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       Darwin 
      Evolution 
      
          
      Chapter
      IV of The Origin of Species 
      Natural
      Selection; or the Survival of the Fittest 
      
      
      
      
      
      Natural Selection— 
      its power compared with man's selection— 
      its power on characters of trifling importance— 
      its power at all ages
      and on both sexes— 
      Sexual selection—On the generality of intercrosses 
      between individuals of the same species— 
      Circumstances favourable
      and unfavourable 
      to the results of natural selection, namely,
      intercrossing, 
      isolation, number of individuals— 
      Slow action—Extinction caused by
      natural selection—Divergence of character, related to the diversity 
      of inhabitants of any small area,
      and to naturalization—Action of
      natural selection, through divergence 
      of character and extinction,
      on the descendants 
      from a common parent— 
      Explains the grouping
      of all organic beings— 
      Advance in organization—Low forms preserved— 
      Convergence of character—I 
      definite multiplication of species
      — 
      Summary.
      
       
        
      How will the struggle
      for existence, briefly discussed in the last chapter, act in regard to
      variation? Can the principle of selection which we have seen is so potent
      in the hands of man, apply under nature? I think we shall see that it can
      act most efficiently. Let the endless number of slight variations and
      individual differences occurring in our domestic productions, and, in a
      lesser degree, in those under nature, be borne in mind; as well as the
      strength of the hereditary tendency. Under domestication, it may be truly
      said that the who]e organization becomes in some degree plastic. But the
      variability, which we almost universally meet with in our domestic
      productions, is not directly produced, as Hooker and Asa Gray have well
      remarked, by man; he can neither originate varieties, nor prevent their
      occurrence; he can preserve and accumulate such as do occur. Unintentionally
      he exposes organic beings to new and changing conditions of life, and
      variability ensues, but similar changes of conditions might and do occur
      under nature. Let it also be borne mind how infinitely complex and
      close-fitting arc the mutual relations of all organic beings to each other
      and to their physical conditions of life; and consequently what infinitely
      varied diversities of structure might be of use to each being under
      changing conditions of life. Can it, then, be thought improbable seeing
      that variations useful to man have undoubtedly occurred that other
      variations useful in some way to each being in the great and complex
      battle of life, should occur in the course of many successive generations.
      If such do occur, can we doubt (remembering that many more individuals are
      born than can possibly survive) that individuals having any advantage,
      however slight, over others should have the best chance of surviving and
      of procreating their kind? On the other hand, we may feel sure that any
      variation in the least degree injurious would be rigidly destroyed. This
      preservation of favourable individual differences and variations, and the
      destruction of those which are injurious, I have called Natural Selection
      or the Survival of the Fittest. Variations neither useful nor injurious
      would not be affected by natural selection, and would be left either a
      fluctuating element, as perhaps we see in certain polymorphic species, or
      would ultimately become fixed, owing to the nature of the organism and the
      nature of the conditions.
      
       
      Several writers have
      misapprehended or objected to the term Natural Selection. Some have even
      imagined that natural selection induces variability, whereas it implies
      only the preservation of such variations as arise and are beneficial to
      the being under its conditions of life. No one objects to agriculturists
      speaking of the potent effects of man's selection; and in this case the
      individual differences given by nature, which man for some object selects
      must of necessity first occur. Others have objected that the term
      selection implies conscious choice in the animals which become modified;
      and It had even been urged that, as plants have no volition, natural
      selection is not applicable to them! In the literal sense of the word, no
      doubt, natural selection is a false term, but who ever objected to
      chemists speaking of the elective affinities of the various
      elements?—and yet an acid cannot strictly be said to elect the base with
      which it in preference combines. It has been said that I speak of natural
      selection as an active power or Deity; but who objects to an author
      speaking of the attraction of gravity as ruling the movements of the
      planets? Every one knows what is meant and is implied by such metaphorical
      expressions; and they are almost necessary for brevity. So again it Is
      difficult to avoid personifying the word Nature; but I mean by Nature,
      only the aggregate action and product of many natural laws, and by laws
      the sequence of events as ascertained by us. With a little familiarity
      such superficial objections will be forgotten. We shall best understand
      the probable course of natural selection by taking the case of a country
      undergoing some slight physical change, for instance, of climate. The
      proportional numbers of Its inhabitants will almost immediately undergo a
      change, and some species will probably become extinct. We may conclude,
      from what we have seen of the intimate and complex manner in which the
      inhabitants of each country are bound together, that any change in the
      numerical proportions of the inhabitants, independently of the change of
      climate itself, would seriously affect the others If the country were open
      on its borders, new forms would certainly immigrate, and this would
      likewise seriously disturb the relations of some of the former
      inhabitants. Let it be remembered how powerful the influence of a single
      introduced tree or mammal has been shown to be. But in the case of an
      island, or of a country partly surrounded by barriers, into which new and
      better adapted forms could not freely enter, we should then have places in
      the economy of nature which would assuredly be better filled up, if some
      of the original inhabitants were in some manner modified; for, had the
      area been open to immigration, these same places would have been seized on
      by intruders. In such cases, slight modifications, which in any way
      favoured the individuals of any species, by better adapting them to their
      altered conditions, would tend to be preserved; and natural selection
      would have free scope for the work of improvement. 
      We have good reason to
      believe, as shown in the first chapter, that changes in the conditions of
      life give a tendency to increased variability—and in the foregoing cases
      the conditions have changed, and this would manifestly be favourable to
      natural selection, by affording a better chance of the occurrence of
      profitable variations. Unless such occur, natural selection can do
      nothing. Under the term of “variations,” it must never be forgotten
      that mere individual differences are included. As man can produce a great
      result with his domestic animals and plants by adding up in any given
      direction individual differences, so could natural selection, but ar more
      easily from having incomparably longer time for action. Nor do I believe
      that any great physical change, as of climate, or any unusual degree of
      isolation to check immigration, is necessary in order that new and
      unoccupied places should be left, for natural selection to fill up by
      improving some of the varying inhabitants with nicely balanced forces,
      extremely slight modifications in the structure or habits of one species
      would often give it an advantage over others; and still further
      modifications of the same kind would often still further increase the
      advantage, as long as the species continued under the same conditions of
      life and profited by similar means of subsistence and defence. No country
      can be named m which all the native inhabitants are now so perfectly
      adapted to each other and to the physical conditions under which they
      live, that none of them could be still better adapted or improved; for in
      all countries, the natives have been so far conquered by naturalized
      productions, that they have allowed some foreigners to take firm
      possession of the land. And as foreigners have thus in every country
      beaten some of the natives, we may safely conclude that the natives might
      have been modified with advantage, so as to have better resisted the
      intruders.
      
       
      As man can produce, and
      certainly has produced, a great result by his methodical and unconscious
      means of selection, what may not natural selection effect? Man can act
      only on external and visible characters: Nature, if I may be allowed to
      personify the natural preservation or survival of the fittest, cares
      nothing for appearances, except in so far as they are useful to any being.
      She can act on every internal organ, on every shade of constitutional
      difference, on the whole machinery of life. Man selects only for his own
      good: Nature only for that of the being which she tends. Every selected
      character Is fully exercised by her, as is implied by the fact of their
      selection. Man keeps the natives of many climates in the same country; he
      seldom exercises each selected character in some peculiar and fitting
      manner; he feeds a long and a short beaked pigeon on the same food; he
      does not exercise a long-backed or long-legged quadruped in any peculiar
      manner; he exposes sheep with long and short wool to the same climate. He
      does not allow the most vigorous males to struggle for the females. He
      does not rigidly destroy all inferior animals, but protects during each
      varying season as far as lies in his power, all his productions. He often
      begins his selection by some half-monstrous form; or at least by some
      modification prominent enough to catch the eye or to be plainly useful to
      him. Under Nature, the slightest differences of structure or constitution
      may well turn the nicely balanced scale in the struggle for life, and so
      be preserved. How fleeting are the wishes and efforts of man, how short
      his time! And consequently how poor will be his results, compared with
      those accumulated by Nature during whole geological periods! Can we
      wonder, then, that Nature's productions should be far “truer” in
      character than man's productions that they should be infinitely better
      adapted to the most complex conditions of life and should plainly bear the
      stamp of far higher workmanship?
      
       
      It may metaphorically
      be said that natural selection is daily and hourly scrutinising,
      throughout the world, the slightest variations; rejecting those that are
      bad, preserving and adding up all that are good, silently and insensibly
      working, whenever and wherever opportunity offers, at the improvement of
      each organic being in relation to its organic and inorganic conditions of
      life. We see nothing of these slow changes in progress, until the hand of
      time has marked the lapse of ages, and then so imperfect is our view into
      long-past geological ages, that we see only that the forms of life are now
      different from what they formerly were. In order that any great amount of
      modification should be seen in a species, a variety when once formed must
      again, perhaps after a long interval of time, vary or present individual
      differences of the same favourable nature as before; and these must be
      again preserved, and so onwards step by step. Seeing that individual
      differences of the same kind perpetually recur, this can hardly be
      considered as an unwarrantable assumption. But whether it is true, we can
      judge only by seeing how far the hypothesis accords with and explains the
      general phenomena of nature. On the other hand, the ordinary belief that
      the amount of possible variation is a strictly limited quantity is
      likewise a simple assumption. Although natural selection can act only
      through and for the good of each being, yet characters and structures,
      which we are apt to consider as of very trifling importance, may thus be
      acted on. When we see leaf-eating insects green, and bark-feeders mottled-grey;
      the alpine ptarmigan white in winter, the red-grouse the colour of
      heather, we must believe that these tints are of service to these birds
      and insects in preserving them from danger. Grouse, if not destroyed at
      some period of their lives would increase in countless numbers- they are
      known to suffer largely from birds of prey; and hawks are guided by
      eyesight to their prey--so much so, that on parts of the Continent persons
      are warned not to keep white pigeons, as being the most liable to
      destruction. Hence natural selection might be effective in giving the
      proper colour to each kind of grouse, and in keeping that colour, when
      once acquired, true and constant. Nor ought we to think that the
      occasional destruction of an animal of any particular colour would produce
      little effect: we should remember how essential it is in a flock of white
      sheep to destroy a lamb with the faintest trace of black. We have seen how
      the colour of the hogs, which feed on the “paint-root” in Virginia,
      determines whether they shall live or die. In plants, the down on the
      fruit and the colour of the flesh are considered by botanists as
      characters of the most trifling importance: yet we hear from an excellent
      horticulturist, Downing, that in the United States, smooth-skinned fruits
      suffer far more from a beetle, a Curculio, than those with down; that
      purple plums suffer far more from a certain disease than yellow plums;
      whereas another disease attacks yellow-fleshed peaches far more than those
      with other coloured flesh. If, with all the aids of art, these slight
      differences make a great difference in cultivating the several varieties,
      assuredly, in a state of nature, where the trees would have to struggle
      with other trees, and with a host of enemies, such differences would
      effectually settle which variety, whether a smooth or downy, a yellow or
      purple fleshed fruit, should succeed.
      
       
      In looking at many
      small points of difference between species which, as far as our ignorance
      permits us to judge, seem quite unimportant, we must not forget that
      climate, food, &c., have no doubt produced some direct effect. It is
      also necessary to bear in mind that, owing to the law of correlation, when
      one part varies an the variations are accumulated through natural
      selection, other modifications, often of the most unexpected nature, will
      ensue.
      
       
      As we see that those
      variations which, under domestication, appear at any particular period of
      life, tend to reappear in the offspring at the same period;--for instance,
      in the shape, size, and flavour of the seeds of the many varieties of our
      culinary and agricultural plants; in the caterpillar and cocoon stages of
      the varieties of the silk-worm; in the eggs of poultry, and in the colour
      of the down of their chickens; in the horns of our sheep and cattle when
      nearly adult;—so in a state of nature natural selection will be enabled
      to act on and modify organic beings at any age, by the accumulation of
      variations profitable at that age, and by their inheritance at a
      corresponding age. If it profit a plant to have its seeds more and more
      widely disseminated by the wind, I can see no greater difficulty in this
      being effected through natural selection, than in the cotton-planter
      increasing and improving by selection the down in the pods on his
      cotton-trees. Natural selection may modify and adapt the larva of an
      insect to a score of contingencies, wholly different from those which
      concern the mature insect, and these modifications may effect, through
      correlation, the structure of the adult So, conversely, modifications in
      the adult may affect the structure of the larva; but m all cases natural
      selection will ensure that they shall not be injurious: for if they were
      so, the species would become extinct.
      
       
      Natural selection will
      modify the structure of the young in relation to the parent, and of the
      parent in relation to the young. In social animals it will adapt the
      structure of each individual for the benefit of the whole community; if
      the community profits by the selected change. What natural selection
      cannot do, is to modify the structure of one species, without giving it
      any advantage, for the good of another species; and though statements to
      this effect may be found in works of natural history, I cannot find one
      case which will bear investigation. A structure used only once in an
      animal's life, if of high importance to it, might be modified to any
      extent by natural selection; for instance, the great jaws possessed by
      certain insects, used exclusively for opening the cocoon—or the hard tip
      to the beak of unhatched birds, used for breaking the egg. It has been
      asserted, that of the best short-beaked tumbler-pigeons a greater number
      perish in the egg than are able to get out n it; so that fanciers assist
      in the act of hatching. Now if nature had to make the beak of a full-grown
      pigeon very short for the bird's own advantage, the process of
      modification would be very slow, and there would be simultaneously the
      most rigorous selection of all the young birds within the egg, which had
      the most powerful and hardest beaks, for all with weak beaks would
      inevitably perish; or, more delicate and more easily broken shells might
      be selected, the thickness of the shell being known to vary like every
      other structure. It may be well here to remark that with all beings there
      must be much fortuitous destruction, which can have little or no influence
      on the course of natural selection. For instance a vast number of eggs or
      seeds are annually devoured, and these could be modified through natural
      selection only if they varied in some manner which protected them from
      their enemies. Yet many of these eggs or seeds would perhaps, if not
      destroyed, have yielded individuals better adapted to their conditions of
      life than any of those which happened to survive. So again a vast number
      of mature animals and plants, whether or not they be the best adapted to
      their conditions, must be annually destroyed by accidental causes, which
      would not be in the least degree mitigated by certain changes of structure
      or constitution which would in other ways be beneficial to the species.
      But let the destruction of the adults be ever so heavy, If the number
      which can exist in any district be not wholly kept down by such
      causes,—or again let the destruction of eggs or seeds be so great that
      only a hundredth or a thousandth part are developed,—yet of those which
      do survive, the best adapted individuals, supposing that there is any
      variability in a favourable direction, will tend to propagate their kind
      in larger numbers than the less well adapted. If the numbers be wholly
      kept down by the causes just indicated, as will often have been the case,
      natural selection will be powerless in certain beneficial directions; but
      this is no valid objection to its efficiency at other times and in other
      ways; for we are far from having any reason to suppose that many species
      ever undergo modification and improvement at the same time in the same
      area.
      
       
       
      
       
      Sexual
      Selection
      
       
      Inasmuch as
      peculiarities often appear under domestication in one sex and become
      hereditarily attached to that sex, so no doubt it will be under nature.
      Thus it is rendered possible for the two sexes to be modified through
      natural selection in relation to different habits of life, as Is sometimes
      the case; or for one sex to be modified in relation to the other sex, as
      commonly occurs. This leads me to say a few words on what I have called
      Sexual Selection. This form of selection depends, not on a struggle for
      existence in relation to other organic beings or to external conditions,
      but on a struggle between the individuals of one sex, generally the males,
      for the possession of the other sex. The result is not death to the
      unsuccessful competitor, but few or no offspring. Sexual selection is,
      therefore less rigorous than natural selection. Generally, the most
      vigorous males, those which are best fitted for their places in nature,
      will leave most progeny. But in many cases, victory depends not so much on
      general vigor, as on having special weapons, confined to the male sex. A
      hornless stag or spurless cock would have poor chance of leaving numerous
      offspring. Sexual selection, by always allowing the victor to breed, might
      surely give indomitable courage, length to the spur, and strength to the
      wing to strike in the spurred leg, in nearly the same manner as does the
      brutal cock-fighter by the careful selection of his best cocks. How low in
      the scale of nature the law of battle descends, I know not; male
      alligators have been described as fighting, bellowing, and whirling round,
      like Indians in a war dance, for the possession of the females; male
      salmons have been observed fighting all day long; male stag-beetles
      sometimes bear wounds from the huge mandibles of other males, the males of
      certain hymenopterous insects have been frequently seen by that Inimitable
      observer M. Fabre, fighting for a particular female who sits by, an
      apparently unconcerned beholder of the struggle, and then retires with the
      conqueror. The war is, perhaps, severest between the males of polygamous
      animals, and these seem oftenest provided with special weapons. The males
      of carnivorous animals are already well armed; though to them and to
      others, special means of defence may be given through means of sexual
      selection, as the mane of the lion, and the hooked jaw to the male salmon,
      for the shield may be as important for victory, as the sword or spear
      
       
      Amongst birds, the
      contest is often of a more peaceful character All those who have attended
      to the subject, believe that there is the severest rivalry between the
      males of many species to attract, by singing, the females. The rock-thrush
      of Guiana, birds of paradise and some others, congregate; and successive
      males display with the most elaborate care, and show off in the best
      manner, their gorgeous plumage; they likewise perform strange antics
      before the females which, standing by as spectators, at last choose the
      most attractive partner. Those who have closely attended to birds in
      confinement well know that they often take individual preferences and
      dislikes: Sir R. Heron has described how a pied peacock was eminently
      attractive to all his hen birds. I cannot here enter on the necessary
      details, but if man can in a short time give beauty and an elegant
      carriage to his bantams, according to the standard of beauty, I can see no
      good reason to doubt that female birds, by selecting, during thousands of
      generations, the most melodious or beautiful males according to their
      standard of beauty, might produce a marked effect. Some well-known laws,
      with respect to the plumage of male and female birds, in comparison with
      the plumage of the young, can partly be explained through the action of
      sexual selection on variations occurring at different ages, and
      transmitted to the males alone or to both sexes at corresponding ages; but
      I have not space here to enter on this subject.
      
       
      Thus it is, as I
      believe, that when the males and females of any animal have the same
      general habits of life, but differ in structure, colour, or ornament, such
      differences have been mainly caused by sexual selection: that is, by
      individual males having had, in successive generations, some slight
      advantage over other males, in their weapons, means of defence, or charms,
      which they have transmitted to their male offspring alone. Yet, I would
      not wish to attribute a sexual differences to this agency: for we see in
      our domestic animals peculiarities arising and becoming attached to the
      male sex, which apparently have not been augmented through selection by
      man. The tuft of hair on the breast of the wild turkey-cock cannot be of
      any use, and it is doubtful whether it can be ornamental in the eyes of
      the female bird;—indeed, had the tuft appeared under domestication, it
      would have been called a monstrosity.
      
       
       
      
       
      Illustrations
      of the Action of Natural Selection, or the Survival of the Fittest
      
       
      In order to make it
      clear how, as I believe, natural selection acts, I must beg permission to
      give one or two imaginary illustrations. Let us take the case of a wolf,
      which preys on various animals, securing some by craft, some by strength
      and some by fleetness; and let us suppose that the fleetest prey, a deer
      for instance, had from any change in the country increased in numbers or
      that other prey had decreased in numbers, during that season of the year
      when the wolf was hardest pressed for food. Under such circumstances the
      swiftest and slimmest wolves would have the best chance of surviving and
      so be preserved or selected,--provided always that they retained strength
      to master their prey at this or some other period of the year, when they
      were compelled to prey on other animals, can see no more reason to doubt
      that this would be the result, than that man should be able to improve the
      fleetness of his greyhounds y careful and methodical selection, or by that
      kind of unconscious selection which follows from each man trying to keep
      the best dogs without any thought of modifying the breed. I may add, that,
      according to Mr. Pierce, there are two varieties of the wolf inhabiting
      the Catskill Mountains, in the United States, one with a light
      greyhound-like form, which pursues deer, and the other more bulky, with
      shorter legs, which more frequently attacks the sheep
      
       
      It should be observed
      that, in the above illustration, I speak of the slimmest individual
      wolves, and not of any single strongly-marked variation having been
      preserved. In former editions of this work I sometimes spoke as if this
      latter alternative had frequently occurred. I saw the great importance of
      individual differences, and this led me fully to discuss the results of
      unconscious selection by man, which depends on the preservation of all the
      more or less valuable individuals, and on the destruction of the worst. I
      saw also, that the preservation in a state of nature of any occasional
      deviation of structure, such as a monstrosity, would be a rare event and
      that, If at first preserved, it would generally be lost by subsequent
      Intercrossing with ordinary individuals. Nevertheless, until reading an
      able and valuable article in the ‘North British Review’ (l867), I did
      not appreciate how rarely single variations, whether slight or strongly
      marked, could be perpetuated. The author takes the case of a pair of
      animals, producing during their lifetime two hundred offspring, of which,
      from various causes of destruction, only two on an average survive to
      procreate their kind. This is rather an extreme estimate for most of the
      higher animals, but by no means so for many of the lower organisms. He
      then shows that if a single Individual were born, which varied in some
      manner, giving it twice as good a chance of life as that of the other
      individuals, yet the chances would be strongly against its survival.
      Supposing it to survive and to breed, and that half its young inherited
      the favourable variation; still, as the Reviewer goes on to show, the
      young would have only a slightly better chance of surviving and breeding;
      and this chance would go on decreasing in the succeeding generations. The
      justice of these remarks cannot, I think, be disputed. If for instance, a
      bird of some kind could procure its food more easily by having its beak
      curved, and if one were born with its beak strongly curved, and which
      consequently flourished, nevertheless ere would be a very poor chance of
      this one individual perpetuating its kind to the exclusion of the common
      form; but there can hardly be a doubt, judging by what we see taking place
      under domestication, that this result would follow from the presentation
      during many generations of a large number of individuals with more or less
      strongly curved beaks, and from the destruction of a still larger number
      with the straightest beaks.
      
       
      It should not, however,
      be overlooked that certain rather strongly marked variations, which no one
      would rank as mere individual differences, frequently recur owing to a
      similar organization being similarly acted on—of which fact numerous
      instances could be given with our domestic productions. In such cases, If
      the varying individual did not actually transmit to its offspring its
      newly-acquired character, it would undoubtedly transmit to them, as long
      as the existing conditions remained the same, a still stronger tendency to
      vary in the same manner. There can also be little doubt that the tendency
      to vary in the same manner has often been so strong that all the
      individuals of the same species have been similarly modified without the
      aid of any form of selection. Or only a third, fifth, or tenth part of the
      individuals may have been thus affected, of which fact several instances
      could be given. Thus Graba estimates that about one-fifth of the
      guillemots in the Faroe Islands consist of a variety so well marked, that
      it was formerly ranked as a distinct species under the name of Uria
      lacrymans. In cases of this kind, if the variation were of a beneficial
      nature, the original form would soon be supplanted by the modified form,
      through the survival of the fittest.
      
       
      To the effects of
      intercrossing in eliminating variations of all kinds, I shall have to
      recur; but it may be here remarked that most animals and plants keep to
      their proper homes, and do not needlessly wander about; we see this even
      with migratory birds, which almost always return to the same spot.
      Consequently each newly-formed variety would generally be at first local,
      as seems to be the common rule with varieties in a state of nature; so
      that similarly modified individuals would soon exist in a small body
      together, and would often breed together. If the new variety were
      successful in its battle for life, it would slowly spread from a central
      district, competing with and conquering the unchanged individuals on the
      margins of an ever-increasing circle.
      
       
      It may be worth while
      to give another and more complex illustration of the action of natural
      selection. Certain plants excrete sweet juice, apparently for the sake of
      eliminating something injurious from the sap: this is effected, for
      instance, by glands at the base of the stipules in some Leguminosa, and at
      the backs of the leaves of the common laurel. This juice, though small in
      quantity, is greedily sought by insects; but their visits do not in any
      way benefit the plant . Now, let us suppose that the juice or nectar was
      excreted from the inside of the flowers of a certain number of plants of
      any species. Insects in seeking the nectar would get dusted with pollen,
      and would often transport it from one flower to another. The flowers of
      two distinct individuals of the same species would thus get crossed, and
      the act of crossing, as can be fully proved, gives rise to vigorous
      seedlings which consequently would have the best chance of flourishing and
      surviving. The plants which produced flowers with the largest glands or
      nectaries, excreting most nectar, would oftenest be visited by insects,
      and would oftenest be crossed; and so in the long-run would gain the upper
      hand and form a local variety. The flowers, also, which had their stamens
      and pistils placed, in relation to the size and habits of the particular
      insects which visited them, so as to favour in any degree the transportal
      of the pollen, would likewise be favoured. We might have taken the case of
      insects visiting flowers for the sake of collecting pollen instead of
      nectar; and as pollen is formed for the sole purpose of fertilization, its
      destruction appears to be a simple loss to the plant; yet if a little
      pollen were carried, at first occasionally and then habitually, by the
      pollen-devouring insects from flower to flower, and a cross thus effected,
      although nine-tenths of the pollen were destroyed it might still be a
      great gain to the plant to be thus robbed, and the individuals which
      produced more and more pollen, and had larger anthers, would be selected.
      
       
      When our plant, by the
      above process long continued, had been rendered highly attractive to
      insects, they would, unintentionally on their part, regularly carry pollen
      from flower to flower, and that they do this effectually, I could easily
      show by many striking facts. I will give only one, as likewise
      illustrating one step in the separation of the sexes of plants. Some
      holly-trees bear only male flowers, which have four stamens producing a
      rather small quantity of pollen, and a rudimentary pistil: other
      holly-trees bear only female flowers; these have a full-sized pistil, and
      four stamens with shrivelled anthers, in which not a grain of pollen can
      be detected. Having found a female tree exactly sixty yards from a male
      tree, I put the stigmas of twenty flowers, taken from different branches
      under the microscope, and on all, without exception, there were a few
      pollen-grains, and on some a profusion. As the wind had set for several
      days from the female to the male tree, the pollen could not thus have been
      carried. The weather had been cold and boisterous and therefore not
      favourable to bees, nevertheless every female flower which I examined had
      been effectually fertilized by the bees which had flown from tree to tree
      in search of nectar. But to return to our imaginary case: as soon as the
      plant had been rendered so highly attractive to insects that pollen was
      regularly carried from flower to flower, another process might commence.
      No naturalist doubts the advantage of what has been called the
      "physiological division of labour"; hence we may believe that it
      would be advantageous to a plant to produce stamens alone in one flower or
      on one whole plant, and pistils alone in another flower or on another
      plant. In plants under culture and placed under new conditions of life
      sometimes the male organs and sometimes the female organs become more or
      less impotent; now if we suppose this to occur in ever so slight a degree
      under nature, then, as pollen is already carried regularly from flower to
      flower, and as a more complete separation of the sexes of our plant would
      be advantageous on the principle of
      
       
      the division of labour,
      individuals with this tendency more and more increased would be
      continually favoured or selected, until at last a complete separation of
      the sexes might be effected. It would take up too much space to show the
      various steps, through dimorphism and other means, by which the separation
      of the sexes m plants of various kinds is apparently now in progress; but
      I may add that some of the species of holly in North America, are,
      according to Asa Gray, in an exactly intermediate condition, or, as he
      expresses It, are more or less dioeciously polygamous.
      
       
      Let us now turn to the
      nectar-feeding insects; we may suppose the plant, of which we have been
      slowly increasing the nectar by continued selection, to be a common plant;
      and that certain insects depended in main part on its nectar for food. I
      could give many facts showing how anxious bees are to save time: for
      instance, their habit of cutting holes and sucking the nectar at the bases
      of certain flowers, which, with a very little more trouble, they can enter
      by the mouth. Bearing such facts in mind, it may be believed that under
      certain circumstances individual differences in the curvature or length of
      the proboscis, &c., too slight to be appreciated by us, might profit a
      bee or other insect, so that certain individuals would be able to obtain
      their food more quickly than others; and thus the communities to which
      they belonged would flourish and throw off many swarms inheriting the same
      peculiarities. The tubes of the corolla of the common red and incarnate
      clovers (Trifolium pratense and incarnatum) do not on a hasty glance
      appear to differ in length; yet the hive-bee can easily suck the nectar
      out of the incarnate clover, but not out of the common red clover, which
      is visible by humble-bees alone; so that whole fields of red clover offer
      in vain an abundant supply of precious nectar to the hive-bee. That this
      nectar is much liked by the hive-bee is certain; for I have repeatedly
      seen, but only in the autumn, many hive-bees sucking the flowers through
      holes bitten in the base of the tube by bumble-bees. The difference in the
      length of the corolla in the two kinds of clover, which determines the
      visits of the hive-bee, must be very trifling; for I have been assured
      that when red clover has been mown, the flowers of the second crop are
      somewhat smaller, and that these are visited by many hive-bees. I do not
      know whether this statement Is accurate, nor whether another published
      statement can be trusted, namely, that the Ligurian bee which is generally
      considered a mere variety of the common hive-bee, and which freely crosses
      with It, Is able to reach and suck the nectar of the red clover. Thus, in
      a country where this kind of clover abounded, it might be a great
      advantage to the hive-bee to have a slightly longer or differently
      constructed proboscis. On the other hand, as the fertility of this clover
      absolutely depends on bees visiting the flowers, if humble-bees were to
      become rare in any country, it might be a great advantage to the plant to
      have a shorter or more deeply divided corolla, so that the hive-bees
      should be enabled to suck its flowers. Thus I can understand how a flower
      and a bee might slowly become, either simultaneously or one after the
      other, modified and adapted to each other In the most perfect manner, by
      the continued preservation of all the individuals which presented slight
      deviations of structure mutually favourable to each other.
      
       
      I am well aware that
      this doctrine of natural selection, exemplified in the above imaginary
      instances, is open to the same objections which were first urged against
      Sir Charles Lyell's noble views on the modern changes of the earth, as
      illustrative of “geology”; but we now seldom hear the agencies which
      we see still at work spoken of as trifling or insignificant, when used in
      explaining the excavation of the deepest valleys of the formation of long
      lines of inland cliffs. Natural selection acts only by the preservation
      and accumulation of small inherited modifications, each profitable to the
      preserved being; and as modern geology has almost banished such views as
      the excavation of a great valley by a single diluvial wave, so will
      natural selection banish the belief of the continued creation of new
      organic beings, or of any great and sudden modification in their structure. 
      
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