be discovered, have been in action year
after year for tens of thousands of years, it would, I think, be a
marvellous fact if many plants had not thus become widely transported.
These means of transport are sometimes called accidental, but this is not
strictly correct: the currents of the sea are not accidental, nor is the
direction of prevalent gales of wind. It should be observed that scarcely
any means of transport would carry seeds for very great distances; for
seeds do not retain their vitality when exposed for a great length of time
to the action of sea water; nor could they be long carried in the crops or
intestines of birds. These means, however, would suffice for occasional
transport across tracts of sea some hundred miles in breadth, or from
island to island, or from a continent to a neighbouring island, but not
from one distant continent to another. The floras of distant continents
would not by such means become mingled; but would remain as distinct as
they now are. The currents, from their course, would never bring seeds
from North America to Britain, though they might and do bring seeds from
the West Indies to our western shores, where, if not killed by their very
long immersion in salt water, they could not endure our climate. Almost
every year, one or two land-birds are blown across the whole Atlantic
Ocean, from North America to the western shores of Ireland and England;
but seeds could be transported by these rare wanderers only by one means,
namely, by dirt adhering to their feet or beaks, which is in itself a rare
accident. Even in this case, how small would be the chance of a seed
falling on favourable soil, and coming to maturity! But it would be a
great error to argue that because a well-stocked island, like Great
Britain, has not, as far as is known (and it would be very difficult to
prove this), received within the last few centuries, through occasional
means of transport, immigrants from Europe or any other continent, that a
poorly-stocked island, though standing more remote from the mainland,
would not receive colonists by similar means. Out of a hundred kinds of
seeds or animals transported to an island, even if far less well-stocked
than Britain, perhaps not more than one would be so well fitted to its new
home, as to become naturalised. But this is no valid argument against what
would be effected by occasional means of transport, during the long lapse
of geological time, whilst the island was being upheaved, and before it
had become fully stocked with inhabitants. On almost bare land, with few
or no destructive insects or birds living there, nearly every seed which
chanced to arrive, if fitted for the climate, would germinate and survive.</p>
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