John Lothrop Motley - "History of the United Netherlands, 1600 02"

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John Lothrop Motley - "History of the United Netherlands, 1588"

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Books: On the Origin of Species

C >> Charles Darwin >> On the Origin of Species




It is most difficult always to remember that the increase of every
living being is constantly being checked by unperceived injurious
agencies; and that these same unperceived agencies are amply
sufficient to cause rarity, and finally extinction. We see in many
cases in the more recent tertiary formations, that rarity precedes
extinction; and we know that this has been the progress of events with
those animals which have been exterminated, either locally or wholly,
through man's agency. I may repeat what I published in 1845, namely,
that to admit that species generally become rare before they become
extinct--to feel no surprise at the rarity of a species, and yet to
marvel greatly when it ceases to exist, is much the same as to admit
that sickness in the individual is the forerunner of death--to feel no
surprise at sickness, but when the sick man dies, to wonder and to
suspect that he died by some unknown deed of violence.

The theory of natural selection is grounded on the belief that each
new variety, and ultimately each new species, is produced and
maintained by having some advantage over those with which it comes
into competition; and the consequent extinction of less-favoured forms
almost inevitably follows. It is the same with our domestic
productions: when a new and slightly improved variety has been raised,
it at first supplants the less improved varieties in the same
neighbourhood; when much improved it is transported far and near, like
our short-horn cattle, and takes the place of other breeds in other
countries. Thus the appearance of new forms and the disappearance of
old forms, both natural and artificial, are bound together. In certain
flourishing groups, the number of new specific forms which have been
produced within a given time is probably greater than that of the old
forms which have been exterminated; but we know that the number of
species has not gone on indefinitely increasing, at least during the
later geological periods, so that looking to later times we may
believe that the production of new forms has caused the extinction of
about the same number of old forms.

The competition will generally be most severe, as formerly explained
and illustrated by examples, between the forms which are most like
each other in all respects. Hence the improved and modified
descendants of a species will generally cause the extermination of the
parent-species; and if many new forms have been developed from any one
species, the nearest allies of that species, i.e. the species of the
same genus, will be the most liable to extermination. Thus, as I
believe, a number of new species descended from one species, that is a
new genus, comes to supplant an old genus, belonging to the same
family. But it must often have happened that a new species belonging
to some one group will have seized on the place occupied by a species
belonging to a distinct group, and thus caused its extermination; and
if many allied forms be developed from the successful intruder, many
will have to yield their places; and it will generally be allied
forms, which will suffer from some inherited inferiority in common.
But whether it be species belonging to the same or to a distinct
class, which yield their places to other species which have been
modified and improved, a few of the sufferers may often long be
preserved, from being fitted to some peculiar line of life, or from
inhabiting some distant and isolated station, where they have escaped
severe competition. For instance, a single species of Trigonia, a
great genus of shells in the secondary formations, survives in the
Australian seas; and a few members of the great and almost extinct
group of Ganoid fishes still inhabit our fresh waters. Therefore the
utter extinction of a group is generally, as we have seen, a slower
process than its production.

With respect to the apparently sudden extermination of whole families
or orders, as of Trilobites at the close of the palaeozoic period and
of Ammonites at the close of the secondary period, we must remember
what has been already said on the probable wide intervals of time
between our consecutive formations; and in these intervals there may
have been much slow extermination. Moreover, when by sudden
immigration or by unusually rapid development, many species of a new
group have taken possession of a new area, they will have exterminated
in a correspondingly rapid manner many of the old inhabitants; and the
forms which thus yield their places will commonly be allied, for they
will partake of some inferiority in common.

Thus, as it seems to me, the manner in which single species and whole
groups of species become extinct, accords well with the theory of
natural selection. We need not marvel at extinction; if we must
marvel, let it be at our presumption in imagining for a moment that we
understand the many complex contingencies, on which the existence of
each species depends. If we forget for an instant, that each species
tends to increase inordinately, and that some check is always in
action, yet seldom perceived by us, the whole economy of nature will
be utterly obscured. Whenever we can precisely say why this species is
more abundant in individuals than that; why this species and not
another can be naturalised in a given country; then, and not till
then, we may justly feel surprise why we cannot account for the
extinction of this particular species or group of species.

ON THE FORMS OF LIFE CHANGING ALMOST SIMULTANEOUSLY THROUGHOUT THE
WORLD.

Scarcely any palaeontological discovery is more striking than the
fact, that the forms of life change almost simultaneously throughout
the world. Thus our European Chalk formation can be recognised in many
distant parts of the world, under the most different climates, where
not a fragment of the mineral chalk itself can be found; namely, in
North America, in equatorial South America, in Tierra del Fuego, at
the Cape of Good Hope, and in the peninsula of India. For at these
distant points, the organic remains in certain beds present an
unmistakeable degree of resemblance to those of the Chalk. It is not
that the same species are met with; for in some cases not one species
is identically the same, but they belong to the same families, genera,
and sections of genera, and sometimes are similarly characterised in
such trifling points as mere superficial sculpture. Moreover other
forms, which are not found in the Chalk of Europe, but which occur in
the formations either above or below, are similarly absent at these
distant points of the world. In the several successive palaeozoic
formations of Russia, Western Europe and North America, a similar
parallelism in the forms of life has been observed by several authors:
so it is, according to Lyell, with the several European and North
American tertiary deposits. Even if the few fossil species which are
common to the Old and New Worlds be kept wholly out of view, the
general parallelism in the successive forms of life, in the stages of
the widely separated palaeozoic and tertiary periods, would still be
manifest, and the several formations could be easily correlated.

These observations, however, relate to the marine inhabitants of
distant parts of the world: we have not sufficient data to judge
whether the productions of the land and of fresh water change at
distant points in the same parallel manner. We may doubt whether they
have thus changed: if the Megatherium, Mylodon, Macrauchenia, and
Toxodon had been brought to Europe from La Plata, without any
information in regard to their geological position, no one would have
suspected that they had coexisted with still living sea-shells; but as
these anomalous monsters coexisted with the Mastodon and Horse, it
might at least have been inferred that they had lived during one of
the latter tertiary stages.

When the marine forms of life are spoken of as having changed
simultaneously throughout the world, it must not be supposed that this
expression relates to the same thousandth or hundred-thousandth year,
or even that it has a very strict geological sense; for if all the
marine animals which live at the present day in Europe, and all those
that lived in Europe during the pleistocene period (an enormously
remote period as measured by years, including the whole glacial
epoch), were to be compared with those now living in South America or
in Australia, the most skilful naturalist would hardly be able to say
whether the existing or the pleistocene inhabitants of Europe
resembled most closely those of the southern hemisphere. So, again,
several highly competent observers believe that the existing
productions of the United States are more closely related to those
which lived in Europe during certain later tertiary stages, than to
those which now live here; and if this be so, it is evident that
fossiliferous beds deposited at the present day on the shores of North
America would hereafter be liable to be classed with somewhat older
European beds. Nevertheless, looking to a remotely future epoch, there
can, I think, be little doubt that all the more modern MARINE
formations, namely, the upper pliocene, the pleistocene and strictly
modern beds, of Europe, North and South America, and Australia, from
containing fossil remains in some degree allied, and from not
including those forms which are only found in the older underlying
deposits, would be correctly ranked as simultaneous in a geological
sense.

The fact of the forms of life changing simultaneously, in the above
large sense, at distant parts of the world, has greatly struck those
admirable observers, MM. de Verneuil and d'Archiac. After referring to
the parallelism of the palaeozoic forms of life in various parts of
Europe, they add, "If struck by this strange sequence, we turn our
attention to North America, and there discover a series of analogous
phenomena, it will appear certain that all these modifications of
species, their extinction, and the introduction of new ones, cannot be
owing to mere changes in marine currents or other causes more or less
local and temporary, but depend on general laws which govern the whole
animal kingdom." M. Barrande has made forcible remarks to precisely
the same effect. It is, indeed, quite futile to look to changes of
currents, climate, or other physical conditions, as the cause of these
great mutations in the forms of life throughout the world, under the
most different climates. We must, as Barrande has remarked, look to
some special law. We shall see this more clearly when we treat of the
present distribution of organic beings, and find how slight is the
relation between the physical conditions of various countries, and the
nature of their inhabitants.

This great fact of the parallel succession of the forms of life
throughout the world, is explicable on the theory of natural
selection. New species are formed by new varieties arising, which have
some advantage over older forms; and those forms, which are already
dominant, or have some advantage over the other forms in their own
country, would naturally oftenest give rise to new varieties or
incipient species; for these latter must be victorious in a still
higher degree in order to be preserved and to survive. We have
distinct evidence on this head, in the plants which are dominant, that
is, which are commonest in their own homes, and are most widely
diffused, having produced the greatest number of new varieties. It is
also natural that the dominant, varying, and far-spreading species,
which already have invaded to a certain extent the territories of
other species, should be those which would have the best chance of
spreading still further, and of giving rise in new countries to new
varieties and species. The process of diffusion may often be very
slow, being dependent on climatal and geographical changes, or on
strange accidents, but in the long run the dominant forms will
generally succeed in spreading. The diffusion would, it is probable,
be slower with the terrestrial inhabitants of distinct continents than
with the marine inhabitants of the continuous sea. We might therefore
expect to find, as we apparently do find, a less strict degree of
parallel succession in the productions of the land than of the sea.

Dominant species spreading from any region might encounter still more
dominant species, and then their triumphant course, or even their
existence, would cease. We know not at all precisely what are all the
conditions most favourable for the multiplication of new and dominant
species; but we can, I think, clearly see that a number of
individuals, from giving a better chance of the appearance of
favourable variations, and that severe competition with many already
existing forms, would be highly favourable, as would be the power of
spreading into new territories. A certain amount of isolation,
recurring at long intervals of time, would probably be also
favourable, as before explained. One quarter of the world may have
been most favourable for the production of new and dominant species on
the land, and another for those in the waters of the sea. If two great
regions had been for a long period favourably circumstanced in an
equal degree, whenever their inhabitants met, the battle would be
prolonged and severe; and some from one birthplace and some from the
other might be victorious. But in the course of time, the forms
dominant in the highest degree, wherever produced, would tend
everywhere to prevail. As they prevailed, they would cause the
extinction of other and inferior forms; and as these inferior forms
would be allied in groups by inheritance, whole groups would tend
slowly to disappear; though here and there a single member might long
be enabled to survive.

Thus, as it seems to me, the parallel, and, taken in a large sense,
simultaneous, succession of the same forms of life throughout the
world, accords well with the principle of new species having been
formed by dominant species spreading widely and varying; the new
species thus produced being themselves dominant owing to inheritance,
and to having already had some advantage over their parents or over
other species; these again spreading, varying, and producing new
species. The forms which are beaten and which yield their places to
the new and victorious forms, will generally be allied in groups, from
inheriting some inferiority in common; and therefore as new and
improved groups spread throughout the world, old groups will disappear
from the world; and the succession of forms in both ways will
everywhere tend to correspond.

There is one other remark connected with this subject worth making. I
have given my reasons for believing that all our greater fossiliferous
formations were deposited during periods of subsidence; and that blank
intervals of vast duration occurred during the periods when the bed of
the sea was either stationary or rising, and likewise when sediment
was not thrown down quickly enough to embed and preserve organic
remains. During these long and blank intervals I suppose that the
inhabitants of each region underwent a considerable amount of
modification and extinction, and that there was much migration from
other parts of the world. As we have reason to believe that large
areas are affected by the same movement, it is probable that strictly
contemporaneous formations have often been accumulated over very wide
spaces in the same quarter of the world; but we are far from having
any right to conclude that this has invariably been the case, and that
large areas have invariably been affected by the same movements. When
two formations have been deposited in two regions during nearly, but
not exactly the same period, we should find in both, from the causes
explained in the foregoing paragraphs, the same general succession in
the forms of life; but the species would not exactly correspond; for
there will have been a little more time in the one region than in the
other for modification, extinction, and immigration.

I suspect that cases of this nature have occurred in Europe. Mr.
Prestwich, in his admirable Memoirs on the eocene deposits of England
and France, is able to draw a close general parallelism between the
successive stages in the two countries; but when he compares certain
stages in England with those in France, although he finds in both a
curious accordance in the numbers of the species belonging to the same
genera, yet the species themselves differ in a manner very difficult
to account for, considering the proximity of the two areas,--unless,
indeed, it be assumed that an isthmus separated two seas inhabited by
distinct, but contemporaneous, faunas. Lyell has made similar
observations on some of the later tertiary formations. Barrande, also,
shows that there is a striking general parallelism in the successive
Silurian deposits of Bohemia and Scandinavia; nevertheless he finds a
surprising amount of difference in the species. If the several
formations in these regions have not been deposited during the same
exact periods,--a formation in one region often corresponding with a
blank interval in the other,--and if in both regions the species have
gone on slowly changing during the accumulation of the several
formations and during the long intervals of time between them; in this
case, the several formations in the two regions could be arranged in
the same order, in accordance with the general succession of the form
of life, and the order would falsely appear to be strictly parallel;
nevertheless the species would not all be the same in the apparently
corresponding stages in the two regions.
$
ON THE AFFINITIES OF EXTINCT SPECIES TO EACH OTHER, AND TO LIVING
FORMS.

Let us now look to the mutual affinities of extinct and living
species. They all fall into one grand natural system; and this fact is
at once explained on the principle of descent. The more ancient any
form is, the more, as a general rule, it differs from living forms.
But, as Buckland long ago remarked, all fossils can be classed either
in still existing groups, or between them. That the extinct forms of
life help to fill up the wide intervals between existing genera,
families, and orders, cannot be disputed. For if we confine our
attention either to the living or to the extinct alone, the series is
far less perfect than if we combine both into one general system. With
respect to the Vertebrata, whole pages could be filled with striking
illustrations from our great palaeontologist, Owen, showing how
extinct animals fall in between existing groups. Cuvier ranked the
Ruminants and Pachyderms, as the two most distinct orders of mammals;
but Owen has discovered so many fossil links, that he has had to alter
the whole classification of these two orders; and has placed certain
pachyderms in the same sub-order with ruminants: for example, he
dissolves by fine gradations the apparently wide difference between
the pig and the camel. In regard to the Invertebrata, Barrande, and a
higher authority could not be named, asserts that he is every day
taught that palaeozoic animals, though belonging to the same orders,
families, or genera with those living at the present day, were not at
this early epoch limited in such distinct groups as they now are.

Some writers have objected to any extinct species or group of species
being considered as intermediate between living species or groups. If
by this term it is meant that an extinct form is directly intermediate
in all its characters between two living forms, the objection is
probably valid. But I apprehend that in a perfectly natural
classification many fossil species would have to stand between living
species, and some extinct genera between living genera, even between
genera belonging to distinct families. The most common case,
especially with respect to very distinct groups, such as fish and
reptiles, seems to be, that supposing them to be distinguished at the
present day from each other by a dozen characters, the ancient members
of the same two groups would be distinguished by a somewhat lesser
number of characters, so that the two groups, though formerly quite
distinct, at that period made some small approach to each other.

It is a common belief that the more ancient a form is, by so much the
more it tends to connect by some of its characters groups now widely
separated from each other. This remark no doubt must be restricted to
those groups which have undergone much change in the course of
geological ages; and it would be difficult to prove the truth of the
proposition, for every now and then even a living animal, as the
Lepidosiren, is discovered having affinities directed towards very
distinct groups. Yet if we compare the older Reptiles and Batrachians,
the older Fish, the older Cephalopods, and the eocene Mammals, with
the more recent members of the same classes, we must admit that there
is some truth in the remark.

Let us see how far these several facts and inferences accord with the
theory of descent with modification. As the subject is somewhat
complex, I must request the reader to turn to the diagram in the
fourth chapter. We may suppose that the numbered letters represent
genera, and the dotted lines diverging from them the species in each
genus. The diagram is much too simple, too few genera and too few
species being given, but this is unimportant for us. The horizontal
lines may represent successive geological formations, and all the
forms beneath the uppermost line may be considered as extinct. The
three existing genera, a14, q14, p14, will form a small family; b14
and f14 a closely allied family or sub-family; and o14, e14, m14, a
third family. These three families, together with the many extinct
genera on the several lines of descent diverging from the parent-form
A, will form an order; for all will have inherited something in common
from their ancient and common progenitor. On the principle of the
continued tendency to divergence of character, which was formerly
illustrated by this diagram, the more recent any form is, the more it
will generally differ from its ancient progenitor. Hence we can
understand the rule that the most ancient fossils differ most from
existing forms. We must not, however, assume that divergence of
character is a necessary contingency; it depends solely on the
descendants from a species being thus enabled to seize on many and
different places in the economy of nature. Therefore it is quite
possible, as we have seen in the case of some Silurian forms, that a
species might go on being slightly modified in relation to its
slightly altered conditions of life, and yet retain throughout a vast
period the same general characteristics. This is represented in the
diagram by the letter F14.

All the many forms, extinct and recent, descended from A, make, as
before remarked, one order; and this order, from the continued effects
of extinction and divergence of character, has become divided into
several sub-families and families, some of which are supposed to have
perished at different periods, and some to have endured to the present
day.

By looking at the diagram we can see that if many of the extinct
forms, supposed to be embedded in the successive formations, were
discovered at several points low down in the series, the three
existing families on the uppermost line would be rendered less
distinct from each other. If, for instance, the genera a1, a5, a10,
f8, m3, m6, m9 were disinterred, these three families would be so
closely linked together that they probably would have to be united
into one great family, in nearly the same manner as has occurred with
ruminants and pachyderms. Yet he who objected to call the extinct
genera, which thus linked the living genera of three families
together, intermediate in character, would be justified, as they are
intermediate, not directly, but only by a long and circuitous course
through many widely different forms. If many extinct forms were to be
discovered above one of the middle horizontal lines or geological
formations--for instance, above Number VI.--but none from beneath this
line, then only the two families on the left hand (namely, a14, etc.,
and b14, etc.) would have to be united into one family; and the two
other families (namely, a14 to f14 now including five genera, and o14
to m14) would yet remain distinct. These two families, however, would
be less distinct from each other than they were before the discovery
of the fossils. If, for instance, we suppose the existing genera of
the two families to differ from each other by a dozen characters, in
this case the genera, at the early period marked VI., would differ by
a lesser number of characters; for at this early stage of descent they
have not diverged in character from the common progenitor of the
order, nearly so much as they subsequently diverged. Thus it comes
that ancient and extinct genera are often in some slight degree
intermediate in character between their modified descendants, or
between their collateral relations.