CONTRIBUTIONS TO THE THEORY OF NATURAL SELECTION.
A Series of Essays.
ALFRED RUSSEL WALLACE,
"THE MALAY ARCHIPELAGO," ETC., ETC.
SECOND EDITION, WITH CORRECTIONS AND ADDITIONS.
MACMILLAN AND CO.
[The Right of Translation and Reproduction is reserved.]
PRINTED BY HEAD, HOLE & CO., FARRINGDON STREET,
AND IVY LANE, E.C.
The present volume consists of essays which I have contributed to various periodicals, or read before scientific societies during the last fifteen years, with others now printed for the first time. The two first of the series are printed without alteration, because, having gained me the reputation of being an independent originator of the theory of "natural selection," they may be considered to have some historical value. I have added to them one or two very short explanatory notes, and have given headings to subjects, to make them uniform with the rest of the book. The other essays have been carefully corrected, often considerably enlarged, and in some cases almost rewritten, so as to express more fully and more clearly the views which I hold at the present time; and as most of them originally appeared in publications which have a very limited circulation, I believe that the larger portion of this volume will be new to many of my friends and to most of my readers.
I now wish to say a few words on the reasons which have led me to publish this work. The second essay, especially when taken in connection with the first, contains an outline sketch of the theory of the origin of species (by means of what was afterwards termed by Mr. Darwin—"natural selection,") as conceived by me before I had the least notion of the scope and nature of Mr. Darwin's labours. They were published in a way not likely to attract the attention of any but working naturalists, and I feel sure that many who have heard of them, have never had the opportunity of ascertaining how much or how little they really contain. It therefore happens, that, while some writers give me more credit than I deserve, others may very naturally class me with Dr. Wells and Mr. Patrick Matthew, who, as Mr. Darwin has shown in the historical sketch given in the 4th and 5th Editions of the "Origin of Species," certainly propounded the fundamental principle of "natural selection" before himself, but who made no further use of that principle, and failed to see its wide and immensely important applications.
The present work will, I venture to think, prove, that I both saw at the time the value and scope of the law which I had discovered, and have since been able to apply it to some purpose in a few original lines of investigation. But here my claims cease. I have felt all my life, and I still feel, the most sincere satisfaction that Mr. Darwin had been at work long before me, and that it was not left for me to attempt to write "The Origin of Species." I have long since measured my own strength, and know well that it would be quite unequal to that task. Far abler men than myself may confess, that they have not that untiring patience in accumulating, and that wonderful skill in using, large masses of facts of the most varied kind,—that wide and accurate physiological knowledge,—that acuteness in devising and skill in carrying out experiments,—and that admirable style of composition, at once clear, persuasive and judicial,—qualities, which in their harmonious combination mark out Mr. Darwin as the man, perhaps of all men now living, best fitted for the great work he has undertaken and accomplished.
My own more limited powers have, it is true, enabled me now and then to seize on some conspicuous group of unappropriated facts, and to search out some generalization which might bring them under the reign of known law; but they are not suited to that more scientific and more laborious process of elaborate induction, which in Mr. Darwin's hands has led to such brilliant results.
Another reason which has led me to publish this volume at the present time is, that there are some important points on which I differ from Mr. Darwin, and I wish to put my opinions on record in an easily accessible form, before the publication of his new work, (already announced,) in which I believe most of these disputed questions will be fully discussed.
I will now give the date and mode of publication of each of the essays in this volume, as well as the amount of alteration they have undergone.
I.—ON THE LAW WHICH HAS REGULATED THE INTRODUCTION OF NEW SPECIES.
First published in the "Annals and Magazine of Natural History," September, 1855. Reprinted without alteration of the text.
II.—ON THE TENDENCY OF VARIETIES TO DEPART INDEFINITELY FROM THE ORIGINAL TYPE.
First published in the "Journal of the Proceedings of the Linnaean Society," August, 1858. Reprinted without alteration of the text, except one or two grammatical emendations.
III.—MIMICRY AND OTHER PROTECTIVE RESEMBLANCES AMONG ANIMALS.
First published in the "Westminster Review," July, 1867. Reprinted with a few corrections and some important additions, among which I may especially mention Mr. Jenner Weir's observations and experiments on the colours of the caterpillars eaten or rejected by birds.
IV.—THE MALAYAN PAPILIONIDAE, OR SWALLOW-TAILED BUTTERFLIES, AS ILLUSTRATIVE OF THE THEORY OF NATURAL SELECTION.
First published in the "Transactions of the Linnaean Society," Vol. XXV. (read March, 1864), under the title, "On the Phenomena of Variation and Geographical Distribution, as illustrated by the Papilionidae of the Malayan Region."
The introductory part of this essay is now reprinted, omitting tables, references to plates, &c., with some additions, and several corrections. Owing to the publication of Dr. Felder's "Voyage of the Novara" (Lepidoptera) in the interval between the reading of my paper and its publication, several of my new species must have their names changed for those given to them by Dr. Felder, and this will explain the want of agreement in some cases between the names used in this volume and those of the original paper.
V.—ON INSTINCT IN MAN AND ANIMALS.
Not previously published.
VI.—THE PHILOSOPHY OF BIRDS' NESTS.
First published in the "Intellectual Observer," July, 1867. Reprinted with considerable emendations and additions.
VII.—A THEORY OF BIRDS' NESTS; SHOWING THE RELATION OF CERTAIN DIFFERENCES OF COLOUR IN BIRDS TO THEIR MODE OF NIDIFICATION.
First published in the "Journal of Travel and Natural History" (No. 2), 1868. Now reprinted with considerable emendations and additions, by which I have endeavoured more clearly to express, and more fully to illustrate, my meaning in those parts which have been misunderstood by my critics.
VIII.—CREATION BY LAW.
First published in the "Quarterly Journal of Science," October, 1867. Now reprinted with a few alterations and additions.
IX.—THE DEVELOPMENT OF HUMAN RACES UNDER THE LAW OF NATURAL SELECTION.
First published in the "Anthropological Review," May, 1864. Now reprinted with a few important alterations and additions. I had intended to have considerably extended this essay, but on attempting it I found that I should probably weaken the effect without adding much to the argument. I have therefore preferred to leave it as it was first written, with the exception of a few ill-considered passages which never fully expressed my meaning. As it now stands, I believe it contains the enunciation of an important truth.
X.—THE LIMITS OF NATURAL SELECTION AS APPLIED TO MAN.
This is the further development of a few sentences at the end of an article on "Geological Time and the Origin of Species," which appeared in the "Quarterly Review," for April, 1869. I have here ventured to touch on a class of problems which are usually considered to be beyond the boundaries of science, but which, I believe, will one day be brought within her domain.
* * * * *
For the convenience of those who are acquainted with any of my essays in their original form, I subjoin references to the more important additions and alterations now made to them.
ADDITIONS AND CORRECTIONS TO THE ESSAYS AS ORIGINALLY PUBLISHED.
Essays I. and II. are unaltered, but short notes are added at pp. 19, 24, 29, and 40.
III.—Mimicry, and other Protective Resemblances among Animals.
53 Additional illustration of protective colouring in the case of the wood-dove and the robin.
63 On moths resembling bird's dung and mortar.
86 Correction of some names of African Papilios and a reference to Mr. Trimen's observations.
89 Mr. Jenner Weir's observation on birds which refused to eat Spilosoma menthrasti.
102 An additional case of snake mimicry in Oxyrhopus trigeminus.
107 Mr. Salvin's case of mimicry among hawks.
113 Name, Diadema anomala, added.
117 to 122. Use of gay colours in caterpillars, with an account of Mr. Jenner Weir's and Mr. Butler's observations.
IV.—The Malayan Papilionidae or Swallow-tailed Butterflies, as illustrative of the Theory of Natural Selection.
135 to 140. Additions to the discussion on the rank of the Papilionidae, and on the principles which determine the comparative rank of groups in the animal kingdom.
164 Illustration of variability from Mr. Baker's revision of the British Roses.
173 Additional facts, on local variations of colour.
196 Additional genus of birds (Ceycopsis) peculiar to Celebes.
199, 200. Concluding remarks.
VI.—The Philosophy of Birds' Nests.
218 On nesting of Terns and Gulls, rewritten.
220 to 222. Daines Barrington, and others, on the song of birds.
223 On young birds learning to build, by memory and imitation.
224 Levaillant, on mode of nest-building.
229 On imperfect adaptation in birds' nests.
VII.—A Theory of Birds' Nests.
231, 232. Introductory passages modified, with some omissions.
233 How modifications of organization would affect the form of the nest.
235 Illustration from the habits of children and savages.
235, 236. Objection to term "hereditary habit" answered.
237 Passage rewritten, on more or less variable characters in relation to nidification.
248 On males choosing or rejecting females, and on the various modes in which colour may be acquired by female birds.
249 On probable ancestral colours of female birds.
255 Protective colouring of the Waxwing.
VIII.—Creation by Law.
293 Amount of variation in dogs.
296, 297. The "Times" on Natural Selection.
298 to 300. On intermediate or generalized forms of extinct animals as an indication of transmutation or development.
302 Tabular demonstration of the Origin of Species by Natural Selection.
IX.—The development of Human Races, under the law of Natural Selection.
316 On colour as perhaps correlated with immunity from disease in man.
326, 327. On the probable future development of man.
330 Concluding paragraph rewritten.
London, March, 1870.
PREFACE TO THE SECOND EDITION.
The flattering reception of my Essays by the public and the press having led to a second edition being called for within a year of its first publication, I have taken the opportunity to make a few necessary corrections. I have also added a few passages to the 6th and 7th Essays, and have given two notes, explanatory of some portions of the last chapter which appear to have been not always understood. These additions are as follows:—
+ -+ To avoid altering the paging the additional pages now given have been lettered. + -+ -+ -+ 1st Ed. 2nd Ed. + -+ -+ 221 221 Additional facts as to birds acquiring the song of other species. 223 223A } Mr. Spruce's remarks on young birds 223B } pairing with old. 228 228A } Pouchet's observations on a change 228B } in the nests of swallows. 229 Passage omitted about nest of Golden Crested Warbler, which had been inserted on Rennie's authority, but has not been confirmed by any later observers. 261 261 Daines Barrington, on importance of protection to the female bird. 372 Note A. 372B Note B. + -+ -+ -+
I.—On the Law which has regulated the introduction of New Species. Pp. 1-25
Geographical distribution dependent on Geologic Changes
A Law deduced from well-known Geographical and Geological facts
The form of a true system of Classification determined by this Law
Geographical Distribution of Organisms
Geological Distribution of the forms of Life
High Organization of very ancient Animals consistent with this Law
Objections to Forbes' Theory of Polarity
II.—On the Tendency of Varieties to depart indefinitely from the Original Type. Pp. 26-44
Instability of Varieties supposed to prove the permanent distinctness of Species
The Struggle for Existence
The Law of Population of Species
The Abundance or Rarity of a Species dependent upon its more or less perfect Adaptation to the Conditions of Existence
Useful Variations will tend to Increase, useless or hurtful Variations to Diminish
Superior Varieties will ultimately extirpate the Original Species
The Partial Reversion of Domesticated Varieties explained
Lamarck's Hypothesis very different from that now advanced
III.—Mimicry, and other Protective Resemblances among Animals. Pp. 45-129
Test of true and false Theories
Importance of the Principle of Utility
Popular Theories of Colour in Animals
Importance of Concealment as influencing Colour
Special modifications of Colour
Theory of Protective Colouring
Objection that Colour as being dangerous should not exist in Nature
Mimicry among Lepidoptera
Lepidoptera mimicking other Insects
Mimicry among Beetles
Beetles mimicking other Insects
Insects mimicking Species of other Orders
Cases of Mimicry among the Vertebrata
Mimicry among Snakes
Mimicry among Birds
Mimicry among Mammals
Objections to Mr. Bates' Theory of Mimicry
Mimicry by Female Insects only
Cause of the dull Colours of Female Birds
Use of the gaudy Colours of many Caterpillars
General deductions as to Colour in Nature
IV.—The Malayan Papilionidae, or Swallow-tailed Butterflies, as illustrative of the Theory of Natural Selection. Pp. 130-200
Special value of the Diurnal Lepidoptera for inquiries of this Nature
Question of the rank of the Papilionidae
Distribution of the Papilionidae
Definition of the word Species
Laws and Modes of Variation
Polymorphism or Dimorphism
Local form or variety
Race or Subspecies
Variation as specially influenced by Locality
Local Variation of Size
Local Variation of Form
Local Variations of Colour
Remarks on the facts of Local Variation
Concluding Remarks on Variation in Lepidoptera
Arrangement and Geographical Distribution of the Malayan Papilionidae
Range of the Groups of Malayan Papilionidae
Remarkable peculiarities of the island of Celebes
V.—On Instinct in Man and Animals. Pp. 201-210
How Instinct may be best Studied
Definition of Instinct
Does Man possess Instincts?
How Indians travel through unknown and trackless Forests
VI.—The Philosophy of Birds' Nests. Pp. 211-230
Instinct or Reason in the Construction of Birds' Nests
Do Men build by Reason or by Imitation?
Why does each Bird build a peculiar kind of Nest?
How do young Birds learn to build their first Nest?
Do Birds sing by Instinct or by Imitation?
Man's Works mainly Imitative
How young Birds may learn to build Nests.
Birds do Alter and Improve their Nests when altered conditions require it
VII.—A Theory of Birds' Nests; showing the relation of certain differences of colour in female birds to their mode of nidification. Pp. 231-263
Changed Conditions and persistent Habits as influencing Nidification
Classification of Nests
Sexual differences of Colour in Birds
The Law which connects the Colours of Female Birds with the mode of Nidification
What the Facts Teach us Colour more variable than Structure or Habits, and therefore the Character which has generally been modified
Exceptional cases confirmatory of the above Explanation
Real or apparent exceptions to the Law stated at p. 240
Various modes of Protection of Animals
Females of some groups require and obtain more Protection than the Males
VIII.—Creation by Law. Pp. 264-301
Laws from which the Origin of Species may be deduced
Mr. Darwin's Metaphors liable to Misconception
A case of Orchis-structure explained by Natural Selection
Adaptation brought about by General Laws
Beauty in Nature
How new Forms are produced by Variation and Selection
The Objection that there are Limits to Variation
Objection to the argument from Classification
The Times on Natural Selection
Intermediate or generalized forms of Extinct Animals an indication of Transmutation or Development
A Demonstration of the Origin of Species
IX.—The Development of Human Races under the Law of Natural Selection. Pp. 302-331
Wide difference of Opinion as to Man's Origin
Outline of the Theory of Natural Selection
Different effects of Natural Selection on Animals and on Man
Influence of External Nature in the development of the Human Mind
Extinction of Lower Races
The Origin of the Races of Man
The Bearing of these views on the Antiquity of Man
Their Bearing on the Dignity and Supremacy of Man
Their Bearing on the future Development of Man
X.—The Limits of Natural Selection as applied to Man. Pp. 333—371
What Natural Selection can Not do
The Brain of the Savage shown to be Larger than he Needs it to be
Size of Brain an important Element of Mental Power
Comparison of the Brains of Man and of Anthropoid Apes
Range of intellectual power in Man
Intellect of Savages and of Animals compared
The use of the Hairy Covering of Mammalia
The Constant absence of Hair from certain parts of Man's body a remarkable Phenomenon
Savage Man feels the want of this Hairy Covering
Man's Naked Skin could not have been produced by Natural Selection
Feet and Hands of Man considered as Difficulties on the Theory of Natural Selection
The Origin of Some of Man's Mental Faculties, by the preservation of Useful Variations, not possible
Difficulty as to the Origin of the Moral Sense
Summary of the Argument as to the Insufficiency of Natural Selection to account for the Development of Man
The Origin of Consciousness
The Nature of Matter
Matter is Force
All Force is probably Will-force
ON THE LAW WHICH HAS REGULATED THE INTRODUCTION OF NEW SPECIES.[A]
[A] Written at Sarawak in February, 1855, and published in the "Annals and Magazine of Natural History," September, 1855.
Geographical Distribution dependent on Geologic Changes.
Every naturalist who has directed his attention to the subject of the geographical distribution of animals and plants, must have been interested in the singular facts which it presents. Many of these facts are quite different from what would have been anticipated, and have hitherto been considered as highly curious, but quite inexplicable. None of the explanations attempted from the time of Linnaeus are now considered at all satisfactory; none of them have given a cause sufficient to account for the facts known at the time, or comprehensive enough to include all the new facts which have since been, and are daily being added. Of late years, however, a great light has been thrown upon the subject by geological investigations, which have shown that the present state of the earth and of the organisms now inhabiting it, is but the last stage of a long and uninterrupted series of changes which it has undergone, and consequently, that to endeavour to explain and account for its present condition without any reference to those changes (as has frequently been done) must lead to very imperfect and erroneous conclusions.
The facts proved by geology are briefly these:—That during an immense, but unknown period, the surface of the earth has undergone successive changes; land has sunk beneath the ocean, while fresh land has risen up from it; mountain chains have been elevated; islands have been formed into continents, and continents submerged till they have become islands; and these changes have taken place, not once merely, but perhaps hundreds, perhaps thousands of times:—That all these operations have been more or less continuous, but unequal in their progress, and during the whole series the organic life of the earth has undergone a corresponding alteration. This alteration also has been gradual, but complete; after a certain interval not a single species existing which had lived at the commencement of the period. This complete renewal of the forms of life also appears to have occurred several times:—That from the last of the geological epochs to the present or historical epoch, the change of organic life has been gradual: the first appearance of animals now existing can in many cases be traced, their numbers gradually increasing in the more recent formations, while other species continually die out and disappear, so that the present condition of the organic world is clearly derived by a natural process of gradual extinction and creation of species from that of the latest geological periods. We may therefore safely infer a like gradation and natural sequence from one geological epoch to another.
Now, taking this as a fair statement of the results of geological inquiry, we see that the present geographical distribution of life upon the earth must be the result of all the previous changes, both of the surface of the earth itself and of its inhabitants. Many causes, no doubt, have operated of which we must ever remain in ignorance, and we may, therefore, expect to find many details very difficult of explanation, and in attempting to give one, must allow ourselves to call into our service geological changes which it is highly probable may have occurred, though we have no direct evidence of their individual operation.
The great increase of our knowledge within the last twenty years, both of the present and past history of the organic world, has accumulated a body of facts which should afford a sufficient foundation for a comprehensive law embracing and explaining them all, and giving a direction to new researches. It is about ten years since the idea of such a law suggested itself to the writer of this essay, and he has since taken every opportunity of testing it by all the newly-ascertained facts with which he has become acquainted, or has been able to observe himself. These have all served to convince him of the correctness of his hypothesis. Fully to enter into such a subject would occupy much space, and it is only in consequence of some views having been lately promulgated, he believes, in a wrong direction, that he now ventures to present his ideas to the public, with only such obvious illustrations of the arguments and results as occur to him in a place far removed from all means of reference and exact information.
A Law deduced from well-known Geographical and Geological Facts.
The following propositions in Organic Geography and Geology give the main facts on which the hypothesis is founded.
1. Large groups, such as classes and orders, are generally spread over the whole earth, while smaller ones, such as families and genera, are frequently confined to one portion, often to a very limited district.
2. In widely distributed families the genera are often limited in range; in widely distributed genera, well marked groups of species are peculiar to each geographical district.
3. When a group is confined to one district, and is rich in species, it is almost invariably the case that the most closely allied species are found in the same locality or in closely adjoining localities, and that therefore the natural sequence of the species by affinity is also geographical.
4. In countries of a similar climate, but separated by a wide sea or lofty mountains, the families, genera and species of the one are often represented by closely allied families, genera and species peculiar to the other.
5. The distribution of the organic world in time is very similar to its present distribution in space.
6. Most of the larger and some small groups extend through several geological periods.
7. In each period, however, there are peculiar groups, found nowhere else, and extending through one or several formations.
8. Species of one genus, or genera of one family occurring in the same geological time are more closely allied than those separated in time.
9. As generally in geography no species or genus occurs in two very distant localities without being also found in intermediate places, so in geology the life of a species or genus has not been interrupted. In other words, no group or species has come into existence twice.
10. The following law may be deduced from these facts:—Every species has come into existence coincident both in space and time with a pre-existing closely allied species.
This law agrees with, explains and illustrates all the facts connected with the following branches of the subject:—1st. The system of natural affinities. 2nd. The distribution of animals and plants in space. 3rd. The same in time, including all the phaenomena of representative groups, and those which Professor Forbes supposed to manifest polarity. 4th. The phaenomena of rudimentary organs. We will briefly endeavour to show its bearing upon each of these.
The Form of a true system of Classification determined by this Law.
If the law above enunciated be true, it follows that the natural series of affinities will also represent the order in which the several species came into existence, each one having had for its immediate antitype a closely allied species existing at the time of its origin. It is evidently possible that two or three distinct species may have had a common antitype, and that each of these may again have become the antitypes from which other closely allied species were created. The effect of this would be, that so long as each species has had but one new species formed on its model, the line of affinities will be simple, and may be represented by placing the several species in direct succession in a straight line. But if two or more species have been independently formed on the plan of a common antitype, then the series of affinities will be compound, and can only be represented by a forked or many branched line. Now, all attempts at a Natural classification and arrangement of organic beings show, that both these plans have obtained in creation. Sometimes the series of affinities can be well represented for a space by a direct progression from species to species or from group to group, but it is generally found impossible so to continue. There constantly occur two or more modifications of an organ or modifications of two distinct organs, leading us on to two distinct series of species, which at length differ so much from each other as to form distinct genera or families. These are the parallel series or representative groups of naturalists, and they often occur in different countries, or are found fossil in different formations. They are said to have an analogy to each other when they are so far removed from their common antitype as to differ in many important points of structure, while they still preserve a family resemblance. We thus see how difficult it is to determine in every case whether a given relation is an analogy or an affinity, for it is evident that as we go back along the parallel or divergent series, towards the common antitype, the analogy which existed between the two groups becomes an affinity. We are also made aware of the difficulty of arriving at a true classification, even in a small and perfect group;—in the actual state of nature it is almost impossible, the species being so numerous and the modifications of form and structure so varied, arising probably from the immense number of species which have served as antitypes for the existing species, and thus produced a complicated branching of the lines of affinity, as intricate as the twigs of a gnarled oak or the vascular system of the human body. Again, if we consider that we have only fragments of this vast system, the stem and main branches being represented by extinct species of which we have no knowledge, while a vast mass of limbs and boughs and minute twigs and scattered leaves is what we have to place in order, and determine the true position each originally occupied with regard to the others, the whole difficulty of the true Natural System of classification becomes apparent to us.
We shall thus find ourselves obliged to reject all these systems of classification which arrange species or groups in circles, as well as these which fix a definite number for the divisions of each group. The latter class have been very generally rejected by naturalists, as contrary to nature, notwithstanding the ability with which they have been advocated; but the circular system of affinities seems to have obtained a deeper hold, many eminent naturalists having to some extent adopted it. We have, however, never been able to find a case in which the circle has been closed by a direct and close affinity. In most cases a palpable analogy has been substituted, in others the affinity is very obscure or altogether doubtful. The complicated branching of the lines of affinities in extensive groups must also afford great facilities for giving a show of probability to any such purely artificial arrangements. Their death-blow was given by the admirable paper of the lamented Mr. Strickland, published in the "Annals of Natural History," in which he so clearly showed the true synthetical method of discovering the Natural System.
Geographical Distribution of Organisms.
If we now consider the geographical distribution of animals and plants upon the earth, we shall find all the facts beautifully in accordance with, and readily explained by, the present hypothesis. A country having species, genera, and whole families peculiar to it, will be the necessary result of its having been isolated for a long period, sufficient for many series of species to have been created on the type of pre-existing ones, which, as well as many of the earlier-formed species, have become extinct, and thus made the groups appear isolated. If in any case the antitype had an extensive range, two or more groups of species might have been formed, each varying from it in a different manner, and thus producing several representative or analogous groups. The Sylviadae of Europe and the Sylvicolidae of North America, the Heliconidae of South America and the Euploeas of the East, the group of Trogons inhabiting Asia, and that peculiar to South America, are examples that may be accounted for in this manner.
Such phaenomena as are exhibited by the Galapagos Islands, which contain little groups of plants and animals peculiar to themselves, but most nearly allied to those of South America, have not hitherto received any, even a conjectural explanation. The Galapagos are a volcanic group of high antiquity, and have probably never been more closely connected with the continent than they are at present. They must have been first peopled, like other newly-formed islands, by the action of winds and currents, and at a period sufficiently remote to have had the original species die out, and the modified prototypes only remain. In the same way we can account for the separate islands having each their peculiar species, either on the supposition that the same original emigration peopled the whole of the islands with the same species from which differently modified prototypes were created, or that the islands were successively peopled from each other, but that new species have been created in each on the plan of the pre-existing ones. St. Helena is a similar case of a very ancient island having obtained an entirely peculiar, though limited, flora. On the other hand, no example is known of an island which can be proved geologically to be of very recent origin (late in the Tertiary, for instance), and yet possesses generic or family groups, or even many species peculiar to itself.
When a range of mountains has attained a great elevation, and has so remained during a long geological period, the species of the two sides at and near their bases will be often very different, representative species of some genera occurring, and even whole genera being peculiar to one side only, as is remarkably seen in the case of the Andes and Rocky Mountains. A similar phaenomenon occurs when an island has been separated from a continent at a very early period. The shallow sea between the Peninsula of Malacca, Java, Sumatra and Borneo was probably a continent or large island at an early epoch, and may have become submerged as the volcanic ranges of Java and Sumatra were elevated. The organic results we see in the very considerable number of species of animals common to some or all of these countries, while at the same time a number of closely allied representative species exist peculiar to each, showing that a considerable period has elapsed since their separation. The facts of geographical distribution and of geology may thus mutually explain each other in doubtful cases, should the principles here advocated be clearly established.
In all those cases in which an island has been separated from a continent, or raised by volcanic or coralline action from the sea, or in which a mountain-chain has been elevated in a recent geological epoch, the phaenomena of peculiar groups or even of single representative species will not exist. Our own island is an example of this, its separation from the continent being geologically very recent, and we have consequently scarcely a species which is peculiar to it; while the Alpine range, one of the most recent mountain elevations, separates faunas and floras which scarcely differ more than may be due to climate and latitude alone.
The series of facts alluded to in Proposition (3), of closely allied species in rich groups being found geographically near each other, is most striking and important. Mr. Lovell Reeve has well exemplified it in his able and interesting paper on the Distribution of the Bulimi. It is also seen in the Humming-birds and Toucans, little groups of two or three closely allied species being often found in the same or closely adjoining districts, as we have had the good fortune of personally verifying. Fishes give evidence of a similar kind: each great river has its peculiar genera, and in more extensive genera its groups of closely allied species. But it is the same throughout Nature; every class and order of animals will contribute similar facts. Hitherto no attempt has been made to explain these singular phenomena, or to show how they have arisen. Why are the genera of Palms and of Orchids in almost every case confined to one hemisphere? Why are the closely allied species of brown-backed Trogons all found in the East, and the green-backed in the West? Why are the Macaws and the Cockatoos similarly restricted? Insects furnish a countless number of analogous examples;—the Goliathi of Africa, the Ornithopterae of the Indian Islands, the Heliconidae of South America, the Danaidae of the East, and in all, the most closely allied species found in geographical proximity. The question forces itself upon every thinking mind,—why are these things so? They could not be as they are had no law regulated their creation and dispersion. The law here enunciated not merely explains, but necessitates the facts we see to exist, while the vast and long-continued geological changes of the earth readily account for the exceptions and apparent discrepancies that here and there occur. The writer's object in putting forward his views in the present imperfect manner is to submit them to the test of other minds, and to be made aware of all the facts supposed to be inconsistent with them. As his hypothesis is one which claims acceptance solely as explaining and connecting facts which exist in nature, he expects facts alone to be brought to disprove it, not a priori arguments against its probability.
Geological Distribution of the Forms of Life.
The phaenomena of geological distribution are exactly analogous to those of geography. Closely allied species are found associated in the same beds, and the change from species to species appears to have been as gradual in time as in space. Geology, however, furnishes us with positive proof of the extinction and production of species, though it does not inform us how either has taken place. The extinction of species, however, offers but little difficulty, and the modus operandi has been well illustrated by Sir C. Lyell in his admirable "Principles." Geological changes, however gradual, must occasionally have modified external conditions to such an extent as to have rendered the existence of certain species impossible. The extinction would in most cases be effected by a gradual dying-out, but in some instances there might have been a sudden destruction of a species of limited range. To discover how the extinct species have from time to time been replaced by new ones down to the very latest geological period, is the most difficult, and at the same time the most interesting problem in the natural history of the earth. The present inquiry, which seeks to eliminate from known facts a law which has determined, to a certain degree, what species could and did appear at a given epoch, may, it is hoped, be considered as one step in the right direction towards a complete solution of it.
High Organization of very ancient Animals consistent with this Law.
Much discussion has of late years taken place on the question, whether the succession of life upon the globe has been from a lower to a higher degree of organization. The admitted facts seem to show that there has been a general, but not a detailed progression. Mollusca and Radiata existed before Vertebrata, and the progression from Fishes to Reptiles and Mammalia, and also from the lower mammals to the higher, is indisputable. On the other hand, it is said that the Mollusca and Radiata of the very earliest periods were more highly organized than the great mass of those now existing, and that the very first fishes that have been discovered are by no means the lowest organised of the class. Now it is believed the present hypothesis will harmonize with all these facts, and in a great measure serve to explain them; for though it may appear to some readers essentially a theory of progression, it is in reality only one of gradual change. It is, however, by no means difficult to show that a real progression in the scale of organization is perfectly consistent with all the appearances, and even with apparent retrogression, should such occur.
Returning to the analogy of a branching tree, as the best mode of representing the natural arrangement of species and their successive creation, let us suppose that at an early geological epoch any group (say a class of the Mollusca) has attained to a great richness of species and a high organization. Now let this great branch of allied species, by geological mutations, be completely or partially destroyed. Subsequently a new branch springs from the same trunk, that is to say, new species are successively created, having for their antitypes the same lower organized species which had served as the antitypes for the former group, but which have survived the modified conditions which destroyed it. This new group being subject to these altered conditions, has modifications of structure and organization given to it, and becomes the representative group of the former one in another geological formation. It may, however, happen, that though later in time, the new series of species may never attain to so high a degree of organization as those preceding it, but in its turn become extinct, and give place to yet another modification from the same root, which may be of higher or lower organization, more or less numerous in species, and more or less varied in form and structure than either of those which preceded it. Again, each of these groups may not have become totally extinct, but may have left a few species, the modified prototypes of which have existed in each succeeding period, a faint memorial of their former grandeur and luxuriance. Thus every case of apparent retrogression may be in reality a progress, though an interrupted one: when some monarch of the forest loses a limb, it may be replaced by a feeble and sickly substitute. The foregoing remarks appear to apply to the case of the Mollusca, which, at a very early period, had reached a high organization and a great development of forms and species in the testaceous Cephalopoda. In each succeeding age modified species and genera replaced the former ones which had become extinct, and as we approach the present aera, but few and small representatives of the group remain, while the Gasteropods and Bivalves have acquired an immense preponderance. In the long series of changes the earth has undergone, the process of peopling it with organic beings has been continually going on, and whenever any of the higher groups have become nearly or quite extinct, the lower forms which have better resisted the modified physical conditions have served as the antitypes on which to found the new races. In this manner alone, it is believed, can the representative groups at successive periods, and the risings and fallings in the scale of organization, be in every case explained.
Objections to Forbes' Theory of Polarity.
The hypothesis of polarity, recently put forward by Professor Edward Forbes to account for the abundance of generic forms at a very early period and at present, while in the intermediate epochs there is a gradual diminution and impoverishment, till the minimum occurred at the confines of the Palaeozoic and Secondary epochs, appears to us quite unnecessary, as the facts may be readily accounted for on the principles already laid down. Between the Palaeozoic and Neozoic periods of Professor Forbes, there is scarcely a species in common, and the greater part of the genera and families also disappear to be replaced by new ones. It is almost universally admitted that such a change in the organic world must have occupied a vast period of time. Of this interval we have no record; probably because the whole area of the early formations now exposed to our researches was elevated at the end of the Palaeozoic period, and remained so through the interval required for the organic changes which resulted in the fauna and flora of the Secondary period. The records of this interval are buried beneath the ocean which covers three-fourths of the globe. Now it appears highly probable that a long period of quiescence or stability in the physical conditions of a district would be most favourable to the existence of organic life in the greatest abundance, both as regards individuals and also as to variety of species and generic group, just as we now find that the places best adapted to the rapid growth and increase of individuals also contain the greatest profusion of species and the greatest variety of forms,—the tropics in comparison with the temperate and arctic regions. On the other hand, it seems no less probable that a change in the physical conditions of a district, even small in amount if rapid, or even gradual if to a great amount, would be highly unfavourable to the existence of individuals, might cause the extinction of many species, and would probably be equally unfavourable to the creation of new ones. In this too we may find an analogy with the present state of our earth, for it has been shown to be the violent extremes and rapid changes of physical conditions, rather than the actual mean state in the temperate and frigid zones, which renders them less prolific than the tropical regions, as exemplified by the great distance beyond the tropics to which tropical forms penetrate when the climate is equable, and also by the richness in species and forms of tropical mountain regions which principally differ from the temperate zone in the uniformity of their climate. However this may be, it seems a fair assumption that during a period of geological repose the new species which we know to have been created would have appeared; that the creations would then exceed in number the extinctions, and therefore the number of species would increase. In a period of geological activity, on the other hand, it seems probable that the extinctions might exceed the creations, and the number of species consequently diminish. That such effects did take place in connexion with the causes to which we have imputed them, is shown in the case of the Coal formation, the faults and contortions of which show a period of great activity and violent convulsions, and it is in the formation immediately succeeding this that the poverty of forms of life is most apparent. We have then only to suppose a long period of somewhat similar action during the vast unknown interval at the termination of the Palaeozoic period, and then a decreasing violence or rapidity through the Secondary period, to allow for the gradual repopulation of the earth with varied forms, and the whole of the facts are explained.[B] We thus have a clue to the increase of the forms of life during certain periods, and their decrease during others, without recourse to any causes but those we know to have existed, and to effects fairly deducible from them. The precise manner in which the geological changes of the early formations were effected is so extremely obscure, that when we can explain important facts by a retardation at one time and an acceleration at another of a process which we know from its nature and from observation to have been unequal,—a cause so simple may surely be preferred to one so obscure and hypothetical as polarity.
[B] Professor Ramsay has since shown that a glacial epoch probably occurred at the time of the Permian formation, which will more satisfactorily account for the comparative poverty of species.
I would also venture to suggest some reasons against the very nature of the theory of Professor Forbes. Our knowledge of the organic world during any geological epoch is necessarily very imperfect. Looking at the vast numbers of species and groups that have been discovered by geologists, this may be doubted; but we should compare their numbers not merely with those that now exist upon the earth, but with a far larger amount. We have no reason for believing that the number of species on the earth at any former period was much less than at present; at all events the aquatic portion, with which geologists have most acquaintance, was probably often as great or greater. Now we know that there have been many complete changes of species; new sets of organisms have many times been introduced in place of old ones which have become extinct, so that the total amount which have existed on the earth from the earliest geological period must have borne about the same proportion to those now living, as the whole human race who have lived and died upon the earth, to the population at the present time. Again, at each epoch, the whole earth was no doubt, as now, more or less the theatre of life, and as the successive generations of each species died, their exuviae and preservable parts would be deposited over every portion of the then existing seas and oceans, which we have reason for supposing to have been more, rather than less, extensive than at present. In order then to understand our possible knowledge of the early world and its inhabitants, we must compare, not the area of the whole field of our geological researches with the earth's surface, but the area of the examined portion of each formation separately with the whole earth. For example, during the Silurian period all the earth was Silurian, and animals were living and dying, and depositing their remains more or less over the whole area of the globe, and they were probably (the species at least) nearly as varied in different latitudes and longitudes as at present. What proportion do the Silurian districts bear to the whole surface of the globe, land and sea (for far more extensive Silurian districts probably exist beneath the ocean than above it), and what portion of the known Silurian districts has been actually examined for fossils? Would the area of rock actually laid open to the eye be the thousandth or the ten-thousandth part of the earth's surface? Ask the same question with regard to the Oolite or the Chalk, or even to particular beds of these when they differ considerably in their fossils, and you may then get some notion of how small a portion of the whole we know.
But yet more important is the probability, nay almost the certainty, that whole formations containing the records of vast geological periods are entirely buried beneath the ocean, and for ever beyond our reach. Most of the gaps in the geological series may thus be filled up, and vast numbers of unknown and unimaginable animals, which might help to elucidate the affinities of the numerous isolated groups which are a perpetual puzzle to the zoologist, may there be buried, till future revolutions may raise them in their turn above the waters, to afford materials for the study of whatever race of intelligent beings may then have succeeded us. These considerations must lead us to the conclusion, that our knowledge of the whole series of the former inhabitants of the earth is necessarily most imperfect and fragmentary,—as much so as our knowledge of the present organic world would be, were we forced to make our collections and observations only in spots equally limited in area and in number with those actually laid open for the collection of fossils. Now, the hypothesis of Professor Forbes is essentially one that assumes to a great extent the completeness of our knowledge of the whole series of organic beings which have existed on the earth. This appears to be a fatal objection to it, independently of all other considerations. It may be said that the same objections exist against every theory on such a subject, but this is not necessarily the case. The hypothesis put forward in this paper depends in no degree upon the completeness of our knowledge of the former condition of the organic world, but takes what facts we have as fragments of a vast whole, and deduces from them something of the nature and proportions of that whole which we can never know in detail. It is founded upon isolated groups of facts, recognizes their isolation, and endeavours to deduce from them the nature of the intervening portions.
Another important series of facts, quite in accordance with, and even necessary deductions from, the law now developed, are those of rudimentary organs. That these really do exist, and in most cases have no special function in the animal oeconomy, is admitted by the first authorities in comparative anatomy. The minute limbs hidden beneath the skin in many of the snake-like lizards, the anal hooks of the boa constrictor, the complete series of jointed finger-bones in the paddle of the Manatus and whale, are a few of the most familiar instances. In botany a similar class of facts has been long recognised. Abortive stamens, rudimentary floral envelopes and undeveloped carpels, are of the most frequent occurrence. To every thoughtful naturalist the question must arise, What are these for? What have they to do with the great laws of creation? Do they not teach us something of the system of Nature? If each species has been created independently, and without any necessary relations with pre-existing species, what do these rudiments, these apparent imperfections mean? There must be a cause for them; they must be the necessary results of some great natural law. Now, if, as it has been endeavoured to be shown, the great law which has regulated the peopling of the earth with animal and vegetable life is, that every change shall be gradual; that no new creature shall be formed widely differing from anything before existing; that in this, as in everything else in Nature, there shall be gradation and harmony,—then these rudimentary organs are necessary, and are an essential part of the system of Nature. Ere the higher Vertebrata were formed, for instance, many steps were required, and many organs had to undergo modifications from the rudimental condition in which only they had as yet existed. We still see remaining an antitypal sketch of a wing adapted for flight in the scaly flapper of the penguin, and limbs first concealed beneath the skin, and then weakly protruding from it, were the necessary gradations before others should be formed fully adapted for locomotion.[C] Many more of these modifications should we behold, and more complete series of them, had we a view of all the forms which have ceased to live. The great gaps that exist between fishes, reptiles, birds, and mammals would then, no doubt, be softened down by intermediate groups, and the whole organic world would be seen to be an unbroken and harmonious system.
[C] The theory of Natural Selection has now taught us that these are not the steps by which limbs have been formed; and that most rudimentary organs have been produced by abortion, owing to disuse, as explained by Mr. Darwin.
It has now been shown, though most briefly and imperfectly, how the law that "Every species has come into existence coincident both in time and space with a pre-existing closely allied species," connects together and renders intelligible a vast number of independent and hitherto unexplained facts. The natural system of arrangement of organic beings, their geographical distribution, their geological sequence, the phaenomena of representative and substituted groups in all their modifications, and the most singular peculiarities of anatomical structure, are all explained and illustrated by it, in perfect accordance with the vast mass of facts which the researches of modern naturalists have brought together, and, it is believed, not materially opposed to any of them. It also claims a superiority over previous hypotheses, on the ground that it not merely explains, but necessitates what exists. Granted the law, and many of the most important facts in Nature could not have been otherwise, but are almost as necessary deductions from it, as are the elliptic orbits of the planets from the law of gravitation.
ON THE TENDENCY OF VARIETIES TO DEPART INDEFINITELY FROM THE ORIGINAL TYPE.[D]
[D] Written at Ternate, February, 1858; and published in the Journal of the Proceedings of the Linnaean Society for August, 1858.
Instability of Varieties supposed to prove the permanent distinctness of Species.
One of the strongest arguments which have been adduced to prove the original and permanent distinctness of species is, that varieties produced in a state of domesticity are more or less unstable, and often have a tendency, if left to themselves, to return to the normal form of the parent species; and this instability is considered to be a distinctive peculiarity of all varieties, even of those occurring among wild animals in a state of nature, and to constitute a provision for preserving unchanged the originally created distinct species.
In the absence or scarcity of facts and observations as to varieties occurring among wild animals, this argument has had great weight with naturalists, and has led to a very general and somewhat prejudiced belief in the stability of species. Equally general, however, is the belief in what are called "permanent or true varieties,"—races of animals which continually propagate their like, but which differ so slightly (although constantly) from some other race, that the one is considered to be a variety of the other. Which is the variety and which the original species, there is generally no means of determining, except in those rare cases in which the one race has been known to produce an offspring unlike itself and resembling the other. This, however, would seem quite incompatible with the "permanent invariability of species," but the difficulty is overcome by assuming that such varieties have strict limits, and can never again vary further from the original type, although they may return to it, which, from the analogy of the domesticated animals, is considered to be highly probable, if not certainly proved.
It will be observed that this argument rests entirely on the assumption, that varieties occurring in a state of nature are in all respects analogous to or even identical with those of domestic animals, and are governed by the same laws as regards their permanence or further variation. But it is the object of the present paper to show that this assumption is altogether false, that there is a general principle in nature which will cause many varieties to survive the parent species, and to give rise to successive variations departing further and further from the original type; and which also produces, in domesticated animals, the tendency of varieties to return to the parent form.
The Struggle for Existence.
The life of wild animals is a struggle for existence. The full exertion of all their faculties and all their energies is required to preserve their own existence and provide for that of their infant offspring. The possibility of procuring food during the least favourable seasons, and of escaping the attacks of their most dangerous enemies, are the primary conditions which determine the existence both of individuals and of entire species. These conditions will also determine the population of a species; and by a careful consideration of all the circumstances we may be enabled to comprehend, and in some degree to explain, what at first sight appears so inexplicable—the excessive abundance of some species, while others closely allied to them are very rare.
The Law of Population of Species.
The general proportion that must obtain between certain groups of animals is readily seen. Large animals cannot be so abundant as small ones; the carnivora must be less numerous than the herbivora; eagles and lions can never be so plentiful as pigeons and antelopes; and the wild asses of the Tartarian deserts cannot equal in numbers the horses of the more luxuriant prairies and pampas of America. The greater or less fecundity of an animal is often considered to be one of the chief causes of its abundance or scarcity; but a consideration of the facts will show us that it really has little or nothing to do with the matter. Even the least prolific of animals would increase rapidly if unchecked, whereas it is evident that the animal population of the globe must be stationary, or perhaps, through the influence of man, decreasing. Fluctuations there may be; but permanent increase, except in restricted localities, is almost impossible. For example, our own observation must convince us that birds do not go on increasing every year in a geometrical ratio, as they would do, were there not some powerful check to their natural increase. Very few birds produce less than two young ones each year, while many have six, eight, or ten; four will certainly be below the average; and if we suppose that each pair produce young only four times in their life, that will also be below the average, supposing them not to die either by violence or want of food. Yet at this rate how tremendous would be the increase in a few years from a single pair! A simple calculation will show that in fifteen years each pair of birds would have increased to nearly ten millions![E] whereas we have no reason to believe that the number of the birds of any country increases at all in fifteen or in one hundred and fifty years. With such powers of increase the population must have reached its limits, and have become stationary, in a very few years after the origin of each species. It is evident, therefore, that each year an immense number of birds must perish—as many in fact as are born; and as on the lowest calculation the progeny are each year twice as numerous as their parents, it follows that, whatever be the average number of individuals existing in any given country, twice that number must perish annually,—a striking result, but one which seems at least highly probable, and is perhaps under rather than over the truth. It would therefore appear that, as far as the continuance of the species and the keeping up the average number of individuals are concerned, large broods are superfluous. On the average all above one become food for hawks and kites, wild cats or weasels, or perish of cold and hunger as winter comes on. This is strikingly proved by the case of particular species; for we find that their abundance in individuals bears no relation whatever to their fertility in producing offspring.
[E] This is under estimated. The number would really amount to more than two thousand millions!
Perhaps the most remarkable instance of an immense bird population is that of the passenger pigeon of the United States, which lays only one, or at most two eggs, and is said to rear generally but one young one. Why is this bird so extraordinarily abundant, while others producing two or three times as many young are much less plentiful? The explanation is not difficult. The food most congenial to this species, and on which it thrives best, is abundantly distributed over a very extensive region, offering such differences of soil and climate, that in one part or another of the area the supply never fails. The bird is capable of a very rapid and long-continued flight, so that it can pass without fatigue over the whole of the district it inhabits, and as soon as the supply of food begins to fail in one place is able to discover a fresh feeding-ground. This example strikingly shows us that the procuring a constant supply of wholesome food is almost the sole condition requisite for ensuring the rapid increase of a given species, since neither the limited fecundity, nor the unrestrained attacks of birds of prey and of man are here sufficient to check it. In no other birds are these peculiar circumstances so strikingly combined. Either their food is more liable to failure, or they have not sufficient power of wing to search for it over an extensive area, or during some season of the year it becomes very scarce, and less wholesome substitutes have to be found; and thus, though more fertile in offspring, they can never increase beyond the supply of food in the least favourable seasons.
Many birds can only exist by migrating, when their food becomes scarce, to regions possessing a milder, or at least a different climate, though, as these migrating birds are seldom excessively abundant, it is evident that the countries they visit are still deficient in a constant and abundant supply of wholesome food. Those whose organization does not permit them to migrate when their food becomes periodically scarce, can never attain a large population. This is probably the reasons why woodpeckers are scarce with us, while in the tropics they are among the most abundant of solitary birds. Thus the house sparrow is more abundant than the redbreast, because its food is more constant and plentiful,—seeds of grasses being preserved during the winter, and our farm-yards and stubble-fields furnishing an almost inexhaustible supply. Why, as a general rule, are aquatic, and especially sea birds, very numerous in individuals? Not because they are more prolific than others, generally the contrary; but because their food never fails, the sea-shores and river-banks daily swarming with a fresh supply of small mollusca and crustacea. Exactly the same laws will apply to mammals. Wild cats are prolific and have few enemies; why then are they never as abundant as rabbits? The only intelligible answer is, that their supply of food is more precarious. It appears evident, therefore, that so long as a country remains physically unchanged, the numbers of its animal population cannot materially increase. If one species does so, some others requiring the same kind of food must diminish in proportion. The numbers that die annually must be immense; and as the individual existence of each animal depends upon itself, those that die must be the weakest—the very young, the aged, and the diseased—while those that prolong their existence can only be the most perfect in health and vigour—those who are best able to obtain food regularly, and avoid their numerous enemies. It is, as we commenced by remarking, "a struggle for existence," in which the weakest and least perfectly organized must always succumb.
The Abundance or Rarity of a Species dependent upon its more or less perfect Adaptation to the Conditions of Existence.
It seems evident that what takes place among the individuals of a species must also occur among the several allied species of a group,—viz., that those which are best adapted to obtain a regular supply of food, and to defend themselves against the attacks of their enemies and the vicissitudes of the seasons, must necessarily obtain and preserve a superiority in population; while those species which from some defect of power or organization are the least capable of counteracting the vicissitudes of food-supply, &c., must diminish in numbers, and, in extreme cases, become altogether extinct. Between these extremes the species will present various degrees of capacity for ensuring the means of preserving life; and it is thus we account for the abundance or rarity of species. Our ignorance will generally prevent us from accurately tracing the effects to their causes; but could we become perfectly acquainted with the organization and habits of the various species of animals, and could we measure the capacity of each for performing the different acts necessary to its safety and existence under all the varying circumstances by which it is surrounded, we might be able even to calculate the proportionate abundance of individuals which is the necessary result.
If now we have succeeded in establishing these two points—1st, that the animal population of a country is generally stationary, being kept down by a periodical deficiency of food, and other checks; and, 2nd, that the comparative abundance or scarcity of the individuals of the several species is entirely due to their organization and resulting habits, which, rendering it more difficult to procure a regular supply of food and to provide for their personal safety in some cases than in others, can only be balanced by a difference in the population which have to exist in a given area—we shall be in a condition to proceed to the consideration of varieties, to which the preceding remarks have a direct and very important application.
Useful Variations will tend to Increase; useless or hurtful Variations to Diminish.
Most or perhaps all the variations from the typical form of a species must have some definite effect, however slight, on the habits or capacities of the individuals. Even a change of colour might, by rendering them more or less distinguishable, affect their safety; a greater or less development of hair might modify their habits. More important changes, such as an increase in the power or dimensions of the limbs or any of the external organs, would more or less affect their mode of procuring food or the range of country which they could inhabit. It is also evident that most changes would affect, either favourably or adversely, the powers of prolonging existence. An antelope with shorter or weaker legs must necessarily suffer more from the attacks of the feline carnivora; the passenger pigeon with less powerful wings would sooner or later be affected in its powers of procuring a regular supply of food; and in both cases the result must necessarily be a diminution of the population of the modified species. If, on the other hand, any species should produce a variety having slightly increased powers of preserving existence, that variety must inevitably in time acquire a superiority in numbers. These results must follow as surely as old age, intemperance, or scarcity of food produce an increased mortality. In both cases there may be many individual exceptions; but on the average the rule will invariably be found to hold good. All varieties will therefore fall into two classes—those which under the same conditions would never reach the population of the parent species, and those which would in time obtain and keep a numerical superiority. Now, let some alteration of physical conditions occur in the district—a long period of drought, a destruction of vegetation by locusts, the irruption of some new carnivorous animal seeking "pastures new"—any change in fact tending to render existence more difficult to the species in question, and tasking its utmost powers to avoid complete extermination; it is evident that, of all the individuals composing the species, those forming the least numerous and most feebly organized variety would suffer first, and, were the pressure severe, must soon become extinct. The same causes continuing in action, the parent species would next suffer, would gradually diminish in numbers, and with a recurrence of similar unfavourable conditions might also become extinct. Tho superior variety would then alone remain, and on a return to favourable circumstances would rapidly increase in numbers and occupy the place of the extinct species and variety.
Superior Varieties will ultimately Extirpate the original Species.
The variety would now have replaced the species, of which it would be a more perfectly developed and more highly organized form. It would be in all respects better adapted to secure its safety, and to prolong its individual existence and that of the race. Such a variety could not return to the original form; for that form is an inferior one, and could never compete with it for existence. Granted, therefore, a "tendency" to reproduce the original type of the species, still the variety must ever remain preponderant in numbers, and under adverse physical conditions again alone survive. But this new, improved, and populous race might itself, in course of time, give rise to new varieties, exhibiting several diverging modifications of form, any of which, tending to increase the facilities for preserving existence, must, by the same general law, in their turn become predominant. Here, then, we have progression and continued divergence deduced from the general laws which regulate the existence of animals in a state of nature, and from the undisputed fact that varieties do frequently occur. It is not, however, contended that this result would be invariable; a change of physical conditions in the district might at times materially modify it, rendering the race which had been the most capable of supporting existence under the former conditions now the least so, and even causing the extinction of the newer and, for a time, superior race, while the old or parent species and its first inferior varieties continued to flourish. Variations in unimportant parts might also occur, having no perceptible effect on the life-preserving powers; and the varieties so furnished might run a course parallel with the parent species, either giving rise to further variations or returning to the former type. All we argue for is, that certain varieties have a tendency to maintain their existence longer than the original species, and this tendency must make itself felt; for though the doctrine of chances or averages can never be trusted to on a limited scale, yet, if applied to high numbers, the results come nearer to what theory demands, and, as we approach to an infinity of examples, become strictly accurate. Now the scale on which nature works is so vast—the numbers of individuals and the periods of time with which she deals approach so near to infinity, than any cause, however slight, and however liable to be veiled and counteracted by accidental circumstances, must in the end produce its full legitimate results.
The Partial Reversion of Domesticated Varieties explained.
Let us now turn to domesticated animals, and inquire how varieties produced among them are affected by the principles here enunciated. The essential difference in the condition of wild and domestic animals is this,—that among the former, their well-being and very existence depend upon the full exercise and healthy condition of all their senses and physical powers, whereas, among the latter, these are only partially exercised, and in some cases are absolutely unused. A wild animal has to search, and often to labour, for every mouthful of food—to exercise sight, hearing, and smell in seeking it, and in avoiding dangers, in procuring shelter from the inclemency of the seasons, and in providing for the subsistence and safety of its offspring. There is no muscle of its body that is not called into daily and hourly activity; there is no sense or faculty that is not strengthened by continual exercise. The domestic animal, on the other hand, has food provided for it, is sheltered, and often confined, to guard it against the vicissitudes of the seasons, is carefully secured from the attacks of its natural enemies, and seldom even rears its young without human assistance. Half of its senses and faculties become quite useless, and the other half are but occasionally called into feeble exercise, while even its muscular system is only irregularly brought into action.
Now when a variety of such an animal occurs, having increased power or capacity in any organ or sense, such increase is totally useless, is never called into action, and may even exist without the animal ever becoming aware of it. In the wild animal, on the contrary, all its faculties and powers being brought into full action for the necessities of existence, any increase becomes immediately available, is strengthened by exercise, and must even slightly modify the food, the habits, and the whole economy of the race. It creates as it were a new animal, one of superior powers, and which will necessarily increase in numbers and outlive those which are inferior to it.
Again, in the domesticated animal all variations have an equal chance of continuance; and those which would decidedly render a wild animal unable to compete with its fellows and continue its existence are no disadvantage whatever in a state of domesticity. Our quickly fattening pigs, short-legged sheep pouter pigeons, and poodle dogs could never have come into existence in a state of nature, because the very first step towards such inferior forms would have led to the rapid extinction of the race; still less could they now exist in competition with their wild allies. The great speed but slight endurance of the race horse, the unwieldy strength of the ploughman's team, would both be useless in a state of nature. If turned wild on the pampas, such animals would probably soon become extinct, or under favourable circumstances might each gradually lose those extreme qualities which would never be called into action, and in a few generations revert to a common type, which must be that in which the various powers and faculties are so proportioned to each other as to be best adapted to procure food and secure safety,—that in which by the full exercise of every part of its organisation the animal can alone continue to live. Domestic varieties, when turned wild, must return to something near the type of the original wild stock, or become altogether extinct.[F]
[F] That is, they will vary, and the variations which tend to adapt them to the wild state, and therefore approximate them to wild animals, will be preserved. Those individuals which do not vary sufficiently will perish.
We see, then, that no inferences as to the permanence of varieties in a state of nature can be deduced from the observations of those occurring among domestic animals. The two are so much opposed to each other in every circumstance of their existence, that what applies to the one is almost sure not to apply to the other. Domestic animals are abnormal, irregular, artificial; they are subject to variations which never occur and never can occur in a state of nature: their very existence depends altogether on human care; so far are many of them removed from that just proportion of faculties, that true balance of organisation, by means of which alone an animal left to its own resources can preserve its existence and continue its race.
Lamarck's Hypothesis very different from that now advanced.
The hypothesis of Lamarck—that progressive changes in species have been produced by the attempts of animals to increase the development of their own organs, and thus modify their structure and habits—has been repeatedly and easily refuted by all writers on the subject of varieties and species, and it seems to have been considered that when this was done the whole question has been finally settled; but the view here developed renders such hypothesis quite unnecessary, by showing that similar results must be produced by the action of principles constantly at work in nature. The powerful retractile talons of the falcon-and the cat-tribes have not been produced or increased by the volition of those animals; but among the different varieties which occurred in the earlier and less highly organized forms of these groups, those always survived longest which had the greatest facilities for seizing their prey. Neither did the giraffe acquire its long neck by desiring to reach the foliage of the more lofty shrubs, and constantly stretching its neck for the purpose, but because any varieties which occurred among its antitypes with a longer neck than usual at once secured a fresh range of pasture over the same ground as their shorter-necked companions, and on the first scarcity of food were thereby enabled to outlive them. Even the peculiar colours of many animals, more especially of insects, so closely resembling the soil or leaves or bark on which they habitually reside, are explained on the same principle; for though in the course of ages varieties of many tints may have occurred, yet those races having colours best adapted to concealment from their enemies would inevitably survive the longest. We have also here an acting cause to account for that balance so often observed in nature,—a deficiency in one set of organs always being compensated by an increased development of some others—powerful wings accompanying weak feet, or great velocity making up for the absence of defensive weapons; for it has been shown that all varieties in which an unbalanced deficiency occurred could not long continue their existence. The action of this principle is exactly like that of the centrifugal governor of the steam engine, which checks and corrects any irregularities almost before they become evident; and in like manner no unbalanced deficiency in the animal kingdom can ever reach any conspicuous magnitude, because it would make itself felt at the very first step, by rendering existence difficult and extinction almost sure soon to follow. An origin such as is here advocated will also agree with the peculiar character of the modifications of form and structure which obtain in organized beings—the many lines of divergence from a central type, the increasing efficiency and power of a particular organ through a succession of allied species, and the remarkable persistence of unimportant parts, such as colour, texture of plumage and hair, form of horns or crests, through a series of species differing considerably in more essential characters. It also furnishes us with a reason for that "more specialized structure" which Professor Owen states to be a characteristic of recent compared with extinct forms, and which would evidently be the result of the progressive modification of any organ applied to a special purpose in the animal economy.
We believe we have now shown that there is a tendency in nature to the continued progression of certain classes of varieties further and further from the original type—a progression to which there appears no reason to assign any definite limits—and that the same principle which produces this result in a state of nature will also explain why domestic varieties have a tendency, when they become wild, to revert to the original type. This progression, by minute steps, in various directions, but always checked and balanced by the necessary conditions, subject to which alone existence can be preserved, may, it is believed, be followed out so as to agree with all the phaenomena presented by organized beings, their extinction and succession in past ages, and all the extraordinary modifications of form, instinct and habits which they exhibit.
MIMICRY, AND OTHER PROTECTIVE RESEMBLANCES AMONG ANIMALS.
There is no more convincing proof of the truth of a comprehensive theory, than its power of absorbing and finding a place for new facts, and its capability of interpreting phaenomena which had been previously looked upon as unaccountable anomalies. It is thus that the law of universal gravitation and the undulatory theory of light have become established and universally accepted by men of science. Fact after fact has been brought forward as being apparently inconsistent with them, and one after another these very facts have been shown to be the consequences of the laws they were at first supposed to disprove. A false theory will never stand this test. Advancing knowledge brings to light whole groups of facts which it cannot deal with, and its advocates steadily decrease in numbers, notwithstanding the ability and scientific skill with which it may have been supported. The great name of Edward Forbes did not prevent his theory of "Polarity in the distribution of Organic beings in Time" from dying a natural death; but the most striking illustration of the behaviour of a false theory is to be found in the "Circular and Quinarian System" of classification propounded by MacLeay, and developed by Swainson, with an amount of knowledge and ingenuity that have rarely been surpassed. This theory was eminently attractive, both from its symmetry and completeness, and from the interesting nature of the varied analogies and affinities which it brought to light and made use of. The series of Natural History volumes in "Lardner's Cabinet Cyclopaedia," in which Mr. Swainson developed it in most departments of the animal kingdom, made it widely known; and in fact for a long time these were the best and almost the only popular text-books for the rising generation of naturalists. It was favourably received too by the older school, which was perhaps rather an indication of its unsoundness. A considerable number of well-known naturalists either spoke approvingly of it, or advocated similar principles, and for a good many years it was decidedly in the ascendent. With such a favourable introduction, and with such talented exponents, it must have become established if it had had any germ of truth in it; yet it quite died out in a few short years, its very existence is now a matter of history; and so rapid was its fall that its talented creator, Swainson, perhaps lived to be the last man who believed in it.
Such is the course of a false theory. That of a true one is very different, as may be well seen by the progress of opinion on the subject of Natural Selection. In less than eight years "The Origin of Species" has produced conviction in the minds of a majority of the most eminent living men of science. New facts, new problems, new difficulties as they arise are accepted, solved or removed by this theory; and its principles are illustrated by the progress and conclusions of every well established branch of human knowledge. It is the object of the present essay to show how it has recently been applied to connect together and explain a variety of curious facts which had long been considered as inexplicable anomalies.
Importance of the Principle of Utility.
Perhaps no principle has ever been announced so fertile in results as that which Mr. Darwin so earnestly impresses upon us, and which is indeed a necessary deduction from the theory of Natural Selection, namely—that none of the definite facts of organic nature, no special organ, no characteristic form or marking, no peculiarities of instinct or of habit, no relations between species or between groups of species—can exist, but which must now be or once have been useful to the individuals or the races which possess them. This great principle gives us a clue which we can follow out in the study of many recondite phaenomena, and leads us to seek a meaning and a purpose of some definite character in minutiae which we should be otherwise almost sure to pass over as insignificant or unimportant.
Popular Theories of Colour in Animals.
The adaptation of the external colouring of animals to their conditions of life has long been recognised, and has been imputed either to an originally created specific peculiarity, or to the direct action of climate, soil, or food. Where the former explanation has been accepted, it has completely checked inquiry, since we could never get any further than the fact of the adaptation. There was nothing more to be known about the matter. The second explanation was soon found to be quite inadequate to deal with all the varied phases of the phaenomena, and to be contradicted by many well-known facts. For example, wild rabbits are always of grey or brown tints well suited for concealment among grass and fern. But when these rabbits are domesticated, without any change of climate or food, they vary into white or black, and these varieties may be multiplied to any extent, forming white or black races. Exactly the same thing has occurred with pigeons; and in the case of rats and mice, the white variety has not been shown to be at all dependent on alteration of climate, food, or other external conditions. In many cases the wings of an insect not only assume the exact tint of the bark or leaf it is accustomed to rest on, but the form and veining of the leaf or the exact rugosity of the bark is imitated; and these detailed modifications cannot be reasonably imputed to climate or to food, since in many cases the species does not feed on the substance it resembles, and when it does, no reasonable connexion can be shown to exist between the supposed cause and the effect produced. It was reserved for the theory of Natural Selection to solve all these problems, and many others which were not at first supposed to be directly connected with them. To make these latter intelligible, it will be necessary to give a sketch of the whole series of phaenomena which may be classed under the head of useful or protective resemblances.
Importance of Concealment as Influencing Colour.
Concealment, more or less complete, is useful to many animals, and absolutely essential to some. Those which have numerous enemies from which they cannot escape by rapidity of motion, find safety in concealment. Those which prey upon others must also be so constituted as not to alarm them by their presence or their approach, or they would soon die of hunger. Now it is remarkable in how many cases nature gives this boon to the animal, by colouring it with such tints as may best serve to enable it to escape from its enemies or to entrap its prey. Desert animals as a rule are desert-coloured. The lion is a typical example of this, and must be almost invisible when crouched upon the sand or among desert rocks and stones. Antelopes are all more or less sandy-coloured. The camel is pre-eminently so. The Egyptian cat and the Pampas cat are sandy or earth-coloured. The Australian kangaroos are of the same tints, and the original colour of the wild horse is supposed to have been a sandy or clay-colour.
The desert birds are still more remarkably protected by their assimilative hues. The stonechats, the larks, the quails, the goatsuckers and the grouse, which abound in the North African and Asiatic deserts, are all tinted and mottled so as to resemble with wonderful accuracy the average colour and aspect of the soil in the district they inhabit. The Rev. H. Tristram, in his account of the ornithology of North Africa in the 1st volume of the "Ibis," says: "In the desert, where neither trees, brush-wood, nor even undulation of the surface afford the slightest protection to its foes, a modification of colour which shall be assimilated to that of the surrounding country, is absolutely necessary. Hence without exception the upper plumage of every bird, whether lark, chat, sylvain, or sand-grouse, and also the fur of all the smaller mammals, and the skin of all the snakes and lizards, is of one uniform isabelline or sand colour." After the testimony of so able an observer it is unnecessary to adduce further examples of the protective colours of desert animals.