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Darwinism (1889) by Alfred Russel Wallace

A >> Alfred Russel Wallace >> Darwinism (1889)






There is perhaps no phenomenon of nature that is at once so important,
so universal; and so little understood, as the struggle for existence
continually going on among all organised beings. To most persons nature
appears calm, orderly, and peaceful. They see the birds singing in the
trees, the insects hovering over the flowers, the squirrel climbing
among the tree-tops, and all living things in the possession of health
and vigour, and in the enjoyment of a sunny existence. But they do not
see, and hardly ever think of, the means by which this beauty and
harmony and enjoyment is brought about. They do not see the constant and
daily search after food, the failure to obtain which means weakness or
death; the constant effort to escape enemies; the ever-recurring
struggle against the forces of nature. This daily and hourly struggle,
this incessant warfare, is nevertheless the very means by which much of
the beauty and harmony and enjoyment in nature is produced, and also
affords one of the most important elements in bringing about the origin
of species. We must, therefore, devote some time to the consideration of
its various aspects and of the many curious phenomena to which it gives
rise.

It is a matter of common observation that if weeds are allowed to grow
unchecked in a garden they will soon destroy a number of the flowers.
It is not so commonly known that if a garden is left to become
altogether wild, the weeds that first take possession of it, often
covering the whole surface of the ground with two or three different
kinds, will themselves be supplanted by others, so that in a few years
many of the original flowers and of the earliest weeds may alike have
disappeared. This is one of the very simplest cases of the struggle for
existence, resulting in the successive displacement of one set of
species by another; but the exact causes of this displacement are by no
means of such a simple nature. All the plants concerned may be perfectly
hardy, all may grow freely from seed, yet when left alone for a number
of years, each set is in turn driven out by a succeeding set, till at
the end of a considerable period--a century or a few centuries
perhaps--hardly one of the plants which first monopolised the ground
would be found there.

Another phenomenon of an analogous kind is presented by the different
behaviour of introduced wild plants or animals into countries apparently
quite as well suited to them as those which they naturally inhabit.
Agassiz, in his work on Lake Superior, states that the roadside weeds of
the northeastern United States, to the number of 130 species, are all
European, the native weeds having disappeared westwards; and in New
Zealand there are no less than 250 species of naturalised European
plants, more than 100 species of which have spread widely over the
country, often displacing the native vegetation. On the other hand, of
the many hundreds of hardy plants which produce seed freely in our
gardens, very few ever run wild, and hardly any have become common. Even
attempts to naturalise suitable plants usually fail; for A. de Candolle
states that several botanists of Paris, Geneva, and especially of
Montpellier, have sown the seeds of many hundreds of species of hardy
exotic plants in what appeared to be the most favourable situations, but
that, in hardly a single case, has any one of them become
naturalised.[4] Even a plant like the potato--so widely cultivated, so
hardy, and so well adapted to spread by means of its many-eyed
tubers--has not established itself in a wild state in any part of
Europe. It would be thought that Australian plants would easily run
wild in New Zealand. But Sir Joseph Hooker informs us that the late Mr.
Bidwell habitually scattered Australian seeds during his extensive
travels in New Zealand, yet only two or three Australian plants appear
to have established themselves in that country, and these only in
cultivated or newly moved soil.

These few illustrations sufficiently show that all the plants of a
country are, as De Candolle says, at war with each other, each one
struggling to occupy ground at the expense of its neighbour. But,
besides this direct competition, there is one not less powerful arising
from the exposure of almost all plants to destruction by animals. The
buds are destroyed by birds, the leaves by caterpillars, the seeds by
weevils; some insects bore into the trunk, others burrow in the twigs
and leaves; slugs devour the young seedlings and the tender shoots,
wire-worms gnaw the roots. Herbivorous mammals devour many species
bodily, while some uproot and devour the buried tubers.

In animals, it is the eggs or the very young that suffer most from their
various enemies; in plants, the tender seedlings when they first appear
above the ground. To illustrate this latter point Mr. Darwin cleared and
dug a piece of ground three feet long and two feet wide, and then marked
all the seedlings of weeds and other plants which came up, noting what
became of them. The total number was 357, and out of these no less than
295 were destroyed by slugs and insects. The direct strife of plant with
plant is almost equally fatal when the stronger are allowed to smother
the weaker. When turf is mown or closely browsed by animals, a number of
strong and weak plants live together, because none are allowed to grow
much beyond the rest; but Mr. Darwin found that when the plants which
compose such turf are allowed to grow up freely, the stronger kill the
weaker. In a plot of turf three feet by four, twenty distinct species of
plants were found to be growing, and no less than nine of these perished
altogether when the other species were allowed to grow up to their full
size.[5]

But besides having to protect themselves against competing plants and
against destructive animals, there is a yet deadlier enemy in the
forces of inorganic nature. Each species can sustain a certain amount of
heat and cold, each requires a certain amount of moisture at the right
season, each wants a proper amount of light or of direct sunshine, each
needs certain elements in the soil; the failure of a due proportion in
these inorganic conditions causes weakness, and thus leads to speedy
death. The struggle for existence in plants is, therefore, threefold in
character and infinite in complexity, and the result is seen in their
curiously irregular distribution over the face of the earth. Not only
has each country its distinct plants, but every valley, every hillside,
almost every hedgerow, has a different set of plants from its adjacent
valley, hillside, or hedgerow--if not always different in the actual
species yet very different in comparative abundance, some which are rare
in the one being common in the other. Hence it happens that slight
changes of conditions often produce great changes in the flora of a
country. Thus in 1740 and the two following years the larva of a moth
(Phalaena graminis) committed such destruction in many of the meadows of
Sweden that the grass was greatly diminished in quantity, and many
plants which were before choked by the grass sprang up, and the ground
became variegated with a multitude of different species of flowers. The
introduction of goats into the island of St. Helena led to the entire
destruction of the native forests, consisting of about a hundred
distinct species of trees and shrubs, the young plants being devoured by
the goats as fast as they grew up. The camel is a still greater enemy to
woody vegetation than the goat, and Mr. Marsh believes that forests
would soon cover considerable tracts of the Arabian and African deserts
if the goat and the camel were removed from them.[6] Even in many parts
of our own country the existence of trees is dependent on the absence of
cattle. Mr. Darwin observed, on some extensive heaths near Farnham, in
Surrey, a few clumps of old Scotch firs, but no young trees over
hundreds of acres. Some portions of the heath had, however, been
enclosed a few years before, and these enclosures were crowded with
young fir-trees growing too close together for all to live; and these
were not sown or planted, nothing having been done to the ground beyond
enclosing it so as to keep out cattle. On ascertaining this, Mr. Darwin
was so much surprised that he searched among the heather in the
unenclosed parts, and there he found multitudes of little trees and
seedlings which had been perpetually browsed down by the cattle. In one
square yard, at a point about a hundred yards from one of the old clumps
of firs, he counted thirty-two little trees, and one of them had
twenty-six rings of growth, showing that it had for many years tried to
raise its head above the stems of the heather and had failed. Yet this
heath was very extensive and very barren, and, as Mr. Darwin remarks, no
one would ever have imagined that cattle would have so closely and so
effectually searched it for food.

In the case of animals, the competition and struggle are more obvious.
The vegetation of a given district can only support a certain number of
animals, and the different kinds of plant-eaters will compete together
for it. They will also have insects for their competitors, and these
insects will be kept down by birds, which will thus assist the mammalia.
But there will also be carnivora destroying the herbivora; while small
rodents, like the lemming and some of the field-mice, often destroy so
much vegetation as materially to affect the food of all the other groups
of animals. Droughts, floods, severe winters, storms and hurricanes will
injure these in various degrees, but no one species can be diminished in
numbers without the effect being felt in various complex ways by all the
rest. A few illustrations of this reciprocal action must be given.


_Illustrative Cases of the Struggle for Life_.

Sir Charles Lyell observes that if, by the attacks of seals or other
marine foes, salmon are reduced in numbers, the consequence will be that
otters, living far inland, will be deprived of food and will then
destroy many young birds or quadrupeds, so that the increase of a marine
animal may cause the destruction of many land animals hundreds of miles
away. Mr. Darwin carefully observed the effects produced by planting a
few hundred acres of Scotch fir, in Staffordshire, on part of a very
extensive heath which had never been cultivated. After the planted
portion was about twenty-five years old he observed that the change in
the native vegetation was greater than is often seen in passing from
one quite different soil to another. Besides a great change in the
proportional numbers of the native heath-plants, twelve species which
could not be found on the heath flourished in the plantations. The
effect on the insect life must have been still greater, for six
insectivorous birds which were very common in the plantations were not
to be seen on the heath, which was, however, frequented by two or three
different species of insectivorous birds. It would have required
continued study for several years to determine all the differences in
the organic life of the two areas, but the facts stated by Mr. Darwin
are sufficient to show how great a change may be effected by the
introduction of a single kind of tree and the keeping out of cattle.

The next case I will give in Mr. Darwin's own words: "In several parts
of the world insects determine the existence of cattle. Perhaps Paraguay
offers the most curious instance of this; for here neither cattle nor
horses nor dogs have ever run wild, though they swarm southward and
northward in a feral state; and Azara and Rengger have shown that this
is caused by the greater numbers, in Paraguay, of a certain fly which
lays its eggs in the navels of these animals when first born. The
increase of these flies, numerous as they are, must be habitually
checked by some means, probably by other parasitic insects. Hence, if
certain insectivorous birds were to decrease in Paraguay, the parasitic
insects would probably increase; and this would lessen the number of the
navel-frequenting flies--then cattle and horses would become feral, and
this would greatly alter (as indeed I have observed in parts of South
America) the vegetation: this again would largely affect the insects,
and this, as we have just seen in Staffordshire, the insectivorous
birds, and so onward in ever-increasing circles of complexity. Not that
under nature the relations will ever be as simple as this. Battle within
battle must be continually recurring with varying success; and yet in
the long run the forces are so nicely balanced, that the face of nature
remains for a long time uniform, though assuredly the merest trifle
would give the victory to one organic being over another."[7]

Such cases as the above may perhaps be thought exceptional, but there
is good reason to believe that they are by no means rare, but are
illustrations of what is going on in every part of the world, only it is
very difficult for us to trace out the complex reactions that are
everywhere occurring. The general impression of the ordinary observer
seems to be that wild animals and plants live peaceful lives and have
few troubles, each being exactly suited to its place and surroundings,
and therefore having no difficulty in maintaining itself. Before showing
that this view is, everywhere and always, demonstrably untrue, we will
consider one other case of the complex relations of distinct organisms
adduced by Mr. Darwin, and often quoted for its striking and almost
eccentric character. It is now well known that many flowers require to
be fertilised by insects in order to produce seed, and this
fertilisation can, in some cases, only be effected by one particular
species of insect to which the flower has become specially adapted. Two
of our common plants, the wild heart's-ease (Viola tricolor) and the red
clover (Trifolium pratense), are thus fertilised by humble-bees almost
exclusively, and if these insects are prevented from visiting the
flowers, they produce either no seed at all or exceedingly few. Now it
is known that field-mice destroy the combs and nests of humble-bees, and
Colonel Newman, who has paid great attention to these insects, believes
that more than two-thirds of all the humble-bees' nests in England are
thus destroyed. But the number of mice depends a good deal on the number
of cats; and the same observer says that near villages and towns he has
found the nests of humble-bees more numerous than elsewhere, which he
attributes to the number of cats that destroy the mice. Hence it
follows, that the abundance of red clover and wild heart's-ease in a
district will depend on a good supply of cats to kill the mice, which
would otherwise destroy and keep down the humble-bees and prevent them
from fertilising the flowers. A chain of connection has thus been found
between such totally distinct organisms as flesh-eating mammalia and
sweet-smelling flowers, the abundance or scarcity of the one closely
corresponding to that of the other!

The following account of the struggle between trees in the forests of
Denmark, from the researches of M. Hansten-Blangsted, strikingly
illustrates our subject.[8] The chief combatants are the beech and the
birch, the former being everywhere successful in its invasions. Forests
composed wholly of birch are now only found in sterile, sandy tracts;
everywhere else the trees are mixed, and wherever the soil is favourable
the beech rapidly drives out the birch. The latter loses its branches at
the touch of the beech, and devotes all its strength to the upper part
where it towers above the beech. It may live long in this way, but it
succumbs ultimately in the fight--of old age if of nothing else, for the
life of the birch in Denmark is shorter than that of the beech. The
writer believes that light (or rather shade) is the cause of the
superiority of the latter, for it has a greater development of its
branches than the birch, which is more open and thus allows the rays of
the sun to pass through to the soil below, while the tufted, bushy top
of the beech preserves a deep shade at its base. Hardly any young plants
can grow under the beech except its own shoots; and while the beech can
nourish under the shade of the birch, the latter dies immediately under
the beech. The birch has only been saved from total extermination by the
facts that it had possession of the Danish forests long before the beech
ever reached the country, and that certain districts are unfavourable to
the growth of the latter. But wherever the soil has been enriched by the
decomposition of the leaves of the birch the battle begins. The birch
still flourishes on the borders of lakes and other marshy places, where
its enemy cannot exist. In the same way, in the forests of Zeeland, the
fir forests are disappearing before the beech. Left to themselves, the
firs are soon displaced by the beech. The struggle between the latter
and the oak is longer and more stubborn, for the branches and foliage of
the oak are thicker, and offer much resistance to the passage of light.
The oak, also, has greater longevity; but, sooner or later, it too
succumbs, because it cannot develop in the shadow of the beech. The
earliest forests of Denmark were mainly composed of aspens, with which
the birch was apparently associated; gradually the soil was raised, and
the climate grew milder; then the fir came and formed large forests.
This tree ruled for centuries, and then ceded the first place to the
holm-oak, which is now giving way to the beech. Aspen, birch, fir, oak,
and beech appear to be the steps in the struggle for the survival of the
fittest among the forest-trees of Denmark.

It may be added that in the time of the Romans the beech was the
principal forest-tree of Denmark as it is now, while in the much earlier
bronze age, represented by the later remains found in the peat bogs,
there were no beech-trees, or very few, the oak being the prevailing
tree, while in the still earlier stone period the fir was the most
abundant. The beech is a tree essentially of the temperate zone, having
its northern limit considerably southward of the oak, fir, birch, or
aspen, and its entrance into Denmark was no doubt due to the
amelioration of the climate after the glacial epoch had entirely passed
away. We thus see how changes of climate, which are continually
occurring owing either to cosmical or geographical causes, may initiate
a struggle among plants which may continue for thousands of years, and
which must profoundly modify the relations of the animal world, since
the very existence of innumerable insects, and even of many birds and
mammals, is dependent more or less completely on certain species of
plants.


_The Struggle for Existence on the Pampas_.

Another illustration of the struggle for existence, in which both plants
and animals are implicated, is afforded by the pampas of the southern
part of South America. The absence of trees from these vast plains has
been imputed by Mr. Darwin to the supposed inability of the tropical and
sub-tropical forms of South America to thrive on them, and there being
no other source from which they could obtain a supply; and that
explanation was adopted by such eminent botanists as Mr. Ball and
Professor Asa Gray. This explanation has always seemed to me
unsatisfactory, because there are ample forests both in the temperate
regions of the Andes and on the whole west coast down to Terra del
Fuego; and it is inconsistent with what we know of the rapid variation
and adaptation of species to new conditions. What seems a more
satisfactory explanation has been given by Mr. Edwin Clark, a civil
engineer, who resided nearly two years in the country and paid much
attention to its natural history. He says: "The peculiar characteristics
of these vast level plains which descend from the Andes to the great
river basin in unbroken monotony, are the absence of rivers or
water-storage, and the periodical occurrence of droughts, or 'siccos,'
in the summer months. These conditions determine the singular character
both of its flora and fauna.

"The soil is naturally fertile and favourable for the growth of trees,
and they grow luxuriantly wherever they are protected. The eucalyptus is
covering large tracts wherever it is enclosed, and willows, poplars, and
the fig surround every estancia when fenced in.

"The open plains are covered with droves of horses and cattle, and
overrun by numberless wild rodents, the original tenants of the pampas.
During the long periods of drought, which are so great a scourge to the
country, these animals are starved by thousands, destroying, in their
efforts to live, every vestige of vegetation. In one of these 'siccos,'
at the time of my visit, no less than 50,000 head of oxen and sheep and
horses perished from starvation and thirst, after tearing deep out of
the soil every trace of vegetation, including the wiry roots of the
pampas-grass. Under such circumstances the existence of an unprotected
tree is impossible. The only plants that hold their own, in addition to
the indestructible thistles, grasses, and clover, are a little
herbaceous oxalis, producing viviparous buds of extraordinary vitality,
a few poisonous species, such as the hemlock, and a few tough, thorny
dwarf-acacias and wiry rushes, which even a starving rat refuses.

"Although the cattle are a modern introduction, the numberless
indigenous rodents must always have effectually prevented the
introduction of any other species of plants; large tracts are still
honeycombed by the ubiquitous biscacho, a gigantic rabbit; and numerous
other rodents still exist, including rats and mice, pampas-hares, and
the great nutria and carpincho (capybara) on the river banks."[9]

Mr. Clark further remarks on the desperate struggle for existence which
characterises the bordering fertile zones, where rivers and marshy
plains permit a more luxuriant and varied vegetable and animal life.
After describing how the river sometimes rose 30 feet in eight hours,
doing immense destruction, and the abundance of the larger carnivora and
large reptiles on its banks, he goes on: "But it was among the flora
that the principle of natural selection was most prominently displayed.
In such a district--overrun with rodents and escaped cattle, subject to
floods that carried away whole islands of botany, and especially to
droughts that dried up the lakes and almost the river itself--no
ordinary plant could live, even on this rich and watered alluvial
debris. The only plants that escaped the cattle were such as were either
poisonous, or thorny, or resinous, or indestructibly tough. Hence we had
only a great development of solanums, talas, acacias, euphorbias, and
laurels. The buttercup is replaced by the little poisonous yellow oxalis
with its viviparous buds; the passion-flowers, asclepiads, bignonias,
convolvuluses, and climbing leguminous plants escape both floods and
cattle by climbing the highest trees and towering overhead in a flood of
bloom. The ground plants are the portulacas, turneras, and cenotheras,
bitter and ephemeral, on the bare rock, and almost independent of any
other moisture than the heavy dews. The pontederias, alismas, and
plantago, with grasses and sedges, derive protection from the deep and
brilliant pools; and though at first sight the 'monte' doubtless
impresses the traveller as a scene of the wildest confusion and ruin,
yet, on closer examination, we found it far more remarkable as a
manifestation of harmony and law, and a striking example of the
marvellous power which plants, like animals, possess, of adapting
themselves to the local peculiarities of their habitat, whether in the
fertile shades of the luxuriant 'monte' or on the arid, parched-up
plains of the treeless pampas."

A curious example of the struggle between plants has been communicated
to me by Mr. John Ennis, a resident in New Zealand. The English
water-cress grows so luxuriantly in that country as to completely choke
up the rivers, sometimes leading to disastrous floods, and necessitating
great outlay to keep the stream open. But a natural remedy has now been
found in planting willows on the banks. The roots of these trees
penetrate the bed of the stream in every direction, and the water-cress,
unable to obtain the requisite amount of nourishment, gradually
disappears.

_Increase of Organisms in a Geometrical Ratio_.

The facts which have now been adduced, sufficiently prove that there is
a continual competition, and struggle, and war going on in nature, and
that each species of animal and plant affects many others in complex and
often unexpected ways. We will now proceed to show the fundamental cause
of this struggle, and to prove that it is ever acting over the whole
field of nature, and that no single species of animal or plant can
possibly escape from it. This results from the fact of the rapid
increase, in a geometrical ratio, of all the species of animals and
plants. In the lower orders this increase is especially rapid, a single
flesh-fly (Musca carnaria) producing 20,000 larvae, and these growing so
quickly that they reach their full size in five days; hence the great
Swedish naturalist, Linnaeus, asserted that a dead horse would be
devoured by three of these flies as quickly as by a lion. Each of these
larvae remains in the pupa state about five or six days, so that each
parent fly may be increased ten thousand-fold in a fortnight. Supposing
they went on increasing at this rate during only three months of summer,
there would result one hundred millions of millions of millions for each
fly at the commencement of summer,--a number greater probably than
exists at any one time in the whole world. And this is only one species,
while there are thousands of other species increasing also at an
enormous rate; so that, if they were unchecked, the whole atmosphere
would be dense with flies, and all animal food and much of animal life
would be destroyed by them. To prevent this tremendous increase there
must be incessant war against these insects, by insectivorous birds and
reptiles as well as by other insects, in the larva as well as in the
perfect state, by the action of the elements in the form of rain, hail,
or drought, and by other unknown causes; yet we see nothing of this
ever-present war, though by its means alone, perhaps, we are saved from
famine and pestilence.

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A way with tears makes Magorian a worthy Costa winner
Articles published by guardian.co.uk Books

Perfumes: the Guide – a portal to a whole new art

Michelle Magorian scooped the 2008 Costa Children's Book Award with Just Henry, a huge 700-page book that made me cry. Not many authors can do that but Magorian handles dangerously emotional stuff and pulls it off without slipping into mawkish sentimentality. Hence tears.

The same quality marked out Goodnight Mister Tom, her first novel, which won the 1980 Guardian children's book prize and has been read by every child in year 6 and many others both younger and older – rightly so – ever since. Goodnight Mister Tom is avowedly weepy. Only the hardest heart could remain unmoved. I once met a child who'd sticky-taped three pages together because they made her cry too much – I'm sure everyone who's read the book will know which three.

In Goodnight Mister Tom, Magorian had the external drama of the second world war as an emotional backdrop: put simply, there was a lot to weep over. In Just Henry, however, the setting is 1949 and there should be – and is – a feeling of optimism and hope. It is a period that's rarely used in fiction but Just Henry reveals it to be one that's worth exploring. The effect of the war is still being felt in the social changes it brought about. Life didn't just "slip back": few families were lucky enough to remain unaffected. Fathers were lost or altered; mothers found themselves raising families alone, or having to return abruptly to a subordinate role; children were forced to make adjustments either way.

In her big, bold novel, knitted together with more mysteries and coincidences than are credible, Magorian wonderfully captures that uncertainty and shows children's ability to move forward and embrace change far faster than their parents or grandparents. Lest this realism and the solving of the mysteries is too mundane, Michelle adds an extra layer of emotion by weaving in the stories of film stars from the movies of the day. For once, the current fashion of long, long, long books is justified. Just Henry is a wallowing great read. Just don't forget your hanky.

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Leona Lewis to write autobiography

I touched on Perfumes: the Guide by Luca Turin and Tania Sanchez in today's G2 arts diary. What a wonderful book (I've just opened it and, in a synaesthesic overture, it's offering me Amouage Gold, a spray-sample of which I left between its leaves). It offers a critical analysis – smelling notes, if you will – of 1,500 perfumes. I suppose the authors' language and technique relates somewhat to wine criticism. But the art of writing about something so emotionally rich and elusive (and at the same time entirely unlockable by the proper technical expertise) reminded me a bit of how people write (or try to write) about music (a subject I'm speaking about at this year's Association of British Orchestras annual conference). As it happens, Turin and Sanchez often use musical metaphor to help explain the nature of a perfume (they talk in terms of "brassy" or "melodic line" or "string section"; Shalimar has a "uniquely sweet, penetrating tune"; Yatagan a "high-pitched, hissing tone"). What about a job swap between these two and Andrew Clements or Alexis Petridis, I wonder.

Over Christmas I did a lot of smelling in the various perfumery halls and perfumery shops of London, and had enormous fun trying to get to grips with the artform. The joy of it is that most perfumes are widely available and can be squirted by the curious with impunity. The scent I've been wearing for the past few years – Irisia by Creed – Turin and Sanchez write off, with a cutting one-star review, as a "green floral chypre of exceptional banality and unpleasantness", so I have have had the amusement of trying to find a replacement.

By the way, Sanchez notes the importance of the web in writing and sharing knowledge about perfume. She writes: "Until recently, talking intelligently about the art of perfume seemed impossible. Then suddenly it seemed inevitable. What changed? The obvious: the Internet. Online now you can read historical and technical information, find discontinued or otherwise elusive perfumes, order samples of raw materials to smell out of curiosity, and, most important, find communities of people clustered around this single obsession. Half of what I know I owe to the 24-hour-a-day pajama party that is the fragrance board of Makeup Alley... Online communities can criticise perfume in a way that magazines have never dared: there's no advertising to lose... Perfume blogs now seem to outnumber the sample vials around my desk: there are men and women of intelligence sitting down every day and thinking and writing about perfume."

It's a bit like the way Alex Ross talks about the way the web has affected his relationship with and access to contemporary music. It's that old long tail.

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