Books: On the Origin of Species
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Charles Darwin >> On the Origin of Species
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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 I think safely assume that under such
circumstances their rate of flight would often be 35 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 will pass uninjured through even the
digestive organs of a turkey. In the course of two months, I picked up
in my garden 12 kinds of seeds, out of the excrement of small birds,
and these seemed perfect, and some of them, which I tried, germinated.
But the following fact is more important: the crops of birds do not
secrete gastric juice, and do not in the least injure, as I know by
trial, 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 12 or even 18 hours. A
bird in this interval might easily be blown to the distance of 500
miles, and hawks are known to look out for tired birds, and the
contents of their torn crops might thus readily get scattered. Mr.
Brent informs me that a friend of his had to give up flying
carrier-pigeons from France to England, as the hawks on the English
coast destroyed so many on their arrival. 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. Freshwater 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 their power of germination. Certain seeds, however,
were always killed by this process.
Although the beaks and feet of birds are generally quite clean, I can
show that earth sometimes adheres to them: in one instance I removed
twenty-two grains of dry argillaceous earth from one foot of a
partridge, and in this earth there was a pebble quite as large as the
seed of a vetch. Thus seeds might occasionally be transported to great
distances; for many facts could be given showing that soil almost
everywhere is charged with seeds. Reflect for a moment on the millions
of quails which annually cross the Mediterranean; and can we doubt
that the earth adhering to their feet would sometimes include a few
minute seeds? But I shall presently have to recur to this subject.
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, I
can hardly doubt that they must occasionally have transported seeds
from one part to another of the arctic and antarctic regions, as
suggested by Lyell; and during the Glacial period from one part of the
now temperate regions to another. In the Azores, from the large number
of the species of plants common to Europe, in comparison with the
plants of other oceanic islands nearer to the mainland, and (as
remarked by Mr. H. C. Watson) from the somewhat northern character of
the flora 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 burthens 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.
Considering that the several above means of transport, and that
several other means, which without doubt remain to be discovered, have
been in action year after year, for centuries and 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
seawater; 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 in any great degree; but would remain
as distinct as we now see them to be. 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 so long an 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 wanderers only by one means, namely, in dirt sticking to
their feet, which is in itself a rare accident. Even in this case, how
small would the chance be 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. I do not doubt that out of twenty
seeds or animals transported to an island, even if far less
well-stocked than Britain, scarcely more than one would be so well
fitted to its new home, as to become naturalised. But this, as it
seems to me, is no valid argument against what would be effected by
occasional means of transport, during the long lapse of geological
time, whilst an island was being upheaved and formed, 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, would be sure to germinate and survive.
DISPERSAL DURING THE GLACIAL PERIOD.
The identity of many plants and animals, on mountain-summits,
separated from each other by hundreds of miles of lowlands, where the
Alpine species could not possibly exist, is one of the most striking
cases known of the same species living at distant points, without the
apparent possibility of their having migrated from one to the other.
It is indeed a remarkable fact to see so many of the same plants
living on the snowy regions of the Alps or Pyrenees, and in the
extreme northern parts of Europe; but it is far more remarkable, that
the plants on the White Mountains, in the United States of America,
are all the same with those of Labrador, and nearly all the same, as
we hear from Asa Gray, with those on the loftiest mountains of Europe.
Even as long ago as 1747, such facts led Gmelin to conclude that the
same species must have been independently created at several distinct
points; and we might have remained in this same belief, had not
Agassiz and others called vivid attention to the Glacial period,
which, as we shall immediately see, affords a simple explanation of
these facts. We have evidence of almost every conceivable kind,
organic and inorganic, that within a very recent geological period,
central Europe and North America suffered under an Arctic climate. The
ruins of a house burnt by fire do not tell their tale more plainly,
than do the mountains of Scotland and Wales, with their scored flanks,
polished surfaces, and perched boulders, of the icy streams with which
their valleys were lately filled. So greatly has the climate of Europe
changed, that in Northern Italy, gigantic moraines, left by old
glaciers, are now clothed by the vine and maize. Throughout a large
part of the United States, erratic boulders, and rocks scored by
drifted icebergs and coast-ice, plainly reveal a former cold period.
The former influence of the glacial climate on the distribution of the
inhabitants of Europe, as explained with remarkable clearness by
Edward Forbes, is substantially as follows. But we shall follow the
changes more readily, by supposing a new glacial period to come slowly
on, and then pass away, as formerly occurred. As the cold came on, and
as each more southern zone became fitted for arctic beings and
ill-fitted for their former more temperate inhabitants, the latter
would be supplanted and arctic productions would take their places.
The inhabitants of the more temperate regions would at the same time
travel southward, unless they were stopped by barriers, in which case
they would perish. The mountains would become covered with snow and
ice, and their former Alpine inhabitants would descend to the plains.
By the time that the cold had reached its maximum, we should have a
uniform arctic fauna and flora, covering the central parts of Europe,
as far south as the Alps and Pyrenees, and even stretching into Spain.
The now temperate regions of the United States would likewise be
covered by arctic plants and animals, and these would be nearly the
same with those of Europe; for the present circumpolar inhabitants,
which we suppose to have everywhere travelled southward, are
remarkably uniform round the world. We may suppose that the Glacial
period came on a little earlier or later in North America than in
Europe, so will the southern migration there have been a little
earlier or later; but this will make no difference in the final
result.
As the warmth returned, the arctic forms would retreat northward,
closely followed up in their retreat by the productions of the more
temperate regions. And as the snow melted from the bases of the
mountains, the arctic forms would seize on the cleared and thawed
ground, always ascending higher and higher, as the warmth increased,
whilst their brethren were pursuing their northern journey. Hence,
when the warmth had fully returned, the same arctic species, which had
lately lived in a body together on the lowlands of the Old and New
Worlds, would be left isolated on distant mountain-summits (having
been exterminated on all lesser heights) and in the arctic regions of
both hemispheres.
Thus we can understand the identity of many plants at points so
immensely remote as on the mountains of the United States and of
Europe. We can thus also understand the fact that the Alpine plants of
each mountain-range are more especially related to the arctic forms
living due north or nearly due north of them: for the migration as the
cold came on, and the re-migration on the returning warmth, will
generally have been due south and north. The Alpine plants, for
example, of Scotland, as remarked by Mr. H. C. Watson, and those of
the Pyrenees, as remarked by Ramond, are more especially allied to the
plants of northern Scandinavia; those of the United States to
Labrador; those of the mountains of Siberia to the arctic regions of
that country. These views, grounded as they are on the perfectly
well-ascertained occurrence of a former Glacial period, seem to me to
explain in so satisfactory a manner the present distribution of the
Alpine and Arctic productions of Europe and America, that when in
other regions we find the same species on distant mountain-summits, we
may almost conclude without other evidence, that a colder climate
permitted their former migration across the low intervening tracts,
since become too warm for their existence.
If the climate, since the Glacial period, has ever been in any degree
warmer than at present (as some geologists in the United States
believe to have been the case, chiefly from the distribution of the
fossil Gnathodon), then the arctic and temperate productions will at a
very late period have marched a little further north, and subsequently
have retreated to their present homes; but I have met with no
satisfactory evidence with respect to this intercalated slightly
warmer period, since the Glacial period.
The arctic forms, during their long southern migration and
re-migration northward, will have been exposed to nearly the same
climate, and, as is especially to be noticed, they will have kept in a
body together; consequently their mutual relations will not have been
much disturbed, and, in accordance with the principles inculcated in
this volume, they will not have been liable to much modification. But
with our Alpine productions, left isolated from the moment of the
returning warmth, first at the bases and ultimately on the summits of
the mountains, the case will have been somewhat different; for it is
not likely that all the same arctic species will have been left on
mountain ranges distant from each other, and have survived there ever
since; they will, also, in all probability have become mingled with
ancient Alpine species, which must have existed on the mountains
before the commencement of the Glacial epoch, and which during its
coldest period will have been temporarily driven down to the plains;
they will, also, have been exposed to somewhat different climatal
influences. Their mutual relations will thus have been in some degree
disturbed; consequently they will have been liable to modification;
and this we find has been the case; for if we compare the present
Alpine plants and animals of the several great European
mountain-ranges, though very many of the species are identically the
same, some present varieties, some are ranked as doubtful forms, and
some few are distinct yet closely allied or representative species.
In illustrating what, as I believe, actually took place during the
Glacial period, I assumed that at its commencement the arctic
productions were as uniform round the polar regions as they are at the
present day. But the foregoing remarks on distribution apply not only
to strictly arctic forms, but also to many sub-arctic and to some few
northern temperate forms, for some of these are the same on the lower
mountains and on the plains of North America and Europe; and it may be
reasonably asked how I account for the necessary degree of uniformity
of the sub-arctic and northern temperate forms round the world, at the
commencement of the Glacial period. At the present day, the sub-arctic
and northern temperate productions of the Old and New Worlds are
separated from each other by the Atlantic Ocean and by the extreme
northern part of the Pacific. During the Glacial period, when the
inhabitants of the Old and New Worlds lived further southwards than at
present, they must have been still more completely separated by wider
spaces of ocean. I believe the above difficulty may be surmounted by
looking to still earlier changes of climate of an opposite nature. We
have good reason to believe that during the newer Pliocene period,
before the Glacial epoch, and whilst the majority of the inhabitants
of the world were specifically the same as now, the climate was warmer
than at the present day. Hence we may suppose that the organisms now
living under the climate of latitude 60 deg, during the Pliocene
period lived further north under the Polar Circle, in latitude 66
deg-67 deg; and that the strictly arctic productions then lived on the
broken land still nearer to the pole. Now if we look at a globe, we
shall see that under the Polar Circle there is almost continuous land
from western Europe, through Siberia, to eastern America. And to this
continuity of the circumpolar land, and to the consequent freedom for
intermigration under a more favourable climate, I attribute the
necessary amount of uniformity in the sub-arctic and northern
temperate productions of the Old and New Worlds, at a period anterior
to the Glacial epoch.
Believing, from reasons before alluded to, that our continents have
long remained in nearly the same relative position, though subjected
to large, but partial oscillations of level, I am strongly inclined to
extend the above view, and to infer that during some earlier and still
warmer period, such as the older Pliocene period, a large number of
the same plants and animals inhabited the almost continuous
circumpolar land; and that these plants and animals, both in the Old
and New Worlds, began slowly to migrate southwards as the climate
became less warm, long before the commencement of the Glacial period.
We now see, as I believe, their descendants, mostly in a modified
condition, in the central parts of Europe and the United States. On
this view we can understand the relationship, with very little
identity, between the productions of North America and Europe,--a
relationship which is most remarkable, considering the distance of the
two areas, and their separation by the Atlantic Ocean. We can further
understand the singular fact remarked on by several observers, that
the productions of Europe and America during the later tertiary stages
were more closely related to each other than they are at the present
time; for during these warmer periods the northern parts of the Old
and New Worlds will have been almost continuously united by land,
serving as a bridge, since rendered impassable by cold, for the
inter-migration of their inhabitants.
During the slowly decreasing warmth of the Pliocene period, as soon as
the species in common, which inhabited the New and Old Worlds,
migrated south of the Polar Circle, they must have been completely cut
off from each other. This separation, as far as the more temperate
productions are concerned, took place long ages ago. And as the plants
and animals migrated southward, they will have become mingled in the
one great region with the native American productions, and have had to
compete with them; and in the other great region, with those of the
Old World. Consequently we have here everything favourable for much
modification,--for far more modification than with the Alpine
productions, left isolated, within a much more recent period, on the
several mountain-ranges and on the arctic lands of the two Worlds.
Hence it has come, that when we compare the now living productions of
the temperate regions of the New and Old Worlds, we find very few
identical species (though Asa Gray has lately shown that more plants
are identical than was formerly supposed), but we find in every great
class many forms, which some naturalists rank as geographical races,
and others as distinct species; and a host of closely allied or
representative forms which are ranked by all naturalists as
specifically distinct.
As on the land, so in the waters of the sea, a slow southern migration
of a marine fauna, which during the Pliocene or even a somewhat
earlier period, was nearly uniform along the continuous shores of the
Polar Circle, will account, on the theory of modification, for many
closely allied forms now living in areas completely sundered. Thus, I
think, we can understand the presence of many existing and tertiary
representative forms on the eastern and western shores of temperate
North America; and the still more striking case of many closely allied
crustaceans (as described in Dana's admirable work), of some fish and
other marine animals, in the Mediterranean and in the seas of
Japan,--areas now separated by a continent and by nearly a hemisphere
of equatorial ocean.
These cases of relationship, without identity, of the inhabitants of
seas now disjoined, and likewise of the past and present inhabitants
of the temperate lands of North America and Europe, are inexplicable
on the theory of creation. We cannot say that they have been created
alike, in correspondence with the nearly similar physical conditions
of the areas; for if we compare, for instance, certain parts of South
America with the southern continents of the Old World, we see
countries closely corresponding in all their physical conditions, but
with their inhabitants utterly dissimilar.
But we must return to our more immediate subject, the Glacial period.
I am convinced that Forbes's view may be largely extended. In Europe
we have the plainest evidence of the cold period, from the western
shores of Britain to the Oural range, and southward to the Pyrenees.
We may infer, from the frozen mammals and nature of the mountain
vegetation, that Siberia was similarly affected. Along the Himalaya,
at points 900 miles apart, glaciers have left the marks of their
former low descent; and in Sikkim, Dr. Hooker saw maize growing on
gigantic ancient moraines. South of the equator, we have some direct
evidence of former glacial action in New Zealand; and the same plants,
found on widely separated mountains in this island, tell the same
story. If one account which has been published can be trusted, we have
direct evidence of glacial action in the south-eastern corner of
Australia.
Looking to America; in the northern half, ice-borne fragments of rock
have been observed on the eastern side as far south as lat. 36 deg-37
deg, and on the shores of the Pacific, where the climate is now so
different, as far south as lat. 46 deg; erratic boulders have, also,
been noticed on the Rocky Mountains. In the Cordillera of Equatorial
South America, glaciers once extended far below their present level.
In central Chile I was astonished at the structure of a vast mound of
detritus, about 800 feet in height, crossing a valley of the Andes;
and this I now feel convinced was a gigantic moraine, left far below
any existing glacier. Further south on both sides of the continent,
from lat. 41 deg to the southernmost extremity, we have the clearest
evidence of former glacial action, in huge boulders transported far
from their parent source.
We do not know that the Glacial epoch was strictly simultaneous at
these several far distant points on opposite sides of the world. But
we have good evidence in almost every case, that the epoch was
included within the latest geological period. We have, also, excellent
evidence, that it endured for an enormous time, as measured by years,
at each point. The cold may have come on, or have ceased, earlier at
one point of the globe than at another, but seeing that it endured for
long at each, and that it was contemporaneous in a geological sense,
it seems to me probable that it was, during a part at least of the
period, actually simultaneous throughout the world. Without some
distinct evidence to the contrary, we may at least admit as probable
that the glacial action was simultaneous on the eastern and western
sides of North America, in the Cordillera under the equator and under
the warmer temperate zones, and on both sides of the southern
extremity of the continent. If this be admitted, it is difficult to
avoid believing that the temperature of the whole world was at this
period simultaneously cooler. But it would suffice for my purpose, if
the temperature was at the same time lower along certain broad belts
of longitude.
On this view of the whole world, or at least of broad longitudinal
belts, having been simultaneously colder from pole to pole, much light
can be thrown on the present distribution of identical and allied
species. In America, Dr. Hooker has shown that between forty and fifty
of the flowering plants of Tierra del Fuego, forming no inconsiderable
part of its scanty flora, are common to Europe, enormously remote as
these two points are; and there are many closely allied species. On
the lofty mountains of equatorial America a host of peculiar species
belonging to European genera occur. On the highest mountains of
Brazil, some few European genera were found by Gardner, which do not
exist in the wide intervening hot countries. So on the Silla of
Caraccas the illustrious Humboldt long ago found species belonging to
genera characteristic of the Cordillera. On the mountains of
Abyssinia, several European forms and some few representatives of the
peculiar flora of the Cape of Good Hope occur. At the Cape of Good
Hope a very few European species, believed not to have been introduced
by man, and on the mountains, some few representative European forms
are found, which have not been discovered in the intertropical parts
of Africa. On the Himalaya, and on the isolated mountain-ranges of the
peninsula of India, on the heights of Ceylon, and on the volcanic
cones of Java, many plants occur, either identically the same or
representing each other, and at the same time representing plants of
Europe, not found in the intervening hot lowlands. A list of the
genera collected on the loftier peaks of Java raises a picture of a
collection made on a hill in Europe! Still more striking is the fact
that southern Australian forms are clearly represented by plants
growing on the summits of the mountains of Borneo. Some of these
Australian forms, as I hear from Dr. Hooker, extend along the heights
of the peninsula of Malacca, and are thinly scattered, on the one hand
over India and on the other as far north as Japan.
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