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On the Origin of Species

On the Origin of Species

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On the Origin of Species, 6th Edition

-- by Charles Darwin
On the Origin of Species, 6th Edition

-- by Charles Darwin

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The Project Gutenberg EBook of On the Origin of Species, by Charles Darwin This eBook is for the use of anyone

anywhere at no cost and with almost no restrictions whatsoever. You may copy it, give it away or re-use it under the terms of the Project Gutenberg License included with this eBook or online at www.gutenberg.org Title: On the Origin of Species 6th Edition Author: Charles Darwin Release Date: December, 1999 [Etext #2009] Posting Date: November 23, 2009 Language: English Character set encoding: ASCII *** START OF THIS PROJECT GUTENBERG EBOOK ON THE ORIGIN OF SPECIES ***

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THE ORIGIN OF SPECIES BY MEANS OF NATURAL SELECTION; or, the PRESERVATION OF FAVOURED RACES IN THE STRUGGLE FOR LIFE. By Charles Darwin, M.A., F.R.S., Author of "The Descent of Man," etc., etc. Sixth London Edition, with all Additions and Corrections. The 6th Edition is often considered the definitive edition. Also see Project Gutenberg Etext #1228 for the First Edition.

"But with regard to the material world, we can at least go this--we can perceive that events are brought about not by interpositions of Divine power, exerted in each particular the establishment of general laws."--Whewell: "Bridgewater

so far as insulated case, but by Treatise".

"The only distinct meaning of the word 'natural' is STATED, FIXED or SETTLED; since what is natural as much requires and presupposes an intelligent agent to render it so, i.e., to effect it continually or at stated times, as what is supernatural or miraculous does to effect it for once."--Butler: "Analogy of Revealed Religion". "To conclude, therefore, let no man out of a weak conceit of sobriety,

or an ill-applied moderation, think or maintain, that a man can search too far or be too well studied in the book of God's word, or in the book of God's works; divinity or philosophy; but rather let men endeavour an endless progress or proficience in both."--Bacon: "Advancement of Learning".

AN HISTORICAL SKETCH OF THE PROGRESS OF OPINION ON THE ORIGIN OF SPECIES, PREVIOUSLY TO THE PUBLICATION OF THE FIRST EDITION OF THIS WORK. I will here give a brief sketch of the progress of opinion on the Origin of Species. Until recently the great majority of naturalists believed that species were immutable productions, and had been separately created. This view has been ably maintained by many authors. Some few naturalists, on the other hand, have believed that species undergo modification, and that the existing forms of life are the descendants by true generation of pre existing forms. Passing over allusions to the subject in the classical writers (Aristotle, in his "Physicae Auscultationes" (lib.2, cap.8, s.2), after remarking that rain does not fall in order to make the corn grow, any more than it falls to spoil the farmer's corn when threshed out of doors, applies the same argument to organisation; and adds (as translated by Mr. Clair Grece, who first pointed out the passage to me), "So what hinders the different parts (of the body) from having this merely accidental relation in nature? as the teeth, for example, grow by necessity, the front ones sharp, adapted for dividing, and the grinders flat, and serviceable for masticating the food; since they were not made for the sake of this, but it was the result of accident. And in like manner as to other parts in which there appears to exist an adaptation to an end. Wheresoever, therefore, all things together (that is all the parts of one whole) happened like as if they were made for the sake of something, these were preserved, having been appropriately constituted by an internal spontaneity; and whatsoever things were not thus constituted, perished and still perish." We here see the principle of natural selection shadowed forth, but how little Aristotle fully comprehended the principle, is shown by his remarks on the formation of the teeth.), the first author who in modern times has treated it in a scientific spirit was Buffon. But as his opinions fluctuated greatly at different periods, and as he does not enter on the causes or means of the transformation of species, I need not here enter on details. Lamarck was the first man whose conclusions on the subject excited much attention. This justly celebrated naturalist first published his views in 1801; he much enlarged them in 1809 in his "Philosophie Zoologique", and subsequently, 1815, in the Introduction to his "Hist. Nat. des Animaux sans Vertebres". In these works he up holds the doctrine that all species, including man, are descended from other species. He first did the eminent service of arousing attention to the probability of all change in the organic, as well as in the inorganic world, being the result of law, and not of miraculous interposition. Lamarck seems to have been chiefly led to his conclusion on the gradual change of species, by the difficulty of distinguishing species and varieties, by the almost perfect gradation of forms in certain groups, and by the analogy of domestic productions. With respect to the means of modification, he attributed something to the direct action of the physical conditions of life, something to the crossing of already

existing forms, and much to use and disuse, that is, to the effects of habit. To this latter agency he seems to attribute all the beautiful adaptations in nature; such as the long neck of the giraffe for browsing on the branches of trees. But he likewise believed in a law of progressive development, and as all the forms of life thus tend to progress, in order to account for the existence at the present day of simple productions, he maintains that such forms are now spontaneously generated. (I have taken the date of the first publication of Lamarck from Isidore Geoffroy Saint-Hilaire's ("Hist. Nat. Generale", tom. ii. page 405, 1859) excellent history of opinion on this subject. In this work a full account is given of Buffon's conclusions on the same subject. It is curious how largely my grandfather, Dr. Erasmus Darwin, anticipated the views and erroneous grounds of opinion of Lamarck in his "Zoonomia" (vol. i. pages 500-510), published in 1794. According to Isid. Geoffroy there is no doubt that Goethe was an extreme partisan of similar views, as shown in the introduction to a work written in 1794 and 1795, but not published till long afterward; he has pointedly remarked ("Goethe als Naturforscher", von Dr. Karl Meding, s. 34) that the future question for naturalists will be how, for instance, cattle got their horns and not for what they are used. It is rather a singular instance of the manner in which similar views arise at about the same time, that Goethe in Germany, Dr. Darwin in England, and Geoffroy Saint-Hilaire (as we shall immediately see) in France, came to the same conclusion on the origin of species, in the years 1794-5.) Geoffroy Saint-Hilaire, as is stated in his "Life", written by his son, suspected, as early as 1795, that what we call species are various degenerations of the same type. It was not until 1828 that he published his conviction that the same forms have not been perpetuated since the origin of all things. Geoffroy seems to have relied chiefly on the conditions of life, or the "monde ambiant" as the cause of change. He was cautious in drawing conclusions, and did not believe that existing species are now undergoing modification; and, as his son adds, "C'est donc un probleme a reserver entierement a l'avenir, suppose meme que l'avenir doive avoir prise sur lui." In 1813 Dr. W.C. Wells read before the Royal Society "An Account of a White Female, part of whose skin resembles that of a Negro"; but his paper was not published until his famous "Two Essays upon Dew and Single Vision" appeared in 1818. In this paper he distinctly recognises the principle of natural selection, and this is the first recognition which has been indicated; but he applies it only to the races of man, and to certain characters alone. After remarking that negroes and mulattoes enjoy an immunity from certain tropical diseases, he observes, firstly, that all animals tend to vary in some degree, and, secondly, that agriculturists improve their domesticated animals by selection; and then, he adds, but what is done in this latter case "by art, seems to be done with equal efficacy, though more slowly, by nature, in the formation of varieties of mankind, fitted for the country which they inhabit. Of the accidental varieties of man, which would occur among the first few and scattered inhabitants of the middle regions of Africa, some one would be better fitted than others to bear the diseases of the country. This race would consequently multiply, while the others would decrease; not only from their in ability to sustain the attacks of disease, but from their incapacity of contending with their more vigorous neighbours. The colour of this vigorous race I take for granted, from what has been already said, would be dark. But the same disposition to form varieties still existing, a darker and a darker race would in the course of time occur: and as the darkest would be the best fitted for the climate, this would at length become the most prevalent,

if not the only race, in the particular country in which it had originated." He then extends these same views to the white inhabitants of colder climates. I am indebted to Mr. Rowley, of the United States, for having called my attention, through Mr. Brace, to the above passage of Dr. Wells' work. The Hon. and Rev. W. Herbert, afterward Dean of Manchester, in the fourth volume of the "Horticultural Transactions", 1822, and in his work on the "Amaryllidaceae" (1837, pages 19, 339), declares that "horticultural experiments have established, beyond the possibility of refutation, that botanical species are only a higher and more permanent class of varieties." He extends the same view to animals. The dean believes that single species of each genus were created in an originally highly plastic condition, and that these have produced, chiefly by inter-crossing, but likewise by variation, all our existing species. In 1826 Professor Grant, in the concluding paragraph in his well-known paper ("Edinburgh Philosophical Journal", vol. XIV, page 283) on the Spongilla, clearly declares his belief that species are descended from other species, and that they become improved in the course of modification. This same view was given in his Fifty-fifth Lecture, published in the "Lancet" in 1834. In 1831 Mr. Patrick Matthew published his work on "Naval Timber and Arboriculture", in which he gives precisely the same view on the origin of species as that (presently to be alluded to) propounded by Mr. Wallace and myself in the "Linnean Journal", and as that enlarged in the present volume. Unfortunately the view was given by Mr. Matthew very briefly in scattered passages in an appendix to a work on a different subject, so that it remained unnoticed until Mr. Matthew himself drew attention to it in the "Gardeners' Chronicle", on April 7, 1860. The differences of Mr. Matthew's views from mine are not of much importance: he seems to consider that the world was nearly depopulated at successive periods, and then restocked; and he gives as an alternative, that new forms may be generated "without the presence of any mold or germ of former aggregates." I am not sure that I understand some passages; but it seems that he attributes much influence to the direct action of the conditions of life. He clearly saw, however, the full force of the principle of natural selection. The celebrated geologist and naturalist, Von Buch, in his excellent "Description Physique des Isles Canaries" (1836, page 147), clearly expresses his belief that varieties slowly become changed into permanent species, which are no longer capable of intercrossing. Rafinesque, in his "New Flora of North America", published in 1836, wrote (page 6) as follows: "All species might have been varieties once, and many varieties are gradually becoming species by assuming constant and peculiar characters;" but further on (page 18) he adds, "except the original types or ancestors of the genus." In 1843-44 Professor Haldeman ("Boston Journal of Nat. Hist. U. States", vol. iv, page 468) has ably given the arguments for and against the hypothesis of the development and modification of species: he seems to lean toward the side of change. The "Vestiges of Creation" appeared in 1844. In the tenth and much improved edition (1853) the anonymous author says (page 155): "The proposition determined on after much consideration is, that the several series of animated beings, from the simplest and oldest up to the

highest and most recent, are, under the providence of God, the results, FIRST, of an impulse which has been imparted to the forms of life, advancing them, in definite times, by generation, through grades of organisation terminating in the highest dicotyledons and vertebrata, these grades being few in number, and generally marked by intervals of organic character, which we find to be a practical difficulty in ascertaining affinities; SECOND, of another impulse connected with the vital forces, tending, in the course of generations, to modify organic structures in accordance with external circumstances, as food, the nature of the habitat, and the meteoric agencies, these being the 'adaptations' of the natural theologian." The author apparently believes that organisation progresses by sudden leaps, but that the effects produced by the conditions of life are gradual. He argues with much force on general grounds that species are not immutable productions. But I cannot see how the two supposed "impulses" account in a scientific sense for the numerous and beautiful coadaptations which we see throughout nature; I cannot see that we thus gain any insight how, for instance, a woodpecker has become adapted to its peculiar habits of life. The work, from its powerful and brilliant style, though displaying in the early editions little accurate knowledge and a great want of scientific caution, immediately had a very wide circulation. In my opinion it has done excellent service in this country in calling attention to the subject, in removing prejudice, and in thus preparing the ground for the reception of analogous views. In 1846 the veteran geologist M.J. d'Omalius d'Halloy published in an excellent though short paper ("Bulletins de l'Acad. Roy. Bruxelles", tom. xiii, page 581) his opinion that it is more probable that new species have been produced by descent with modification than that they have been separately created: the author first promulgated this opinion in 1831. Professor Owen, in 1849 ("Nature of Limbs", page 86), wrote as follows: "The archetypal idea was manifested in the flesh under diverse such modifications, upon this planet, long prior to the existence of those animal species that actually exemplify it. To what natural laws or secondary causes the orderly succession and progression of such organic phenomena may have been committed, we, as yet, are ignorant." In his address to the British Association, in 1858, he speaks (page li) of "the axiom of the continuous operation of creative power, or of the ordained becoming of living things." Further on (page xc), after referring to geographical distribution, he adds, "These phenomena shake our confidence in the conclusion that the Apteryx of New Zealand and the Red Grouse of England were distinct creations in and for those islands respectively. Always, also, it may be well to bear in mind that by the word 'creation' the zoologist means 'a process he knows not what.'" He amplifies this idea by adding that when such cases as that of the Red Grouse are "enumerated by the zoologist as evidence of distinct creation of the bird in and for such islands, he chiefly expresses that he knows not how the Red Grouse came to be there, and there exclusively; signifying also, by this mode of expressing such ignorance, his belief that both the bird and the islands owed their origin to a great first Creative Cause." If we interpret these sentences given in the same address, one by the other, it appears that this eminent philosopher felt in 1858 his confidence shaken that the Apteryx and the Red Grouse first appeared in their respective homes "he knew not how," or by some process "he knew not what." This address was delivered after the papers by Mr. Wallace and myself on the Origin of Species, presently to be referred to, had been read before

the Linnean Society. When the first edition of this work was published, I was so completely deceived, as were many others, by such expressions as "the continuous operation of creative power," that I included Professor Owen with other palaeontologists as being firmly convinced of the immutability of species; but it appears ("Anat. of Vertebrates", vol. iii, page 796) that this was on my part a preposterous error. In the last edition of this work I inferred, and the inference still seems to me perfectly just, from a passage beginning with the words "no doubt the type-form," etc.(Ibid., vol. i, page xxxv), that Professor Owen admitted that natural selection may have done something in the formation of a new species; but this it appears (Ibid., vol. iii. page 798) is inaccurate and without evidence. I also gave some extracts from a correspondence between Professor Owen and the editor of the "London Review", from which it appeared manifest to the editor as well as to myself, that Professor Owen claimed to have promulgated the theory of natural selection before I had done so; and I expressed my surprise and satisfaction at this announcement; but as far as it is possible to understand certain recently published passages (Ibid., vol. iii. page 798) I have either partially or wholly again fallen into error. It is consolatory to me that others find Professor Owen's controversial writings as difficult to understand and to reconcile with each other, as I do. As far as the mere enunciation of the principle of natural selection is concerned, it is quite immaterial whether or not Professor Owen preceded me, for both of us, as shown in this historical sketch, were long ago preceded by Dr. Wells and Mr. Matthews. M. Isidore Geoffroy Saint-Hilaire, in his lectures delivered in 1850 (of which a Resume appeared in the "Revue et Mag. de Zoolog.", Jan., 1851), briefly gives his reason for believing that specific characters "sont fixes, pour chaque espece, tant qu'elle se perpetue au milieu des memes circonstances: ils se modifient, si les circonstances ambiantes viennent a changer. En resume, L'OBSERVATION des animaux sauvages demontre deja la variabilite LIMITEE des especes. Les EXPERIENCES sur les animaux sauvages devenus domestiques, et sur les animaux domestiques redevenus sauvages, la demontrent plus clairment encore. Ces memes experiences prouvent, de plus, que les differences produites peuvent etre de VALEUR GENERIQUE." In his "Hist. Nat. Generale" (tom. ii, page 430, 1859) he amplifies analogous conclusions. From a circular lately issued it appears that Dr. Freke, in 1851 ("Dublin Medical Press", page 322), propounded the doctrine that all organic beings have descended from one primordial form. His grounds of belief and treatment of the subject are wholly different from mine; but as Dr. Freke has now (1861) published his Essay on the "Origin of Species by means of Organic Affinity", the difficult attempt to give any idea of his views would be superfluous on my part. Mr. Herbert Spencer, in an Essay (originally published in the "Leader", March, 1852, and republished in his "Essays", in 1858), has contrasted the theories of the Creation and the Development of organic beings with remarkable skill and force. He argues from the analogy of domestic productions, from the changes which the embryos of many species undergo, from the difficulty of distinguishing species and varieties, and from the principle of general gradation, that species have been modified; and he attributes the modification to the change of circumstances. The author (1855) has also treated Psychology on the principle of the necessary acquirement of each mental power and capacity by gradation. In 1852 M. Naudin, a distinguished botanist, expressly stated, in an admirable paper on the Origin of Species ("Revue Horticole", page 102;

since partly republished in the "Nouvelles Archives du Museum", tom. i, page 171), his belief that species are formed in an analogous manner as varieties are under cultivation; and the latter process he attributes to man's power of selection. But he does not show how selection acts under nature. He believes, like Dean Herbert, that species, when nascent, were more plastic than at present. He lays weight on what he calls the principle of finality, "puissance mysterieuse, indeterminee; fatalite pour les uns; pour les autres volonte providentielle, dont l'action incessante sur les etres vivantes determine, a toutes les epoques de l'existence du monde, la forme, le volume, et la duree de chacun d'eux, en raison de sa destinee dans l'ordre de choses dont il fait partie. C'est cette puissance qui harmonise chaque membre a l'ensemble, en l'appropriant a la fonction qu'il doit remplir dans l'organisme general de la nature, fonction qui est pour lui sa raison d'etre." (From references in Bronn's "Untersuchungen uber die Entwickelungs-Gesetze", it appears that the celebrated botanist and palaeontologist Unger published, in 1852, his belief that species undergo development and modification. Dalton, likewise, in Pander and Dalton's work on Fossil Sloths, expressed, in 1821, a similar belief. Similar views have, as is well known, been maintained by Oken in his mystical "Natur-Philosophie". From other references in Godron's work "Sur l'Espece", it seems that Bory St. Vincent, Burdach, Poiret and Fries, have all admitted that new species are continually being produced. I may add, that of the thirty-four authors named in this Historical Sketch, who believe in the modification of species, or at least disbelieve in separate acts of creation, twenty-seven have written on special branches of natural history or geology.) In 1853 a celebrated geologist, Count Keyserling ("Bulletin de la Soc. Geolog.", 2nd Ser., tom. x, page 357), suggested that as new diseases, supposed to have been caused by some miasma have arisen and spread over the world, so at certain periods the germs of existing species may have been chemically affected by circumambient molecules of a particular nature, and thus have given rise to new forms. In this same year, 1853, Dr. Schaaffhausen published an excellent pamphlet ("Verhand. des Naturhist. Vereins der Preuss. Rheinlands", etc.), in which he maintains the development of organic forms on the earth. He infers that many species have kept true for long periods, whereas a few have become modified. The distinction of species he explains by the destruction of intermediate graduated forms. "Thus living plants and animals are not separated from the extinct by new creations, but are to be regarded as their descendants through continued reproduction." A well-known French botanist, M. Lecoq, writes in 1854 ("Etudes sur Geograph." Bot. tom. i, page 250), "On voit que nos recherches sur la fixite ou la variation de l'espece, nous conduisent directement aux idees emises par deux hommes justement celebres, Geoffroy Saint-Hilaire et Goethe." Some other passages scattered through M. Lecoq's large work make it a little doubtful how far he extends his views on the modification of species. The "Philosophy of Creation" has been treated in a masterly manner by the Rev. Baden Powell, in his "Essays on the Unity of Worlds", 1855. Nothing can be more striking than the manner in which he shows that the introduction of new species is "a regular, not a casual phenomenon," or, as Sir John Herschel expresses it, "a natural in contradistinction to a miraculous process."

Referring to such cases. VARIATION UNDER DOMESTICATION. Hooker published his "Introduction to the Australian Flora". and sadly damaged by some of its supporters. their existence would seem to show that the amount of modification which living beings have undergone during geological time is but very small in relation to the whole series of changes which they have suffered. 51) his conviction. the theory of Natural Selection is promulgated by Mr. 1859. by Mr. "Zoologisch-Anthropologische Untersuchungen". or each great type of organisation. that forms now perfectly distinct have descended from a single parent-form. and it is well to recollect that such an assumption is as unsupported by tradition or revelation as it is opposed to the general analogy of nature. The first edition of this work was published on November 24. Von Baer. he remarks. we view "Persistent Types" in relation to that hypothesis which supposes the species living at any time to be the result of the gradual modification of pre-existing species. is yet the only one to which physiology lends any countenance. a hypothesis which. if we suppose that each species of animal and plant. and the second edition on January 7. 1860. In June. "It is difficult to comprehend the meaning of such facts as these. In the first part of this great work he admits the truth of the descent and modification of species.The third volume of the "Journal of the Linnean Society" contains papers. as stated in the introductory remarks to this volume. chiefly grounded on the laws of geographical distribution. 1861. was formed and placed upon the surface of the globe at long intervals by a distinct act of creative power. 1859. read July 1. in which. Professor Huxley gave a lecture before the Royal Institution on the "Persistent Types of Animal Life". their Effects--Methodical and Unconscious Selection--Unknown Origin of our Domestic Productions--Circumstances favourable to Man's power of Selection. . Dr. 1859. on the other hand. though unproven. Wallace with admirable force and clearness. s. expressed about the year 1859 (see Prof. CONTENTS." In December. Wallace and myself. Rudolph Wagner. toward whom all zoologists feel so profound a respect. and supports this doctrine by many original observations. INTRODUCTION CHAPTER I. Causes of Variability--Effects of Habit and the use or disuse of Parts--Correlated Variation--Inheritance--Character of Domestic Varieties--Difficulty of distinguishing between Varieties and Species--Origin of Domestic Varieties from one or more Species--Domestic Pigeons. 1858. anciently followed. If. their Differences and Origin--Principles of Selection.

on the descendants from a common parent--Explains the Grouping of all organic beings--Advance in organisation--Low forms preserved--Convergence of character--Indefinite multiplication of species--Summary. intercrossing. VARIATION UNDER NATURE. Effects of changed conditions--Use and disuse.CHAPTER II. and common species. and in having restricted ranges. CHAPTER V. specific characters more variable than generic. related to each other. LAWS OF VARIATION. NATURAL SELECTION. 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. OR THE SURVIVAL OF THE FITTEST. number of individuals--Slow action--Extinction caused by Natural Selection--Divergence of Character. Its bearing on natural selection--The term used in a wide sense--Geometrical ratio of increase--Rapid increase of naturalised animals and plants--Nature of the checks to increase--Competition universal--Effects of climate--Protection from the number of individuals--Complex relations of all animals and plants throughout nature--Struggle for life most severe between individuals and varieties of the same species. CHAPTER IV. secondary sexual characters variable--Species of the same genus vary in an analogous manner--Reversions to long-lost characters--Summary. CHAPTER III. related to the diversity of inhabitants of any small area and to naturalisation--Action of Natural Selection. rudimentary. Variability--Individual Differences--Doubtful species--Wide ranging. much diffused. isolation. and lowly organised structures variable--Parts developed in an unusual manner are highly variable. CHAPTER VI. often severe between species of the same genus--The relation of organism to organism the most important of all relations. through Divergence of Character and Extinction. combined with natural selection. vary most--Species of the larger genera in each country vary more frequently than the species of the smaller genera--Many of the species of the larger genera resemble varieties in being very closely. but unequally. organs of flight and of vision--Acclimatisation--Correlated variation--Compensation and economy of growth--False correlations--Multiple. namely. . STRUGGLE FOR EXISTENCE.

the most constant--Supposed incompetence of natural selection to account for the incipient stages of useful structures--Causes which interfere with the acquisition through natural selection of useful structures--Gradations of structure with changed functions--Widely different organs in members of the same class. Difficulties of the theory of descent with modification--Absence or rarity of transitional varieties--Transitions in habits of life--Diversified habits in the same species--Species with habits widely different from those of their allies--Organs of extreme perfection--Modes of transition--Cases of difficulty--Natura non facit saltum--Organs of small importance--Organs not in all cases absolutely perfect--The law of Unity of Type and of the Conditions of Existence embraced by the theory of Natural Selection. CHAPTER VIII. . its cell-making instinct--Changes of instinct and structure not necessarily simultaneous--Difficulties on the theory of the Natural Selection of instincts--Neuter or sterile insects--Summary. molothrus. developed from one and the same source--Reasons for disbelieving in great and abrupt modifications. INSTINCT.DIFFICULTIES OF THE THEORY. but different in their origin--Instincts graduated--Aphides and ants--Instincts variable--Domestic instincts. removed by domestication--Laws governing the sterility of hybrids--Sterility not a special endowment. CHAPTER IX. CHAPTER X. affected by close interbreeding. Distinction between the sterility of first crosses and of hybrids--Sterility various in degree. HYBRIDISM. and parasitic bees--Slave-making ants--Hive-bee. ostrich. CHAPTER VII. ON THE IMPERFECTION OF THE GEOLOGICAL RECORD. not universal. MISCELLANEOUS OBJECTIONS TO THE THEORY OF NATURAL SELECTION. Instincts comparable with habits. not accumulated by natural selection--Causes of the sterility of first crosses and of hybrids--Parallelism between the effects of changed conditions of life and of crossing--Dimorphism and Trimorphism--Fertility of varieties when crossed and of their mongrel offspring not universal--Hybrids and mongrels compared independently of their fertility--Summary. Longevity--Modifications not necessarily simultaneous--Modifications apparently of no direct service--Progressive development--Characters of small functional importance. their origin--Natural instincts of the cuckoo. but incidental on other differences.

ON THE GEOLOGICAL SUCCESSION OF ORGANIC BEINGS. explained on the theory of descent with modification--Classification of varieties--Descent always used in classification--Analogical or adaptive characters--Affinities. GEOGRAPHICAL DISTRIBUTION--CONTINUED. CHAPTER XIV. between parts of the same individual--Embryology.On the absence of intermediate varieties at the present day--On the nature of extinct intermediate varieties. between members of the same class. Distribution of fresh-water productions--On the inhabitants of oceanic islands--Absence of Batrachians and of terrestrial Mammals--On the relation of the inhabitants of islands to those of the nearest mainland--On colonisation from the nearest source with subsequent modification--Summary of the last and present chapter. complex and radiating--Extinction separates and defines groups--Morphology. GEOGRAPHICAL DISTRIBUTION. as inferred from the rate of denudation and of deposition--On the lapse of time as estimated in years--On the poorness of our palaeontological collections--On the intermittence of geological formations--On the denudation of granitic areas--On the absence of intermediate varieties in any one formation--On the sudden appearance of groups of species--On their sudden appearance in the lowest known fossiliferous strata--Antiquity of the habitable earth. laws of. on their number--On the lapse of time. general. CHAPTER XII. and by occasional means--Dispersal during the Glacial period--Alternate Glacial periods in the north and south. groups subordinate to groups--Natural system--Rules and difficulties in classification. CHAPTER XIII. On the slow and successive appearance of new species--On their different rates of change--Species once lost do not reappear--Groups of species follow the same general rules in their appearance and disappearance as do single species--On extinction--On simultaneous changes in the forms of life throughout the world--On the affinities of extinct species to each other and to living species--On the state of development of ancient forms--On the succession of the same types within the same areas--Summary of preceding and present chapter. MUTUAL AFFINITIES OF ORGANIC BEINGS: MORPHOLOGY--EMBRYOLOGY--RUDIMENTARY ORGANS. CHAPTER XI. explained by variations not . Present distribution cannot be accounted for by differences in physical conditions--Importance of barriers--Affinity of the productions of the same continent--Centres of creation--Means of dispersal by changes of climate and of the level of the land. Classification.

GLOSSARY OF SCIENTIFIC TERMS.supervening at an early age. which then seemed to me probable: from that period to the present day I have steadily pursued the same object. Hooker. with a request that I would forward it to Sir Charles Lyell. Wallace's excellent memoir. but as it will take me many more years to complete it. On my return home.S. I was much struck with certain facts in the distribution of the organic beings inhabiting South America. Wallace. After five years' work I allowed myself to speculate on the subject. and as my health is far from strong. ORIGIN OF SPECIES. seemed to throw some light on the origin of species--that mystery of mysteries. I have been urged to publish this abstract. who is now studying the natural history of the Malay Archipelago. RECAPITULATION AND CONCLUSION. as I give them to show that I have not been hasty in coming to a decision. as will be seen in the latter chapters of this volume. their origin explained--Summary. as it has been called by one of our greatest philosophers. who both knew of my work--the latter having read my sketch of 1844--honoured me by thinking it advisable to publish. and in the geological relations of the present to the past inhabitants of that continent. These facts. . CHAPTER XV. with Mr. has arrived at almost exactly the same general conclusions that I have on the origin of species. Recapitulation of the objections to the theory of Natural Selection--Recapitulation of the general and special circumstances in its favour--Causes of the general belief in the immutability of species--How far the theory of Natural Selection may be extended--Effects of its adoption on the study of Natural history--Concluding remarks. some brief extracts from my manuscripts. and it is published in the third volume of the Journal of that Society. these I enlarged in 1844 into a sketch of the conclusions. it occurred to me. I have more especially been induced to do this. I hope that I may be excused for entering on these personal details. Lyell and Dr. My work is now (1859) nearly finished. Sir C. When on board H. as Mr. In 1858 he sent me a memoir on this subject. INTRODUCTION. and drew up some short notes.M. who sent it to the Linnean Society. Beagle. in 1837. as naturalist. and being inherited at a corresponding age--Rudimentary Organs. INDEX. that something might perhaps be made out on this question by patiently accumulating and reflecting on all sorts of facts which could possibly have any bearing on it.

It is. with a few facts in illustration. I cannot here give references and authorities for my several statements. food. or of habit. let this opportunity pass without expressing my deep obligations to Dr. Nevertheless. like varieties. although they have been very commonly neglected by naturalists. so admirably adapted to catch insects under the bark of trees. imperfect though it be. so as to acquire that perfection of structure and coadaptation which justly excites our admiration. I may venture to express my conviction of the high value of such studies. or of the volition of the plant itself. Hooker. on which my conclusions have been grounded. tail. A fair result can be obtained only by fully stating and balancing the facts and arguments on both sides of each question. and I hope in a future work to do this. in most cases will suffice. the structure. with its feet. . and this is here impossible. for instance. of variation under domestication. afforded the best and safest clue. even if well founded.This abstract. but which. such a conclusion. for the last fifteen years.. until it could be shown how the innumerable species. I hope. in this and in all other perplexing cases I have invariably found that our knowledge. which has seeds that must be transported by certain birds. who. on their embryological relations. from other species. reflecting on the mutual affinities of organic beings. and which has flowers with separate sexes absolutely requiring the agency of certain insects to bring pollen from one flower to the other. but it is preposterous to attribute to mere external conditions. of the woodpecker. I cannot. some of them personally unknown to me. In the case of the mistletoe. and I must trust to the reader reposing some confidence in my accuracy. of the highest importance to gain a clear insight into the means of modification and coadaptation. For I am well aware that scarcely a single point is discussed in this volume on which facts cannot be adduced. No doubt errors may have crept in. though I hope I have always been cautious in trusting to good authorities alone. often apparently leading to conclusions directly opposite to those at which I have arrived. but had descended. At the commencement of my observations it seemed to me probable that a careful study of domesticated animals and of cultivated plants would offer the best chance of making out this obscure problem. beak. it is equally preposterous to account for the structure of this parasite. such as climate. this may be true. has aided me in every possible way by his large stores of knowledge and his excellent judgment. however. therefore. inhabiting this world have been modified. Nor have I been disappointed. which I now publish. In one limited sense. which draws its nourishment from certain trees. as we shall hereafter see. and other such facts. I much regret that want of space prevents my having the satisfaction of acknowledging the generous assistance which I have received from very many naturalists. with references. with its relations to several distinct organic beings. No one can feel more sensible than I do of the necessity of hereafter publishing in detail all the facts. I can here give only the general conclusions at which I have arrived. Naturalists continually refer to external conditions. as the only possible cause of variation. it is quite conceivable that a naturalist. their geographical distribution. by the effects of external conditions. might come to the conclusion that species had not been independently created. etc. In considering the origin of species. geological succession. and tongue. must necessarily be imperfect. would be unsatisfactory.

under the complex and sometimes varying conditions of life. I will then pass on to the variability of species in a state of nature. In the next chapter I shall consider the geological succession of organic beings throughout time. Still less do we know of the mutual relations of the innumerable inhabitants of the world during the many past geological epochs in its history. and leads to what I have called divergence of character. that the view which most naturalists until recently entertained. in the fourteenth. and which I formerly entertained--namely. or how a simple being or a simple organ can be changed and perfected into a highly developed being or into an elaborately constructed organ. be enabled to discuss what circumstances are most favourable to variation. if he make due allowance for our profound ignorance in regard to the mutual relations of the many beings which live around us. if it vary however slightly in any manner profitable to itself. I am fully convinced that species are not immutable. consequently. in the same manner as the acknowledged varieties of . and will long remain obscure. We shall. No one ought to feel surprise at much remaining as yet unexplained in regard to the origin of species and varieties. In the five succeeding chapters. in the twelfth and thirteenth. In the next chapter I shall discuss the complex and little known laws of variation. there is a frequently recurring struggle for existence. however. that each species has been independently created--is erroneous. I shall devote the first chapter of this abstract to variation under domestication. the most apparent and gravest difficulties in accepting the theory will be given: namely. will have a better chance of surviving. We shall thus see that a large amount of hereditary modification is at least possible. as it can be treated properly only by giving long catalogues of facts. and as. their classification or mutual affinities. thirdly. hybridism. the future success and modification of every inhabitant of this world. any selected variety will tend to propagate its new and modified form. or the mental powers of animals. unfortunately. and we shall then see how natural selection almost inevitably causes much extinction of the less improved forms of life. and thus be NATURALLY SELECTED. and. In the last chapter I shall give a brief recapitulation of the whole work. This is the doctrine of Malthus. what is equally or more important. be compelled to treat this subject far too briefly. and why another allied species has a narrow range and is rare? Yet these relations are of the highest importance. the imperfection of the geological record. first. and fourthly. or the infertility of species and the fertility of varieties when intercrossed. I can entertain no doubt. and a few concluding remarks. it follows that any being. the difficulties of transitions. but I shall. both when mature and in an embryonic condition. Who can explain why one species ranges widely and is very numerous. applied to the whole animal and vegetable kingdoms. as I believe. but that those belonging to what are called the same genera are lineal descendants of some other and generally extinct species. we shall see how great is the power of man in accumulating by his selection successive slight variations. In the next chapter the struggle for existence among all organic beings throughout the world. Although much remains obscure. which inevitably follows from the high geometrical ratio of their increase. for they determine the present welfare and. secondly the subject of instinct.From these considerations. As many more individuals of each species are born than can possibly survive. after the most deliberate study and dispassionate judgment of which I am capable. This fundamental subject of natural selection will be treated at some length in the fourth chapter. From the strong principle of inheritance. their geographical distribution throughout space. will be considered.

for nearly similar variations sometimes arise under. and that. dissimilar variations arise under conditions which appear to be nearly uniform. but not the exclusive. when the organisation has once begun to vary. They may be considered as definite when all or nearly all the offspring of individuals exposed to certain conditions during several generations are modified in the same manner. the nature of the organism and the nature of the conditions. colour from the nature of the food.any one species are the descendants of that species. The former seems to be much the more important. as far as we can judge. we must bear in mind that in every case. and as I have incidently shown in my work on "Variation under Domestication. we are driven to conclude that this great variability is due to our domestic productions having been raised under conditions of life not so uniform as. The effects on the offspring are either definite or in definite. it generally continues varying for many generations. thickness of the skin and hair from . When we compare the individuals of the same variety or sub-variety of our older cultivated plants and animals. as Professor Weismann has lately insisted. one of the first points which strikes us is. There can. on the other hand. Causes of Variability--Effects of Habit and the use and disuse of Parts--Correlated Variation--Inheritance--Character of Domestic Varieties--Difficulty of distinguishing between Varieties and Species--Origin of Domestic Varieties from one or more Species--Domestic Pigeons. CHAPTER I. I am convinced that natural selection has been the most important. their Differences and Origin--Principles of Selection. CAUSES OF VARIABILITY. means of modification. the conditions of life appear to act in two ways--directly on the whole organisation or on certain parts alone and in directly by affecting the reproductive system." there are two factors: namely. Our oldest cultivated plants. dissimilar conditions. such as size from the amount of food. It seems clear that organic beings must be exposed during several generations to new conditions to cause any great amount of variation. With respect to the direct action. and which have varied during all ages under the most different climates and treatment. No case is on record of a variable organism ceasing to vary under cultivation. be little doubt about many slight changes. And if we reflect on the vast diversity of the plants and animals which have been cultivated. however. such as wheat. Furthermore. anciently followed. and. As far as I am able to judge. after long attending to the subject. still yield new varieties: our oldest domesticated animals are still capable of rapid improvement or modification. VARIATION UNDER DOMESTICATION. It is extremely difficult to come to any conclusion in regard to the extent of the changes which have been thus definitely induced. There is. those to which the parent species had been exposed under nature. that this variability may be partly connected with excess of food. that they generally differ more from each other than do the individuals of any one species or variety in a state of nature. also. some probability in the view propounded by Andrew Knight. their Effects--Methodical and Unconscious Selection--Unknown Origin of our Domestic Productions--Circumstances favourable to Man's power of Selection. and somewhat different from.

and in seedlings from the same seed-capsule. Each of the endless variations which we see in the plumage of our fowls must have had some efficient cause. but to show how singular the laws are which determine the reproduction of animals under confinement. deviations of structure so strongly pronounced as to deserve to be called monstrosities arise. causing coughs or colds. we see individuals. between the variability which follows from the crossing of distinct species. and when. namely. Such facts as the complex and extraordinary out growths which variably follow from the insertion of a minute drop of poison by a gall-producing insect. In definite variability is a much more common result of changed conditions than definite variability. breed in this country pretty freely under confinement. may be considered as the in definite effects of the conditions of life on each individual organism. Many exotic plants have pollen utterly worthless. on the one hand. we see domesticated animals and plants. and if the same cause were to act uniformly during a long series of generations on many individuals. but monstrosities cannot be separated by any distinct line from slighter variations. and yet rarely or never seed! In some few cases it has been discovered that a very trifling change. whereas. and healthy (of which I could give numerous . according to their state of body or constitution.climate. and which cannot be accounted for by inheritance from either parent or from some more remote ancestor. partly from the fact of this system being extremely sensitive to any change in the conditions. I may mention that carnivorous animals. Nothing is more easy than to tame an animal. How many animals there are which will not breed. rheumatism. even when the male and female unite. such as a little more or less water at some particular period of growth. which seldom produce young. even from the tropics. long-lived. in the same condition as in the most sterile hybrids. carnivorous birds. etc. whether extremely slight or strongly marked. through the reproductive system of being affected. Even strongly-marked differences occasionally appear in the young of the same litter. on the other hand. will determine whether or not a plant will produce seeds. We see in definite variability in the endless slight peculiarities which distinguish the individuals of the same species. or inflammation of various organs. though often weak and sickly. as Kolreuter and others have remarked. and has probably played a more important part in the formation of our domestic races. all probably would be modified in the same manner. shows us what singular modifications might result in the case of plants from a chemical change in the nature of the sap. though kept in an almost free state in their native country! This is generally. and partly from the similarity. but erroneously attributed to vitiated instincts. I cannot here give the details which I have collected and elsewhere published on this curious subject. With respect to what I have called the in direct action of changed conditions. At long intervals of time. though taken young from a state of nature perfectly tamed. which appear among many individuals living together. with the exception of the plantigrades or bear family. hardly ever lay fertile eggs. All such changes of structure. and few things more difficult than to get it to breed freely under confinement. in nearly the same manner as the chill effects different men in an in definite manner. Many facts clearly show how eminently susceptible the reproductive system is to very slight changes in the surrounding conditions. we may infer that variability is thus induced. with the rarest exception. out of millions of individuals reared in the same country and fed on nearly the same food. and that which may be observed with plants and animals when reared under new or unnatural conditions. When. breeding freely under confinement. Many cultivated plants display the utmost vigour.

that is. With animals the increased use or disuse of parts has had a more marked influence. and the view which has been suggested that the drooping is due to disuse of the muscles of the ear. has been known suddenly to assume a new character. and vary very slightly--perhaps hardly more than in a state of nature. is probably another instance of the effects of use. I will here only allude to what may be called correlated variation. we need not be surprised at this system. as they may be named. Important changes in the embryo or larva will probably entail changes in the mature animal. thus cats which are entirely white and have blue eyes are generally deaf. yet having their reproductive system so seriously affected by unperceived causes as to fail to act. As a single bud out of many thousands produced year after year on the same tree under uniform conditions. have sometimes yielded nearly the same variety--for instance. CORRELATED VARIATION. and this change may be safely attributed to the domestic duck flying much less. and many instances are given in Isidore Geoffroy St. but this is certainly an error. and as buds on distinct trees. EFFECTS OF HABIT AND OF THE USE OR DISUSE OF PARTS. They occur rarely under nature. but are far from rare under culture. of plants which have suddenly produced a single bud with a new and sometimes widely different character from that of the other buds on the same plant. seems probable. Breeders believe that long limbs are almost always accompanied by an elongated head. etc. I may add that as some organisms breed freely under the most unnatural conditions--for instance." as they are called by gardeners. Not one of our domestic animals can be named which has not in some country drooping ears. when it does act under confinement. Many laws regulate variation. Some naturalists have maintained that all variations are connected with the act of sexual reproduction. acting irregularly. rabbits and ferrets kept in hutches--showing that their reproductive organs are not easily affected. and sometimes by seed. buds on peach-trees producing nectarines. has in determining the nature of the flames. perhaps of not more importance than the nature of the spark. but it has been lately stated by Mr. In monstrosities. and buds on common roses producing moss-roses--we clearly see that the nature of the conditions is of subordinate importance in comparison with the nature of the organism in determining each particular form of variation. for I have given in another work a long list of "sporting plants. of which many remarkable cases could be given among animals and plants. Some instances of correlation are quite whimsical. INHERITANCE. and walking more. Hilaire's great work on this subject. so will some animals and plants withstand domestication or cultivation.instances). than its wild parents. the correlations between quite distinct parts are very curious. thus I find in the domestic duck that the bones of the wing weigh less and the bones of the leg more. Colour and constitutional peculiarities go together. Tait that this is confined to the males. than do the same bones in the wild duck. some few of which can be dimly seen. growing under different conditions. in proportion to the whole skeleton. The great and inherited development of the udders in cows and goats in countries where they are habitually milked. Changed habits produce an inherited effect as in the period of the flowering of plants when transported from one climate to another. can be propagated by grafts. From . These bud variations.. and will hereafter be briefly discussed. from the animals being seldom much alarmed. by which a mass of combustible matter is ignited. in comparison with these organs in other countries. and producing offspring somewhat unlike their parents. offsets.

to the males alone. to look at the inheritance of every character whatever as the rule. Virginia squatters) added. that like produces like is his fundamental belief: doubts have been thrown on this principle only by theoretical writers. as on the hyacinth. and which caused the hoofs of all but the black varieties to drop off. apparently exposed to the same conditions. Dr. which I think may be trusted.. both those of slight and those of considerable physiological importance. which coloured their bones pink. If strange and rare deviations of structure are truly inherited. Any variation which is not inherited is unimportant for us. prickly skin. or in different species. at whatever period of life a peculiarity first appears. and it is really surprising to note the endless points of structure and constitution in which the varieties and sub-varieties differ slightly from each other." Hairless dogs have imperfect teeth. it appears that white sheep and pigs are injured by certain plants. once among several million individuals--and it reappears in the child. less strange and commoner deviations may be freely admitted to be inheritable. are endless. the mere doctrine of chances almost compels us to attribute its reappearance to inheritance. no one can say why the same peculiarity in different individuals of the same species. and one of the "crackers" (i. The whole organisation seems to have become plastic. we cannot tell whether it may not be due to the same cause having acted on both. more commonly but not exclusively to the like sex. and we see it in the father and child. long-haired and coarse-haired animals are apt to have. and thus augmenting. long or many horns. as they alone have a good chance of living. why the child often reverts in certain characteristics to its grandfather or grandmother or more remote ancestor. either exclusively or in a much greater degree. potato. and those with long beaks large feet. any peculiarity. in two large volumes. or to one sex alone. Hence if man goes on selecting. The laws governing inheritance are for the most part unknown. owing to the mysterious laws of correlation. any very rare deviation. due to some extraordinary combination of circumstances. pigeons with short beaks have small feet. It is well worth while carefully to study the several treatises on some of our old cultivated plants. is the fullest and the best on this subject. it tends to reappear in the offspring at a corresponding . and departs in a slight degree from that of the parental type. Every one must have heard of cases of albinism. A much more important rule. is sometimes inherited and sometimes not so. etc. When any deviation of structure often appears. as is asserted. that peculiarities appearing in the males of our domestic breeds are often transmitted. even the dahlia. appears in the parent--say. is that. but when among individuals. and non-inheritance as the anomaly. The results of the various. on asking some farmers in Virginia how it was that all their pigs were black. unknown. appearing in several members of the same family.e. Prosper Lucas' treatise. Perhaps the correct way of viewing the whole subject would be. hairy bodies. while dark-coloured individuals escape: Professor Wyman has recently communicated to me a good illustration of this fact. he will almost certainly modify unintentionally other parts of the structure. No breeder doubts how strong is the tendency to inheritance. or but dimly understood laws of variation are infinitely complex and diversified. pigeons with feathered feet have skin between their outer toes. "we select the black members of a litter for raising. why a peculiarity is often transmitted from one sex to both sexes. etc.facts collected by Heusinger.. It is a fact of some importance to us. But the number and diversity of inheritable deviations of structure. they informed him that the pigs ate the paint-root (Lachnanthes).

though sometimes earlier. in . Domestic races often have a somewhat monstrous character. I believe this rule to be of the highest importance in explaining the laws of embryology. any slight deviations in their structure. If it could be shown that our domestic varieties manifested a strong tendency to reversion--that is. it seems to me not improbable that if we could succeed in naturalising. so that free intercrossing might check. and so could not tell whether or not nearly perfect reversion had ensued. that our domestic varieties. and not to the primary cause which may have acted on the ovules or on the male element.age. in nearly the same manner as the increased length of the horns in the offspring from a short-horned cow by a long-horned bull. that only a single variety should be turned loose in its new home. we generally perceive in each domestic race. Whether or not the experiment would succeed is not of great importance for our line of argument. would be opposed to all experience. of the cabbage. and that. by which I mean. Having alluded to the subject of reversion. I may here refer to a statement often made by naturalists--namely. Nevertheless. in very poor soil--in which case. There would be great difficulty in proving its truth: we may safely conclude that very many of the most strongly marked domestic varieties could not possibly live in a wild state. when run wild. When we look to the hereditary varieties or races of our domestic animals and plants. But there is not a shadow of evidence in favour of this view: to assert that we could not breed our cart and race-horses. some effect would have to be attributed to the DEFINITE action of the poor soil--that they would. revert to the wild aboriginal stock. and compare them with closely allied species. But hereditary diseases and some other facts make me believe that the rule has a wider extension. however. or even wholly. and esculent vegetables. to lose their acquired characters. during many generations. as already remarked. for an unlimited number of generations. CHARACTER OF DOMESTIC VARIETIES. yet that it does tend to appear in the offspring at the same period at which it first appeared in the parent. is clearly due to the male element. for by the experiment itself the conditions of life are changed. or were to cultivate. I have in vain endeavoured to discover on what decisive facts the above statement has so often and so boldly been made. by blending together. long and short-horned cattle. although differing from each other and from other species of the same genus. I grant that we could deduce nothing from domestic varieties in regard to species. thus the inherited peculiarities in the horns of cattle could appear only in the offspring when nearly mature. It would be necessary. DIFFICULTY OF DISTINGUISHING BETWEEN VARIETIES AND SPECIES. as our varieties certainly do occasionally revert in some of their characters to ancestral forms. though appearing late in life. ORIGIN OF DOMESTIC VARIETIES FROM ONE OR MORE SPECIES. the several races. Hence it has been argued that no deductions can be drawn from domestic races to species in a state of nature. that. gradually but invariably revert in character to their aboriginal stocks. in such case. to a large extent. In many cases this could not be otherwise. These remarks are of course confined to the first APPEARANCE of the peculiarity. while kept under the same conditions and while kept in a considerable body. in order to prevent the effects of intercrossing. and poultry of various breeds. In many cases we do not know what the aboriginal stock was. for instance. when there is no apparent reason why a peculiarity should appear at any particular age. less uniformity of character than in true species. peculiarities in the silk-worm are known to appear at the corresponding caterpillar or cocoon stage.

when he first tamed an animal. With these exceptions (and with that of the perfect fertility of varieties when crossed--a subject hereafter to be discussed). but how could a savage possibly know. there is presumptive or even strong evidence that all are descended from a single wild stock. The argument mainly relied on by those who believe in the multiple origin of our domestic animals is. spaniel and bull-dog. but the differences in most cases are less in degree. I do not believe. it could be shown that the greyhound. and could be made to breed for an equal number of generations under domestication. I believe that a small part of the difference is due to their being descended from distinct species. This must be admitted as true. It has often been assumed that man has chosen for domestication animals and plants having an extraordinary inherent tendency to vary. would be interesting. but naturalists differ much in determining what characters are of generic value. In the case of strongly marked races of some other domesticated species. we are soon involved in doubt. In the case of most of our anciently domesticated animals and plants. and in the lake-habitations of Switzerland. bloodhound. and belonging to equally diverse classes and countries. and more especially when compared with the species under nature to which they are nearest allied. from not knowing whether they are descended from one or several parent species. it will be seen that we have no right to expect often to find a generic amount of difference in our domesticated races. on the monuments of Egypt. that the whole amount of difference between the several breeds of the dog has been produced under domestication. both when compared one with another. prevented their domestication? I cannot doubt that if other animals and plants. In attempting to estimate the amount of structural difference between allied domestic races. this source of doubt would not so perpetually recur. all such valuations being at present empirical. whether they are descended from one or several wild species. domestic races of the same species differ from each other in the same manner as do the closely allied species of the same genus in a state of nature. whether it would vary in succeeding generations. which we all know propagate their kind truly. if it could be cleared up. as we shall presently see. if. much diversity in the . and by other competent judges as mere varieties. for instance. it is not possible to come to any definite conclusion. It can be shown that this statement is not correct. they would on an average vary as largely as the parent species of our existing domesticated productions have varied. and likewise to withstand diverse climates. and whether it would endure other climates? Has the little variability of the ass and goose. equal in number to our domesticated productions. were the offspring of any single species. or of cold by the common camel. were taken from a state of nature. If any well marked distinction existed between a domestic race and a species.several trifling respects. for the domestic races of many animals and plants have been ranked by some competent judges as the descendants of aboriginally distinct species. they often differ in an extreme degree in some one part. This point. or the small power of endurance of warmth by the reindeer. that we find in the most ancient times. It has often been stated that domestic races do not differ from each other in characters of generic value. When it is explained how genera originate under nature. I do not dispute that these capacities have added largely to the value of most of our domesticated productions. terrier. then such facts would have great weight in making us doubt about the immutability of the many closely allied natural species--for instance. of the many foxes--inhabiting the different quarters of the world.

But I may here state that. etc. In regard to ducks and rabbits. some breeds of which differ much from each other. and we know that at the present day there is hardly a tribe so barbarous as not to have domesticated at least the dog. indeed. kept by different tribes in different districts. that all the races belong to the same species. Spain. One author believes that there formerly existed eleven wild species of sheep peculiar to Great Britain! When we bear in mind that Britain has now not one peculiar mammal. Even in the case of the breeds of the domestic dog throughout the world. flows in the veins of our domestic breeds. Blyth. The origin of most of our domestic animals will probably forever remain vague. on the habits. I have. as well as that of the specific distinction between the humped and common cattle. From facts communicated to me by Mr. all geologists believe that barbarian men existed at an enormously remote period. They also carried on commerce with other nations. which I admit are descended from several wild species. during which the domesticated animals. may. and shows that animals were domesticated at a much earlier period than has hitherto been supposed. and France but few distinct from those of Germany. be looked upon as established by the admirable researches of Professor Rutimeyer. Gallus bankiva. Having kept nearly all the English breeds of the fowl alive. constitution and structure of the humped Indian cattle. and they possessed several domesticated animals. the evidence is clear that they are all descended from the common duck and wild rabbit. and of others who have studied this bird in India. as Heer has remarked. etc. it appears to me almost certain that all are the descendants of the wild Indian fowl. At this rate there must have existed at least a score of species of wild cattle. and that their blood. and this again implies a long continued previous period of less advanced civilisation. This conclusion. in some cases mingled together. or are even identical with.breeds. the pea. the poppy for oil and flax. but that each of these kingdoms possesses several peculiar breeds of cattle. has been carried to an absurd extreme by some authors. that they had at this early age progressed considerably in civilisation. The doctrine of the origin of our several domestic races from several aboriginal stocks. from reasons which I cannot here give. in Europe alone. having bred and crossed them.. looking to the domestic dogs of the whole world. it cannot be doubted that there has been an . and several goats. after a laborious collection of all known facts. and several even within Great Britain. those still existing. let the distinctive characters be ever so slight. Since the discovery of flint tools in the superficial formations of many parts of the world. and this is the conclusion of Mr. come to the conclusion that several wild species of Canidae have been tamed. whether or not these deserve to be called species. as many sheep. and some competent judges believe that these latter have had two or three wild progenitors. With respect to horses. The lake-inhabitants of Switzerland cultivated several kinds of wheat and barley. we must admit that many domestic breeds must have originated in Europe. All this clearly shows. Blyth. They believe that every race which breeds true. In regard to sheep and goats I can form no decided opinion. it is almost certain that they are descended from a different aboriginal stock from our European cattle. and that some of these ancient breeds closely resemble. in opposition to several authors. for whence otherwise could they have been derived? So it is in India. and so with Hungary. I am doubtfully inclined to believe. and examined their skeletons. has had its wild prototype. might have varied and given rise to distinct races. sheep.. But this only throws far backward the history of civilisation. voice.

but. bull-dog. proportionally to its size. bloodhound. The fantail has thirty or even forty tail-feathers. C. instead of a long beak. and if we account for our several domestic races by this process. has a very short and broad one. Believing that it is always best to study some special group. with long. the normal number in all the members of the great pigeon family: these feathers are kept expanded and are carried so erect that in good birds the head and tail touch: the oil-gland is quite aborted. and have been permitted to join two of the London Pigeon Clubs. pug-dog. and have been most kindly favoured with skins from several quarters of the world.. The turbit has a short and conical beak. the bloodhound. as being of considerable antiquity. as the Italian greyhound. in the wild state. very large external orifices to the nostrils. The trumpeter and laugher. but to obtain a race intermediate between two quite distinct races would be very difficult. which it glories in inflating. but when these mongrels are crossed one with another for several generations. after deliberation.--so unlike all wild Canidae--ever existed in a state of nature? It has often been loosely said that all our races of dogs have been produced by the crossing of a few aboriginal species. the possibility of making distinct races by crossing has been greatly exaggerated. and every thing seems simple enough. instead of twelve or fourteen. Sebright expressly experimented with this object and failed. and some of them are very important. Elliot from India. and tumbling in the air head over heels. Many cases are on record showing that a race may be modified by occasional crosses if aided by the careful selection of the individuals which present the desired character. Several other less distinct breeds might be . and it has the habit of continually expanding. Many treatises in different languages have been published on pigeons. Compare the English carrier and the short-faced tumbler. taken up domestic pigeons. The short-faced tumbler has a beak in outline almost like that of a finch. and its enormously developed crop. I have kept every breed which I could purchase or obtain. The diversity of the breeds is something astonishing. elongated wing and tail feathers. and legs. others singularly short tails. The runt is a bird of great size. I have associated with several eminent fanciers. The offspring from the first cross between two pure breeds is tolerably and sometimes (as I have found with pigeons) quite uniform in character. the bull-dog. Moreover. utter a very different coo from the other breeds. slightly. W. hardly two of them are alike. or Blenheim spaniel. The pouter has a much elongated body. Sir J. but by crossing we can only get forms in some degree intermediate between their parents. entailing corresponding differences in their skulls. some of the sub-breeds of runts have very long necks.immense amount of inherited variation. The carrier. etc. The barb is allied to the carrier. I have. we must admit the former existence of the most extreme forms. more especially the male bird. and this is accompanied by greatly elongated eyelids. etc. with a line of reversed feathers down the breast. and see the wonderful difference in their beaks. and then the difficulty of the task becomes manifest. BREEDS OF THE DOMESTIC PIGEON. and the common tumbler has the singular inherited habit of flying at a great height in a compact flock. wings. and a wide gape of mouth. others very long wings and tails. for who will believe that animals closely resembling the Italian greyhound. THEIR DIFFERENCES AND ORIGIN. The Jacobin has the feathers so much reversed along the back of the neck that they form a hood. as their names express. is also remarkable from the wonderful development of the carunculated skin about the head. the upper part of the oesophagus. Murray from Persia. and by the Hon. may well excite astonishment and even laughter. and it has. massive beak and large feet. more especially by the Hon.

the runt. and in some breeds the voice and disposition. As several of the reasons which have led me to this belief are in some degree applicable in other cases. the size of the crop and of the upper part of the oesophagus. The shape and size of the eggs vary. The manner of flight. If the several breeds are not varieties. they must have descended from at least seven or eight aboriginal stocks. which. In the skeletons of the several breeds. could a pouter be produced by crossing two breeds unless one of the parent-stocks possessed the characteristic enormous crop? The supposed aboriginal stocks must all have been rock-pigeons. as he would call them. Hence the supposed aboriginal stocks must either still exist in the countries where they were originally domesticated. and yet be unknown to ornithologists. only two or three other species of rock-pigeons are known. Moreover. which has the same habits with the domestic breeds. as well as the breadth and length of the ramus of the lower jaw. the number of scutellae on the toes. But besides C. or they must have become extinct in the wild state. and these have not any of the characters of the domestic breeds. the males and females have come to differ in a slight degree from each other. Lastly. the number of the primary wing and caudal feathers. I will here briefly give them. as does the number of the ribs. But birds breeding on precipices. namely. which differ from each other in the most trifling respects.specified. The size and shape of the apertures in the sternum are highly variable. together with their relative breadth and the presence of processes. Hence the supposed extermination of so many species having similar habits with . are all points of structure which are variable. The shape. Altogether at least a score of pigeons might be chosen. with its geographical sub-species. could be shown him. more especially as in each of these breeds several truly-inherited sub-breeds. and fantail in the same genus. the development of the bones of the face. and have not proceeded from the rock-pigeon. in length and breadth and curvature. The period at which the perfect plumage is acquired varies. the development of skin between the toes. that all are descended from the rock-pigeon (Columba livia). and the common rock-pigeon. or species. including under this term several geographical races or sub-species. The caudal and sacral vertebrae vary in number. the relative length of the leg and foot. has not been exterminated even on several of the smaller British islets. differs enormously. the proportional length of the eyelids. Great as are the differences between the breeds of the pigeon. livia. the relative length of the wing and tail to each other and to the body. the short-faced tumbler. that is. The proportional width of the gape of mouth. habits and remarkable characters. I am fully convinced that the common opinion of naturalists is correct. varies in a highly remarkable manner. seems improbable. if shown to an ornithologist. as does the state of the down with which the nestling birds are clothed when hatched. so is the degree of divergence and relative size of the two arms of the furcula. differ remarkably. and he were told that they were wild birds. are unlikely to be exterminated. or on the shores of the Mediterranean. they did not breed or willingly perch on trees. and good flyers. and this. would certainly be ranked by him as well-defined species. of the tongue (not always in strict correlation with the length of beak). I do not believe that any ornithologist would in this case place the English carrier. the development and abortion of the oil-gland. for instance. for it is impossible to make the present domestic breeds by the crossing of any lesser number: how. the barb. pouter. of the orifice of the nostrils. considering their size. in certain breeds.

that the above-specified breeds. Some semi-domestic breeds. the several above-named domesticated breeds have been transported to all parts of the world. although no other existing species is thus coloured and marked. which is a white bird with a red tail and red spot on the forehead. The wings have two black bars. though agreeing generally with the wild rock-pigeon in constitution. in every one of the domestic breeds. some of them must have been carried back again into their native country. that these very species have since all become extinct or unknown. and the mongrels were black. voice. Moreover. The rock-pigeon is of a slaty-blue. and in most parts of their structure. C. of generations. but not one has become wild or feral. so that in each separate breed there might be a tendency to revert to the very same colours and markings. but that he intentionally or by chance picked out extraordinarily abnormal species. on the well-known principle of reversion to ancestral characters. if all the domestic breeds are descended from the rock-pigeon. the mongrel offspring are very apt suddenly to acquire these characters. besides the two black bars. not only that half-civilized man succeeded in thoroughly domesticating several species. An argument of great weight. with white loins. for tail-feathers like those of the fantail. double black wing-bar. and applicable in several other cases. and which notoriously breeds very true. habits. or barb. has become feral in several places. have. the wings chequered with black. Now. brown and mottled. though the dovecot-pigeon. even to the white edging of the outer tail-feathers. intermedia of Strickland. therefore. Again. for reversed feathers like those of the Jacobin. Hence it must be assumed. sometimes concur perfectly developed. So many strange contingencies are improbable in the highest degree. all recent experience shows that it is difficult to get wild animals to breed freely under domestication. and further. as any wild rock-pigeon! We can understand these facts. Some facts in regard to the colouring of pigeons well deserve consideration. with some black barbs--and it so happens that blue varieties of barbs are so rare that I never heard of an instance in England. first. it must be assumed that at least seven or eight species were so thoroughly domesticated in ancient times by half-civilized man. or at most within a score. To give one instance out of several which I have observed: I crossed some white fantails. or that of the short-faced tumbler. which breed very true. The tail has a terminal dark bar. been crossed by the rock-pigeon: I say within a dozen . and some truly wild breeds. But if we deny this. for a crop like that of the pouter. that each breed. we may look in vain through the whole great family of Columbidae for a beak like that of the English carrier. all the above marks. I then crossed one of the mongrel barb-fantails with a mongrel barb-spot. when birds belonging to two or more distinct breeds are crossed. none of which are blue or have any of the above-specified marks. with the outer feathers externally edged at the base with white. secondly. that all the several imagined aboriginal stocks were coloured and marked like the rock-pigeon. Moreover. I also crossed a barb with a spot. as to be quite prolific under confinement. even the purest. These several marks do not occur together in any other species of the whole family. the mongrels were dusky and mottled. and barred and white-edged tail-feathers. which is the rock-pigeon in a very slightly altered state. Either. has within a dozen.the rock-pigeon seems a very rash assumption. yet on the hypothesis of the multiple origin of our pigeons. and they produced a bird of as beautiful a blue colour. yet are certainly highly abnormal in other parts. Or. we must make one of the two following highly improbable suppositions. is. taking thoroughly well-bred birds. has this part bluish. and. with the white loins. but the Indian sub-species. colouring.

From these several reasons. we can make. and the explanation of this fact will be obvious when we treat of selection. These two distinct cases of reversion are often confounded together by those who have written on inheritance.. hardly any cases have been ascertained with certainty of hybrids from two quite distinct species of animals being perfectly fertile. as was pointed out to me by Professor Lepsius. both when kept pure and when crossed--and lastly. the earliest known record of pigeons is in the fifth Aegyptian dynasty. pigeons have been watched and tended with the utmost care. Fourthly. as compared with all other Columbidae. if applied to species closely related to each other. the improbability of man having formerly made seven or eight supposed species of pigeons to breed freely under domestication--these supposed species being quite unknown in a wild state. yet that by comparing the several sub-breeds of these two races. as I can state from my own observations. and there is a tendency in the breed to revert to a character which was lost during some former generation. the hybrids or mongrels from between all the breeds of the pigeon are perfectly fertile. so we can in some other cases. more especially those brought from distant countries. and of some other domestic animals. namely. Some authors believe that long-continued domestication eliminates this strong tendency to sterility in species. immense prices were given for pigeons. the mongrel offspring being perfectly fertile--from these several reasons. they are come to this pass. Secondly. but when there has been no cross. we may safely conclude that all our domestic breeds are descended from the rock-pigeon or Columba livia with its geographical sub-species. that although an English carrier or a short-faced tumbler differs immensely in certain characters from the rock-pigeon. and fantails now are. may be transmitted undiminished for an indefinite number of generations. livia has been found capable of domestication in Europe and in India. In a breed which has been crossed only once the tendency to revert to any character derived from such a cross will naturally become less and less. Thirdly. pouters. seems to me rash in the extreme. In the time of the Romans. about 3000 B. But to extend it so far as to suppose that species. the shortness of that of the tumbler.or twenty generations. Lastly. Birch informs me that pigeons are given in a bill of fare in the previous dynasty. and the number of tail-feathers in the fantail. for instance. In favour of this view. but Mr. I may add. the wattle and length of beak of the carrier. on the most distinct breeds. taken together. "nay. this tendency. an almost perfect series. From the history of the dog. as in each succeeding generation there will be less of the foreign blood. removed by a greater number of generations. and that it agrees in habits and in a great number of points of structure with all the domestic breeds. They have been domesticated for thousands of years in several quarters of the world. inter se. for all that we can see to the contrary. and their not having become anywhere feral--these species presenting certain very abnormal characters.C. though so like the rock-pigeon in most other respects--the occasional reappearance of the blue colour and various black marks in all the breeds. that they can reckon up their pedigree and race. those characters which are mainly distinctive of each breed are in each eminently variable. this conclusion is probably quite correct. that the wild C. firstly. should yield offspring perfectly fertile. as we hear from Pliny. tumblers. aboriginally as distinct as carriers. and loved by many people. but not with all the breeds. purposely made. for no instance is known of crossed descendants reverting to an ancestor of foreign blood. between them and the rock-pigeon. Now." Pigeons were much .

Van Mons. or both from a common parent-stock.valued by Akber Khan in India. for instance. either from one or from several allied species. I felt fully as much difficulty in believing that since they had been domesticated they had all proceeded from a common parent. Some variations useful to him have probably arisen suddenly. "The monarchs of Iran and Turan sent him some very rare birds. are firmly convinced that the several breeds to which each has attended. We shall then. not indeed to the animal's or plant's own good. by crossing the breeds. which can not be rivalled by . Let us now briefly consider the steps by which domestic races have been produced. in nature. and though they well know that each race varies slightly. that nearly all the breeders of the various domestic animals and the cultivators of plants. or poultry. when they deride the idea of species in a state of nature being lineal descendants of other species? PRINCIPLES OF SELECTION ANCIENTLY FOLLOWED. and he will laugh you to scorn. see how it is that the several breeds so often have a somewhat monstrous character. as any naturalist could in coming to a similar conclusion in regard to the many species of finches. One circumstance has struck me much. AND THEIR EFFECTS. with its hooks. May not those naturalists who.000 pigeons were taken with the court. never less than 20. but he would be a bold man who would account by such agencies for the differences between a dray and race-horse. with whom I have conversed. but to man's use or fancy. It is also a most favourable circumstance for the production of distinct breeds. about the year 1600. Some effect may be attributed to the direct and definite action of the external conditions of life. yet they ignore all general arguments. I have discussed the probable origin of domestic pigeons at some. well knowing how truly they breed. a greyhound and bloodhound. or other groups of birds. and refuse to sum up in their minds slight differences accumulated during many successive generations. I have never met a pigeon. who was not fully convinced that each main breed was descended from a distinct species. a carrier and tumbler pigeon. believe that the fuller's teasel. shows how utterly he disbelieves that the several sorts. length. has improved them astonishingly. and knowing no more than he does of the intermediate links in the long lines of descent. or duck. as I have asked. whether his cattle might not have descended from Long-horns. and thus different breeds can be kept together in the same aviary. which method was never practised before. will likewise be obvious when we treat of selection. for instance a Ribston-pippin or Codlin-apple. in his treatise on pears and apples. One of the most remarkable features in our domesticated races is that we see in them adaptation. for they win their prizes by selecting such slight differences. or by one step. continues the courtly historian. could ever have proceeded from the seeds of the same tree. a celebrated raiser of Hereford cattle. and some to habit. many botanists. or whose treatises I have read. The paramount importance of these considerations in explaining the immense amount of variation which pigeons have undergone." About this same period the Dutch were as eager about pigeons as were the old Romans. "His Majesty. also. yet admit that many of our domestic races are descended from the same parents--may they not learn a lesson of caution. The explanation. yet quite insufficient. namely. knowing far less of the laws of inheritance than does the breeder. because when I first kept pigeons and watched the several kinds. that male and female pigeons can be easily mated for life. I think." and. Ask. or rabbit fancier. is simple: from long-continued study they are strongly impressed with the differences between the several races. are descended from so many aboriginally distinct species. Innumerable other examples could be given.

But when we compare the dray-horse and race-horse. What English breeders have actually effected is proved by the enormous prices given for animals with a good pedigree. each good for man in different ways. we must. this is done three times at intervals of months. says: "It would seem as if they had chalked out upon a wall a form perfect in itself. which they can model almost as they please. If selection consisted merely in separating some very distinct variety and breeding from it. and the sheep are each time marked and classed. The great power of this principle of selection is not hypothetical. In this sense he may be said to have made for himself useful breeds. like a picture by a connoisseur. culinary. with "everlasting layers" which never desire to sit. I think. modified to a large extent their breeds of cattle and sheep. during successive generations. but to change it altogether. except sometimes among closely allied sub-breeds." Lord Somerville. So it has probably been with the turnspit dog. The key is man's power of accumulative selection: nature gives successive variations. all the best breeders are strongly opposed to this practice. orchard. and with the bantam so small and elegant. that men follow it as a trade: the sheep are placed on a table and are studied. or so beautiful in his eyes. and this is known to have been the case with the ancon sheep. It is certain that several of our eminent breeders have. and flower-garden races of plants. We can not suppose that all the breeds were suddenly produced as perfect and as useful as we now see them. not only to modify the character of his flock. by means of which he may summon into life whatever form and mould he pleases. the dromedary and camel. we know that this has not been their history. even within a single lifetime. Not one man in a thousand has accuracy of eye and judgment sufficient to become an . The improvement is by no means generally due to crossing different breeds. man adds them up in certain directions useful to him. is only a variety of the wild Dipsacus. look further than to mere variability. speaks of the principle of selection as "that which enables the agriculturist. when we compare the game-cock. and these have been exported to almost every quarter of the world. speaking of what breeders have done for sheep. and who was himself a very good judge of animals. and to inspect the animals. but its importance consists in the great effect produced by the accumulation in one direction. and that of another breed for another purpose. so pertinacious in battle. the various breeds of sheep fitted either for cultivated land or mountain pasture. If I had space I could quote numerous passages to this effect from highly competent authorities. And when a cross has been made. and this amount of change may have suddenly arisen in a seedling. with the wool of one breed good for one purpose. It is the magician's wand. so that the very best may ultimately be selected for breeding. Breeders habitually speak of an animal's organisation as something plastic. and then had given it existence. In order fully to realise what they have done it is almost necessary to read several of the many treatises devoted to this subject. most useful to man at different seasons and for different purposes." In Saxony the importance of the principle of selection in regard to merino sheep is so fully recognised.any mechanical contrivance. in many cases. when we compare the many breeds of dogs. indeed. Youatt. with other breeds so little quarrelsome. the closest selection is far more indispensable even than in ordinary cases. the principle would be so obvious as hardly to be worth notice. who was probably better acquainted with the works of agriculturalists than almost any other individual. when we compare the host of agricultural. of differences absolutely inappreciable by an uneducated eye--differences which I for one have vainly attempted to appreciate.

When a race of plants is once pretty well established. and how alike the leaves. and the result has been. or tubers. by comparing the diversity of flowers in the different varieties of the same species in the flower-garden. and they formerly did so. it has certainly been more attended to of late years. how unlike the flowers of the heartsease are. With animals this kind of selection is. it cannot be doubted that the continued selection of slight variations. there is another means of observing the accumulated effects of selection--namely. how much the fruit of the different kinds of gooseberries differ in size. I could give several references to works of high antiquity. but the variations are here often more abrupt. likewise followed. rapid and important. for hardly any one is so careless as to breed from his worst animals. in fact. when the flowers of the present day are compared with drawings made only twenty or thirty years ago. the importance of which should never be overlooked. to give a very trifling instance. and pull up the "rogues. the diversity of leaves. the seed-raisers do not pick out the best plants. Few would readily believe in the natural capacity and years of practice requisite to become even a skilful pigeon-fancier. In rude and barbarous periods of English history choice animals were often imported. and many treatises have been published on the subject. in comparison with the leaves and flowers of the same set of varieties. and how extremely alike the flowers. and he studies his subject for years. he will succeed.eminent breeder. The principle of selection I find distinctly given in an ancient Chinese encyclopaedia. The law of correlated variation. the steadily increasing size of the common gooseberry may be quoted. it is clear that the colour of domestic animals was at that early period attended to. thus. and may make great improvements. The same principles are followed by horticulturists. either in the leaves. We have proofs that this is not so in several cases in which exact records have been kept. See how different the leaves of the cabbage are. and laws were passed to prevent their exportation: the destruction of horses under a certain size was ordered." as they call the plants that deviate from the proper standard. this is hardly ever--I speak after careful observation--perhaps never. as a general rule. If gifted with these qualities. the case. but merely go over their seed-beds. and hairiness. The savages in South Africa match their draught cattle by . he will assuredly fail. if he wants any of these qualities. and devotes his lifetime to it with indomitable perseverance. or the fruit. Explicit rules are laid down by some of the Roman classical writers. pods. and this may be compared to the "roguing" of plants by nurserymen. in the kitchen-garden. We see an astonishing improvement in many florists' flowers. in a corresponding degree. to improve the breed. or whatever part is valued. but. as is attested by passages in Pliny. and yet the flowers present very slight differences. shape. and the diversity of fruit of the same species in the orchard. in which the full importance of the principle is acknowledged. No one supposes that our choicest productions have been produced by a single variation from the aboriginal stock. will ensure some differences. in comparison with the flowers of the same varieties. It may be objected that the principle of selection has been reduced to methodical practice for scarcely more than three-quarters of a century. It is not that the varieties which differ largely in some one point do not differ at all in other points. Savages now sometimes cross their dogs with wild canine animals. In regard to plants. the flowers. colour. But it is very far from true that the principle is a modern discovery. will produce races differing from each other chiefly in these characters. From passages in Genesis.

By comparing the accounts given in various old treatises of the former and present state of carrier and tumbler pigeons in Britain. a form of selection. The two flocks of Leicester sheep kept by Mr. and in this case the change has.. would improve and modify any breed. or but little changed. the forms and qualities of their cattle. By a similar process of selection. as do some of the Esquimaux their teams of dogs. are favoured in the weights which they carry. and which results from every one trying to possess and breed from the best individual animals. At the present time. It would. and afterwards breeds from his own best dogs. by the regulations for the Goodwood Races. Buckley and Mr. the production of the distinct strains. where the breed has been less improved. There is reason to believe that King Charles' spaniel has been unconsciously modified to a large extent since the time of that monarch. for our purpose. Burgess. Borrow has not seen. and Persia. and by careful training.colour. and come to differ so greatly from the rock-pigeon. however. English race-horses have come to surpass in fleetness and size the parent Arabs. Bakewell for upwards of fifty years. individuals of the same breed exist in less civilised districts. that the change has been effected unconsciously and gradually. but he has no wish or expectation of permanently altering the breed. continued during centuries. It is known that the English pointer has been greatly changed within the last century. with a distinct object in view. a man who intends keeping pointers naturally tries to get as good dogs as he can. in so far that the breeders could never have expected. Lord Spencer and others have shown how the cattle of England have increased in weight and in early maturity. is more important. we can trace the stages through which they have insensibly passed. etc. Youatt remarks. for the inheritance of good and bad qualities is so obvious. In some cases. and yet so effectually that. so that the latter. superior to anything of the kind in the country. had attention not been paid to breeding. Some highly competent authorities are convinced that the setter is directly derived from the spaniel. even during their lifetimes. which may serve for comparison. or even wished. only carried on more methodically. by this very same process. which may be called unconscious. Some of these facts do not show actual selection. been chiefly effected by crosses with the foxhound. Thus. and is now attended to by the lowest savages. "Have been purely bred from the original stock of Mr. There is not a suspicion existing in the mind of any one at all acquainted with the subject that . though the old Spanish pointer certainly came from Spain. Mr. Nevertheless we may infer that this process. to produce the result which ensued--namely. as I am informed by him. Livingstone states that good domestic breeds are highly valued by the negroes in the interior of Africa who have not associated with Europeans. compared with the stock formerly kept in this country. Collins. and has probably been slowly altered from it. but what concerns us is. India. Youatt gives an excellent illustration of the effects of a course of selection which may be considered as unconscious. unchanged. eminent breeders try by methodical selection. as Mr. any native dog in Spain like our pointer. it is believed. But. to make a new strain or sub-breed. Slow and insensible changes of this kind could never be recognised unless actual measurements or careful drawings of the breeds in question have been made long ago. have been a strange fact. did greatly modify. UNCONSCIOUS SELECTION. indeed. in the same way as Bakewell. but they show that the breeding of domestic animals was carefully attended to in ancient times.

and therefore do not know. the wild parent-stocks of the plants which have been longest cultivated in our flower and kitchen gardens. as of less value than their dogs. it should not be overlooked that they almost always have to struggle for their own food. for any special purpose. pelargonium. and. who cultivated the best pears which they could procure. and yet the difference between the sheep possessed by these two gentlemen is so great that they have the appearance of being quite different varieties. from Pliny's description. having . if it had come from a garden-stock. thus slowly and unconsciously accumulated.the owner of either of them has deviated in any one instance from the pure blood of Mr. though we owe our excellent fruit in some small degree to their having naturally chosen and preserved the best varieties they could anywhere find. so that in this case there would be a kind of unconscious selection going on. never thought what splendid fruit we should eat. No one would expect to raise a first-rate melting pear from the seed of a wild pear. Bakewell's flock. though cultivated in classical times. in times of dearth. and other plants. by their killing and devouring their old women. as far as the final result is concerned. the Cape of Good Hope. But the gardeners of the classical period. to have been a fruit of very inferior quality. and such choice animals would thus generally leave more offspring than the inferior ones. whether or not sufficiently distinct to be ranked at their first appearance as distinct varieties. the well-known fact. when compared with the older varieties or with their parent-stocks. And in two countries very differently circumstanced. nor any other region inhabited by quite uncivilised man. In plants the same gradual process of improvement through the occasional preservation of the best individuals. when a slightly better variety chanced to appear." If there exist savages so barbarous as never to think of the inherited character of the offspring of their domestic animals. No one would ever expect to get a first-rate heartsease or dahlia from the seed of a wild plant. yet any one animal particularly useful to them. though he might succeed from a poor seedling growing wild. at least during certain seasons. dahlia. appears. but that the native plants have not been improved by continued selection up to a standard of perfection comparable with that acquired by the plants in countries anciently civilised. It has consisted in always cultivating the best known variety. If it has taken centuries or thousands of years to improve or modify most of our plants up to their present standard of usefulness to man. selecting it. I have seen great surprise expressed in horticultural works at the wonderful skill of gardeners in having produced such splendid results from such poor materials. In regard to the domestic animals kept by uncivilised man. as I believe. may plainly be recognised in the increased size and beauty which we now see in the varieties of the heartsease. that in a number of cases we cannot recognise. do not by a strange chance possess the aboriginal stocks of any useful plants. has been followed almost unconsciously. would be carefully preserved during famines and other accidents. explains. has afforded us a single plant worth culture. but the art has been simple. The pear. A large amount of change. rose. sowing its seeds. individuals of the same species. and so onwards. We see the value set on animals even by the barbarians of Tierra del Fuego. and. It is not that these countries. to which savages are so liable. and whether or not two or more species or races have become blended together by crossing. we can understand how it is that neither Australia. so rich in species.

I have no doubt. further understand the frequently abnormal character of our domestic races. two sub-breeds might be formed. however slight." as will hereafter be more fully explained. and from being only slightly valued. and thus by a process of "natural selection. the more likely it would be to catch his attention. or takes more care than usual in matching his best animals. any deviation of structure excepting such as is externally visible. hence the Toulouse and the common breed. would often succeed better in the one country than in the other. We can. and the more abnormal or unusual any character was when it first appeared. which differ only in colour. and likewise their differences being so great in external characters. after several breeds have fairly been established. and thus improves them. in which as many as seventeen tail-feathers have been counted. how it is that our domestic races show adaptation in their structure or in their habits to man's wants or fancies. and the improved animals slowly spread in the immediate neighbourhood. like a dialect of a language. utterly incorrect. in most cases. selection. namely. like the present Java fantail. and will be recognised as something . in fact. or like individuals of other and distinct breeds. never dreamed what the descendants of that pigeon would become through long-continued. Perhaps the parent bird of all fantails had only fourteen tail-feathers somewhat expanded. can hardly be said to have a distinct origin. and relatively so slight in internal parts or organs. The common goose has not given rise to any marked varieties. No man would ever try to make a fantail till he saw a pigeon with a tail developed in some slight degree in an unusual manner. as it is not one of the points of the breed. they will spread more widely. The man who first selected a pigeon with a slightly larger tail.slightly different constitutions or structure. that most fleeting of characters. a breed. It is known that with pigeons many slight variations now occasionally appear. partly explains why the varieties kept by savages. in one's own possession. have lately been exhibited as distinct at our poultry-shows. or only with much difficulty. it becomes at once obvious. Nor let it be thought that some great deviation of structure would be necessary to catch the fancier's eye: he perceives extremely small differences. Perhaps the first pouter-pigeon did not inflate its crop much more than the turbit now does the upper part of its oesophagus--a habit which is disregarded by all fanciers. I think. When further improved by the same slow and gradual process. have more of the character of true species than the varieties kept in civilised countries. perhaps. but these are rejected as faults or deviations from the standard of perfection in each breed. as has been remarked by some authors. and indeed he rarely cares for what is internal. be judged of by the value which is now set on them. But they will as yet hardly have a distinct name. or a pouter till he saw a pigeon with a crop of somewhat unusual size. excepting on variations which are first given to him in some slight degree by nature. their history will have been disregarded. He can never act by selection. But. A man preserves and breeds from an individual with some slight deviation of structure. and it is in human nature to value any novelty. This. On the view here given of the important part which selection by man has played. Man can hardly select. These views appear to explain what has sometimes been noticed. But to use such an expression as trying to make a fantail is. partly unconscious and partly methodical. Nor must the value which would formerly have been set on any slight differences in the individuals of the same species. that we know hardly anything about the origin or history of any of our domestic breeds.

according to the state of civilisation of the inhabitants--slowly to add to the characteristic features of the breed. the spreading of a new sub-breed will be a slow process. that it was most fortunate that the strawberry began to vary just when gardeners began to attend to this plant. as the breed rises or falls in fashion--perhaps more in one district than in another. But the chance will be infinitely small of any record having been preserved of such slow. although so much valued by women and children. No doubt the strawberry had always varied since it was cultivated. such breeds as we do sometimes see are almost always imported from some other . Unless such attention be paid nothing can be effected. the chance of their appearance will be much increased by a large number of individuals being kept. Wandering savages or the inhabitants of open plains rarely possess more than one breed of the same species. In this respect enclosure of the land plays a part. and raised seedlings from them. they never can be improved. for thus many races may be improved and kept true. and again picked out the best seedlings and bred from them. facility in preventing crosses is an important element in the formation of new races--at least. On this principle Marshall formerly remarked. with respect to the sheep of part of Yorkshire. nurserymen. "As they generally belong to poor people. When the individuals are scanty all will be allowed to breed. however. A large number of individuals of an animal or plant can be reared only where the conditions for its propagation are favourable. and insensible changes. as freely giving the materials for selection to work on. Pigeons can be mated for life. I have seen it gravely remarked.distinct and valuable. as gardeners picked out individual plants with slightly larger. cats. On the other hand. As soon as the points of value are once acknowledged. to man's power of selection. Hence number is of the highest importance for success. from their nocturnal rambling habits. whatever they may be. CIRCUMSTANCES FAVOURABLE TO MAN'S POWER OF SELECTION. but the slight varieties had been neglected. and are mostly IN SMALL LOTS. But as variations manifestly useful or pleasing to man appear only occasionally. in a country which is already stocked with other races. Pigeons. with little free communication. from keeping large stocks of the same plant. A high degree of variability is obviously favourable. or better fruit. to allow of the accumulation of a large amount of modification in almost any desired direction. I may add. are generally far more successful than amateurs in raising new and valuable varieties. and this is a great convenience to the fancier. varying. and this circumstance must have largely favoured the formation of new breeds. can not be easily matched. and this will effectually prevent selection. we rarely see a distinct breed long kept up. can be propagated in great numbers and at a very quick rate." On the other hand. But probably the most important element is that the animal or plant should be so highly valued by man. favourable or the reverse. With animals. that the closest attention is paid to even the slightest deviations in its qualities or structure. as I have called it. then (with some aid by crossing distinct species) those many admirable varieties of the strawberry were raised which have appeared during the last half-century. with extreme care. and will then probably first receive a provincial name. I will now say a few words on the circumstances. the principle. In semi-civilised countries. of unconscious selection will always tend--perhaps more at one period than at another. though mingled in the same aviary. earlier. and. and inferior birds may be freely rejected. As soon. not that mere individual differences are not amply sufficient. whatever their quality may be. as when killed they serve for food.

food and feathers. Variability is governed by many unknown laws. effect may be attributed to the increased use or disuse of parts. and this implies variation. When several breeds have once been formed in any country. in donkeys. their occasional intercrossing. No doubt. Some. after remaining fixed for many centuries. Wallace has remarked with much truth. For instance. in peacocks. from not being very easily reared and a large stock not kept. from only a few being kept by poor people. may be attributed to the definite action of the conditions of life. But what concerns us is that the domestic varieties of the same species differ from each other in almost every character. under all circumstances. and probably with the length of hair. than any two natural species belonging to the same genus. under the conditions to which it is exposed when domesticated. may be attributed in main part to selection not having been brought into play: in cats. Isidore Geoffroy St. as this will be determined by the friction to be overcome. as I have elsewhere described. Changed conditions of life are of the highest importance in causing variability. With respect to fleetness. The final result is thus rendered infinitely complex. a limit will be at last reached. no doubt. there must be a limit to the fleetness of any terrestrial animal. goose. for recently in certain parts of Spain and of the United States this animal has been surprisingly modified and improved by careful selection. from being valuable only for two purposes. of which correlated growth is probably the most important. and so it is with colour. Hilaire has proved this in regard to size. and the power of contraction in the muscular fibres. etc. for almost all our animals and plants have been greatly improved in many ways within a recent period. again vary under new conditions of life. The greater or less force of inheritance and reversion determine whether variations shall endure. Some authors have maintained that the amount of variation in our domestic productions is soon reached. yet the rarity or absence of distinct breeds of the cat. The same remark holds good in regard to the fruit of the several varieties of the plum. Eclipse was far fleeter. in geese. peacock. but how much we do not know. the weight of the body to be carried. could not. and still more strongly with the melon. largely aided in the formation of new sub-breeds. has. and more especially from no pleasure having been felt in the display of distinct breeds. In some cases the intercrossing of aboriginally distinct species appears to have played an important part in the origin of our breeds. Although I do not doubt that some domestic animals vary less than others. seems to have a singularly inflexible organisation.. more than do the distinct species of the same genera. It would be equally rash to assert that characters now increased to their utmost limit. and can never afterward be exceeded. with the aid of selection. and little attention paid to their breeding. perhaps a great. So with plants.country. as well as in many other analogous cases. and indirectly by affecting the reproductive system. but the importance of crossing has been much exaggerated. which depends on many bodily characters. the seeds of the different varieties of the bean or maize probably differ more in size than do the seeds of the distinct species in any one genus in the same two families. which man has attended to and selected. from the difficulty in pairing them. It would be somewhat rash to assert that the limit has been attained in any one case. both by acting directly on the organisation. and a dray-horse is comparably stronger. It is not probable that variability is an inherent and necessary contingent. but the goose. both in regard to animals and to those plants which are . as Mr. To sum up on the origin of our domestic races of animals and plants. though it has varied to a slight extent. Something. the donkey.

the accumulative action of selection. yet every naturalist knows vaguely what he means when he speaks of a species. It may be doubted whether sudden and considerable deviations of structure. we must briefly discuss whether these latter are subject to any variation. but here community of descent is almost universally implied. Almost every part of every organic being is so beautifully related to its complex conditions of life that it seems as improbable that any part should have been suddenly produced perfect. but plants not propagated by seed are of little importance to us. We have also what are called monstrosities. cases of monstrosities resembling normal structures in nearly allied forms. VARIATION UNDER NATURE. or dwarfed plants on Alpine summits. a long catalogue of dry facts ought to be given. for their endurance is only temporary. and these alone bear on the question. related to each other. and if any wild species of the same genus had naturally possessed a proboscis. after diligent search. or not useful to the species. generally injurious. such as we occasionally see in our domestic productions. or unconsciously and slowly. Some authors use the term "variation" in a technical sense. seems to have been the predominant power. though it can rarely be proved. buds. but they graduate into varieties. The term "variety" is almost equally difficult to define. No one definition has satisfied all naturalists. more especially with plants.propagated by seed. and common species. Nor shall I here discuss the various definitions which have been given of the term species. or the thicker fur of an animal from far northwards. as implying a modification directly due to the physical conditions of life. are ever permanently propagated in a state of nature. but I have as yet failed to find. whether applied methodically and quickly. and "variations" in this sense are supposed not to be inherited. the importance of crossing is immense. Before applying the principles arrived at in the last chapter to organic beings in a state of nature. but who can say that the dwarfed condition of shells in the brackish waters of the Baltic. as that a complex machine should have been invented by man in a perfect state. it might have been argued that this had appeared as a monstrosity. but these I shall reserve for a future work. for the cultivator may here disregard the extreme variability both of hybrids and of mongrels. would not in some cases be inherited for at least a few generations? And in this case I presume that the form would be called a variety. Thus pigs have occasionally been born with a sort of proboscis. Under domestication monstrosities sometimes occur which resemble normal structures in widely different animals. but unequally. and the sterility of hybrids. etc.. vary most--Species of the larger genera in each country vary more frequently than the species of the smaller genera--Many of the species of the larger genera resemble varieties in being very closely. Variability--Individual differences--Doubtful species--Wide ranging. Over all these causes of change. With plants which are temporarily propagated by cuttings. much diffused. By a monstrosity I presume is meant some considerable deviation of structure. but more efficiently. CHAPTER II. and in having restricted ranges. Generally the term includes the unknown element of a distinct act of creation. To treat this subject properly. If monstrous .

and likewise. Lubbock has shown a degree of variability in these main nerves in Coccus. In most polymorphic genera some of the species have fixed and definite characters. With respect to many of these forms. that parts which must be called important. for these same authors practically rank those parts as important (as some few naturalists have honestly confessed) which do not vary. and. and compare them in many specimens of the same species.forms of this kind ever do appear in a state of nature and are capable of reproduction (which is not always the case). but under any other point of view many instances assuredly can be given. But I shall have to return in a future chapter to the preservation and perpetuation of single or occasional variations. These facts are very perplexing. as they occur rarely and singly. even in important parts of structure. whether viewed under a physiological or classificatory point of view. may be called individual differences. This philosophical naturalist. It should be remembered that systematists are far from being pleased at finding variability in important characters. and that there are not many men who will laboriously examine internal and important organs. or which it may be presumed have thus arisen. and thus their abnormal character would almost inevitably be lost. and of Brachiopod shells. but I could show. Genera which are polymorphic in one country seem to be. no instance will ever be found of an important part varying. polymorphic in other countries. judging from Brachiopod shells. with a few exceptions. for they are often inherited. under this point of view. as must be familiar to every one. INDIVIDUAL DIFFERENCES. has also shown that the muscles in the larvae of certain insects are far from uniform. several genera of insects. yet Sir J. sometimes vary in the individuals of the same species. and they thus afford materials for natural selection to act on and accumulate. and Hieracium among plants. at former periods of time. We may instance Rubus. hardly two naturalists agree whether to rank them as species or as varieties. during a course of years. by a long catalogue of facts. Authors sometimes argue in a circle when they state that important organs never vary. from being observed in the individuals of the same species inhabiting the same confined locality. for they seem to show that this kind of variability is independent of the . in the same manner as man accumulates in any given direction individual differences in his domesticated productions. during the first and succeeding generations cross with the ordinary form." in which species present an inordinate amount of variation. The many slight differences which appear in the offspring from the same parents. These individual differences generally affect what naturalists consider unimportant parts. I am convinced that the most experienced naturalist would be surprised at the number of the cases of variability. They would. I may add. These individual differences are of the highest importance for us. also. which he could collect on good authority. It would never have been expected that the branching of the main nerves close to the great central ganglion of an insect would have been variable in the same species. it might have been thought that changes of this nature could have been effected only by slow degrees. their preservation would depend on unusually favourable circumstances. Rosa. There is one point connected with individual differences which is extremely perplexing: I refer to those genera which have been called "protean" or "polymorphic. which may almost be compared to the irregular branching of the stem of a tree. No one supposes that all the individuals of the same species are cast in the same actual mould. as I have collected.

as I have myself observed. and the other has antennae much more abundantly furnished with smelling-hairs. Practically. the male of a Tanais regularly occurs under two distinct forms. the castes are connected together by finely graduated varieties. it is possible that they were once thus connected. as we shall hereafter see. at least in some of these polymorphic genera. I am inclined to suspect that we see. as far as we know. Thus also with ants. both with animals and plants. but because analogy leads the observer to suppose either that they do . in the Malayan Archipelago. independently of variation. Wallace. the two or three forms. one of these has strong and differently shaped pincers. regularly appear under two or even three conspicuously distinct forms. nor will the commonly assumed hybrid nature of the intermediate forms always remove the difficulty. also. that naturalists do not like to rank them as distinct species. Mr. But cases of great difficulty. and in the immature and larval states of many of the lower animals. as in the two sexes of various animals. Thus. Nevertheless these cases are only exaggerations of the common fact that the female produces offspring of two sexes which sometimes differ from each other in a wonderful manner. but in some cases. has shown that the females of certain species of butterflies.conditions of life. for instance. as have good and true species. even when they are closely connected by intermediate links. DOUBTFUL SPECIES. Fritz Muller has described analogous but more extraordinary cases with the males of certain Brazilian Crustaceans: thus. which I will not here enumerate. not because the intermediate links have actually been found. when a naturalist can unite by means of intermediate links any two forms. he treats the one as a variety of the other. describes a certain butterfly which presents in the same island a great range of varieties connected by intermediate links. are in several respects the most important for us. in the two or three castes of sterile females or workers among insects. sometimes arise in deciding whether or not to rank one form as a variety of another. and the extreme links of the chain closely resemble the two forms of an allied dimorphic species inhabiting another part of the Malay Archipelago. are not now connected by intermediate gradations. In very many cases. one form is ranked as a variety of another. We have every reason to believe that many of these doubtful and closely allied forms have permanently retained their characters for a long time. The forms which possess in some considerable degree the character of species. cases of dimorphism and trimorphism. Wallace. and the other as the variety. and which consequently have not been seized on and rendered definite by natural selection. It certainly at first appears a highly remarkable fact that the same female butterfly should have the power of producing at the same time three distinct female forms and a male. Although in most of these cases. but which are so closely similar to other forms. both with animals and plants. and that an hermaphrodite plant should produce from the same seed-capsule three distinct hermaphrodite forms. for as long. not connected by intermediate varieties. Mr. the several worker-castes are generally quite distinct. however. variations which are of no service or disservice to the species. as hereafter to be explained. or are so closely linked to them by intermediate gradations. but sometimes the one first described as the species. There are. as is known to every one. great differences of structure. Individuals of the same species often present. bearing three different kinds of females and three or even six different kinds of males. who has lately called attention to the subject. So it is. with some dimorphic plants. ranking the most common.

"There is no possible test but individual opinion to determine which of them shall be considered as species and which as varieties. The local forms are moderately constant and distinct in each separate island. Mr. geographical races! Mr. have been ranked by one eminent naturalist as undoubted species. however. Watson. decide by a majority of naturalists. or may formerly have existed. and by another as mere varieties. but as they do not differ from each other by strongly marked and important characters. and see what a surprising number of forms have been ranked by one botanist as good species. inhabiting the islands of the great Malayan Archipelago. the birds from the closely neighbouring islands of the Galapagos Archipelago. and which are highly locomotive. as variable forms. doubtful forms. On the islets of the little Madeira group there are many insects which are characterized as varieties in Mr. no certain criterion can possibly be given by which variable forms. Bentham gives only 112--a difference of 139 doubtful forms! Among animals which unite for each birth. of France. especially on the Lepidoptera. The first or variable forms vary much within the limits of the same island. representative species fill the same place in the natural economy of each island as do the local forms and sub-species. Compare the several floras of Great Britain. whereas Mr." Lastly. and as true representative species. sub species and representative species can be recognised. in determining whether a form should be ranked as a species or a variety. and in making this list he has omitted many trifling varieties. Wallace. or of the United States. but as they are distinguished from each other by a greater amount of difference than that between the local forms and sub-species. one with another. Babington gives 251 species. local forms. Nevertheless. which differ very slightly from each other. the opinion of naturalists having sound judgment and wide experience seems the only guide to follow. has marked for me 182 British plants. to whom I lie under deep obligation for assistance of all kinds. when comparing. The geographical races or sub-species are local forms completely fixed and isolated. ranked by one zoologist as a species and by another as a variety. and here a wide door for the entry of doubt and conjecture is opened. Mr. which are generally considered as varieties. Many years ago.now somewhere exist. namely. That varieties of this doubtful nature are far from uncommon cannot be disputed. but which have all been ranked by botanists as species.C. and he has entirely omitted several highly polymorphic genera. can rarely be found within the same country. How many of the birds and insects in North America and Europe. but which would certainly be ranked as distinct species . they are almost universally ranked by naturalists as true species. as geographical races or sub-species. I was much struck how entirely vague and arbitrary is the distinction between species and varieties. shows that they may be classed under four heads. Wollaston's admirable work. H. and with those from the American mainland. drawn up by different botanists. or. and seeing others compare. and by another as varieties. but when all from the several islands are compared together. but are common in separated areas. in many cases. Under genera. the differences are seen to be so slight and graduated that it is impossible to define or describe them. though at the same time the extreme forms are sufficiently distinct. for few well-marked and well-known varieties can be named which have not been ranked as species by at least some competent judges. including the most polymorphic forms. in several valuable papers on the various animals. We must. but which nevertheless have been ranked by some botanists as species. Hence. as local forms. as they are often called.

resemble species so completely in character that they have been thus ranked by other highly competent judges. which of these Phytophagic forms ought to be called species and which varieties. When. in many cases. will that between Europe and the Azores. A wide distance between the homes of two doubtful forms leads many naturalists to rank them as distinct species. or the Canaries. and those which appear to have lost this power. This likewise necessarily occurs with closely allied organisms. Close investigation. from geographical distribution. analogical variation. it has been well asked. But no observer can determine for another. the forms are ranked by all entomologists as good species. both males and females. or inhabits many islands in the same archipelago. but do not in consequence vary. It is certain that many forms. now generally regarded as varieties. but what distance. whereas the greater number rank it as an undoubted species peculiar to Great Britain. Even Ireland has a few animals. as varieties. there is always a good chance that intermediate forms will be discovered which will link together the extreme states. and when the same identical form is met with in two distant countries. etc. or between the several islets of these small archipelagos. though constant differences in colour. a distinguished entomologist of the United States. on the other hand. it cannot be expected that intermediate links connecting the several forms should now be found.D. for several interesting lines of argument. Most vegetable-feeding insects live on one kind of plant or on one group of plants.. but which have been ranked as species by some zoologists. Many of the cases of strongly marked varieties or doubtful species well deserve consideration. which inhabit distinct continents or islands. considered by highly competent judges to be varieties. When the differences are rather more strongly marked. or in both states. Walsh. in other instances. will suffice if that between America and Europe is ample. Mr. have been observed thus to differ in a slight degree. or Madeira. In some instances the males alone. In several cases. or in the nature of their secretions. and when both sexes and all ages are affected. even if he can do so for himself. The naturalist thus loses his best guide in determining whether to rank doubtful forms as varieties or species. before any definition of these terms has been generally accepted. size. Walsh to present in their larval or mature state. hybridism. will no doubt bring naturalists to agree how to rank doubtful . however. slight. have been brought to bear in the attempt to determine their rank. is vainly to beat the air. But to discuss whether they ought to be called species or varieties. The term species thus comes to be a mere useless abstraction. Some few naturalists maintain that animals never present varieties. and presents different forms in the different areas. they believe that two distinct species are hidden under the same dress. As the differences depend on the insects having long fed on distinct plants. implying and assuming a separate act of creation. as species. but space does not here permit me to discuss them. Several experienced ornithologists consider our British red grouse as only a strongly marked race of a Norwegian species. Walsh ranks the forms which it may be supposed would freely intercross. some feed indiscriminately on many kinds. has described what he calls Phytophagic varieties and Phytophagic species. be sufficient? Mr. and these are then degraded to the rank of varieties. but then these same naturalists rank the slightest difference as of specific value. or in two geological formations.by many entomologists. B. an animal or plant ranges over the same continent. insects found living on different plants. have been observed by Mr.

for he will . as Asa Gray again remarks. I may here allude to a remarkable memoir lately published by A. His general tendency will be to make many species. if these connecting forms which are now rare were to become totally extinct the three sub-species would hold exactly the same relation to each other as do the four or five provisionally admitted species which closely surround the typical Quercus robur. namely Q. and this shows. But if he confine his attention to one class within one country he will soon make up his mind how to rank most of the doubtful forms. Thus Quercus robur has twenty-eight varieties. "and the most accordant with the known facts in palaeontology. how closely it has been studied. so long as a genus was imperfectly known. but concludes that the derivative theory is the most natural one." When a young naturalist commences the study of a group of organisms quite unknown to him he is at first much perplexed in determining what differences to consider as specific and what as varietal. He first gives in detail all the many points of structure which vary in the several species.forms. De Candolle admits that out of the 300 species. This seemed to be true. he emphatically remarks: "They are mistaken. sessiliflora and pubescens." He also adds that it is the best known species which present the greatest number of spontaneous varieties and sub-varieties. that is. Such characters are not of course of specific value. No one ever had more ample materials for the discrimination of the species. and estimates numerically the relative frequency of the variations. at least. at least two-thirds are provisional species. how very generally there is some variation. geographical botany and zoology. Finally. and never found connected by intermediate states. sometimes according to age or development. were provisional. pedunculata. which will be enumerated in his Prodromus as belonging to the oak family. Just as we come to know them better. on the oaks of the whole world. of anatomical structure and classification. or from any cause closely attracts his attention. moreover. which are almost universally considered by other botanists to be varieties. such as generally enter into specific definitions. as Asa Gray has remarked in commenting on this memoir. He specifies above a dozen characters which may be found varying even on the same branch. are not known strictly to fulfil the definition above given of a true species. for he knows nothing of the amount and kind of variation to which the group is subject. excepting six. are clustered round three sub-species. will often be ranked by some authors as species. I have been struck with the fact that if any animal or plant in a state of nature be highly useful to man. Yet it must be confessed that it is in the best known countries that we find the greatest number of them. and doubts as to specific limits augment. varieties of it will almost universally be found recorded. all of which. and its species were founded upon a few specimens. sometimes without any assignable reason. who repeat that the greater part of our species are clearly limited. and. de Candolle. De Candolle then goes on to say that he gives the rank of species to the forms that differ by characters never varying on the same tree. These varieties. and that the doubtful species are in a feeble minority. but they are. or could have worked on them with more zeal and sagacity. After this discussion. Look at the common oak. the result of so much labour. intermediate forms flow in. It should be added that De Candolle no longer believes that species are immutable creations. that is to say. The forms which connect these three sub-species are comparatively rare. and in this country the highest botanical authorities and practical men can be quoted to show that the sessile and pedunculated oaks are either good and distinct species or mere varieties. yet a German author makes more than a dozen species out of forms.

Certainly no clear line of demarcation has as yet been drawn between species and sub-species--that is. but do not quite arrive at. it would then rank as the species. or between lesser varieties and individual differences. for convenience sake. which is given to less distinct and more fluctuating forms. They may become extinct. Wollaston with the varieties of certain fossil land-shells in Madeira. with the amount of difference in the forms which he is continually studying. is also applied arbitrarily. just like the pigeon or poultry fancier before alluded to. When he comes to study allied forms brought from countries not now continuous. as steps toward more strongly marked and permanent varieties. in which case he cannot hope to find intermediate links. A well-marked variety may therefore be called an incipient species. or it might come to supplant and exterminate the parent species. and the species as the variety. and to the effects of the increased use or disuse of parts. again. and the truth of this admission will often be disputed by other naturalists. or. but with respect to the more important and adaptive characters. and then to species. But we shall hereafter return to this subject. These differences blend into each other by an insensible series. again. and at the latter. he will be compelled to trust almost entirely to analogy. the forms which in the opinion of some naturalists come very near to. But if his observations be widely extended he will in the end generally be able to make up his own mind. and a series impresses the mind with the idea of an actual passage. but he will succeed in this at the expense of admitting much variation. From these remarks it will be seen that I look at the term species as one arbitrarily given.become impressed. The passage from one stage of difference to another may. The term variety. Hence I look at individual differences. though of small interest to the systematist. in many cases. or they may endure as varieties for very long periods. as leading to sub-species. the rank of species. . but whether this belief is justifiable must be judged by the weight of the various facts and considerations to be given throughout this work. and with plants by Gaston de Saporta. be the simple result of the nature of the organism and of the different physical conditions to which it has long been exposed. hereafter to be explained. to a set of individuals closely resembling each other. and both rank as independent species. It need not be supposed that all varieties or incipient species attain the rank of species. for the sake of convenience. the passage from one stage of difference to another may be safely attributed to the cumulative action of natural selection. or both might co-exist. in comparison with mere individual differences. If a variety were to flourish so as to exceed in numbers the parent species. for he will encounter a greater number of closely-allied forms. and he has little general knowledge of analogical variation in other groups and in other countries by which to correct his first impressions. And I look at varieties which are in any degree more distinct and permanent. and his difficulties will rise to a climax. and that it does not essentially differ from the term variety. as being the first step towards such slight varieties as are barely thought worth recording in works on natural history. as of the highest importance for us. between sub-species and well-marked varieties. as has been shown to be the case by Mr. As he extends the range of his observations he will meet with more cases of difficulty.

as did subsequently Dr. the former will be found to include a .C. In these remarks on predominence. he thinks that the following statements are fairly well established. by tabulating all the varieties in several well-worked floras. AND COMMON SPECIES VARY MOST. as varieties. it should be understood that reference is made only to the forms which come into competition with each other. is rather perplexing. Guided by theoretical considerations. that is abound most in individuals. in order to become in any degree permanent. the species which are already dominant will be the most likely to yield offspring. and more especially to the members of the same genus or class having nearly similar habits of life." "divergence of character. Watson. but Mr. I shall reserve for a future work the discussion of these difficulties. and more widely diffused. I thought that some interesting results might be obtained in regard to the nature and relations of the species which vary most. might have been anticipated. incipient species. all those in the larger genera (i. A plant of this kind is not the less dominant because some conferva inhabiting the water or some parasitic fungus is infinitely more numerous in individuals. Dr. oftenest give rise to varieties sufficiently well-marked to have been recorded in botanical works. One of the higher plants may be said to be dominant if it be more numerous in individuals and more widely diffused than the other plants of the same country. to whom I am much indebted for valuable advice and assistance on this subject. SPECIES OF THE LARGER GENERA IN EACH COUNTRY VARY MORE FREQUENTLY THAN THE SPECIES OF THE SMALLER GENERA. and all those in the smaller genera on the other side. But if the conferva or parasitic fungus exceeds its allies in the above respects. But my tables further show that. the dominant species--those which range widely. Hooker permits me to add that after having carefully read my manuscript. The whole subject. however. and the species which are most widely diffused within their own country (and this is a different consideration from wide range. Alphonse de Candolle and others have shown that plants which have very wide ranges generally present varieties. and are the most numerous in individuals--which oftenest produce well-marked varieties. and to a certain extent from commonness). which. necessarily have to struggle with the other inhabitants of the country. or. If the plants inhabiting a country as described in any Flora. though in some slight degree modified. And this. treated as it necessarily here is with much brevity. and the tables of the proportional numbers of the varying species. perhaps. as I consider them. in any limited country. as they may be called. which live under nearly the same conditions. are the most diffused in their own country. even in stronger terms. the species which are the most common. and this might have been expected.e. and examined the tables. and as they come into competition (which. Hence it is the most flourishing. the comparison of course relates only to the members of the same group. is a far more important circumstance) with different sets of organic beings. and allusions cannot be avoided to the "struggle for existence. still inherit those advantages that enabled their parents to become dominant over their compatriots. for. At first this seemed a simple task. those including many species) being placed on one side. be divided into two equal masses. H.. soon convinced me that there were many difficulties.WIDE-RANGING. as we shall hereafter see. or. as they are exposed to diverse physical conditions. it will then be dominant within its own class. With respect to the number of individuals or commonness of species." and other questions. Hooker. MUCH DIFFUSED. hereafter to be discussed.

we expect to find saplings. the species of the large genera which present any varieties. there is no apparent reason why more varieties should occur in a group having many species. that I am surprised that my tables show even a small majority on the side of the larger genera. and hence we might expect that the circumstances would generally still be favourable to variation.somewhat larger number of the very common and much diffused or dominant species. species of the same genus) have been formed. as a general rule. for my tables clearly show. into two nearly equal masses. than the species of the smaller genera. But so many causes tend to obscure this result. if we look at each species as a special act of creation.e. as a general rule. This might have been anticipated. but this seems to be connected with the nature of the stations inhabited by them. and has little or no relation to the size of the genera to which the species belong. and. the species of that genus present a number of varieties. wherever many species of a genus have been formed. circumstances have been favourable for variation. are altogether excluded from the tables. It is not that all large genera are now varying much. with from only one to four species. for the mere fact of many species of the same genus inhabiting any country. beyond the average. or that no small genera are now varying and increasing. and when all the least genera. it would have been fatal to my theory. Fresh water and salt-loving plants generally have very wide ranges and are much diffused. and that large genera have often come to their maxima. I was led to anticipate that the species of the larger genera in each country would oftener present varieties. the manufactory of species has been active. for wherever many species of the same genus have been formed. and are thus increasing in the number of their species. And this certainly holds true if varieties be looked at as incipient species. plants low in the scale of organisation are generally much more widely diffused than plants higher in the scale. more especially as we have every reason to believe the process of manufacturing new species to be a slow one. and it has invariably proved to be the case that a larger proportion of the species on the side of the larger genera presented varieties. for if this had been so. I will here allude to only two causes of obscurity. Both these results follow when another division is made. or where. if we may use the expression. From looking at species as only strongly marked and well-defined varieties. These facts are of plain signification on the view that species are only strongly marked and permanent varieties. consequently. for wherever many closely related species (i. than in one having few. that is. Again. inasmuch as geology plainly tells us that small genera have in the lapse of time often increased greatly in size. On the other hand. we might have expected to have found in the larger genera. Where many species of a genus have been formed through variation. or those including many species. declined. of incipient species. many varieties or incipient species ought. than on the side of the smaller genera. and those of the smaller genera on the other side. and here again there is no close relation to the size of the genera. and the coleopterous insects of two districts. The cause of lowly-organised plants ranging widely will be discussed in our chapter on Geographical Distribution. Where many large trees grow. Moreover. shows that there is something in the organic or inorganic conditions of that country favourable to the genus. a larger proportional number of dominant species. invariably present a larger average number of varieties than do the species of the small genera. the species of the larger genera on one side.. that. to be now forming. To test the truth of this anticipation I have arranged the plants of twelve countries. we ought generally to find the manufactory still in action. and .

For instance. when compared with each other or with their parent-species. but which he considers as so closely allied to other species as to be of doubtful value: these sixty-three reputed species range on an average over 6. that the amount of difference between varieties.C. as I call it. is much less than that between the species of the same genus. Watson has divided Great Britain. AND IN HAVING RESTRICTED RANGES. and in so far resemble varieties. as far as my imperfect results go. and. and. No naturalist pretends that all the species of a genus are equally distinct from each other. that where many species of a genus have been formed. Or the case may be put in another way. H.9 of the provinces into which Mr. in the same manner as the varieties of any one species are related to each other. the species of the larger genera resemble varieties. Hence the amount of difference is one very important criterion in settling whether two forms should be ranked as species or varieties. therefore. This statement is indeed scarcely more than a truism. their denominations would be reversed. round their parent-species.disappeared. All that we want to show is. Now. And what are varieties but groups of forms. There is one other point which is worth notice. and this certainly holds good. namely. and clustered round certain forms--that is. for if a variety were found to have a wider range than that of its supposed parent-species. they may generally be divided into sub-genera. But there is reason to believe that the species which are very closely allied to other species. naturalists are compelled to come to a determination by the amount of difference between them. in this . Watson has marked for me in the well-sifted London catalogue of Plants (4th edition) sixty-three plants which are therein ranked as species. or lesser groups. the species of the larger genera are related to each other. RELATED TO EACH OTHER. that in the larger genera. for they differ from each other by a less than the usual amount of difference. unequally related to each other. or sections. on an average many are still forming. BUT UNEQUALLY. in which a number of varieties or incipient species greater than the average are now manufacturing. We have seen that there is no infallible criterion by which to distinguish species and well-marked varieties. and when intermediate links have not been found between doubtful forms. I have endeavoured to test this numerically by averages. There are other relations between the species of large genera and their recorded varieties which deserve notice. But when we come to discuss the principle. Varieties generally have much restricted ranges. after deliberation. judging by analogy whether or not the amount suffices to raise one or both to the rank of species. and it may be said. Mr. that in large genera the amount of difference between the species is often exceedingly small. of divergence of character. In this respect. we shall see how this may be explained. As Fries has well remarked. Moreover. I have also consulted some sagacious and experienced observers. Undoubtedly there is one most important point of difference between varieties and species. and Westwood in regard to insects. and how the lesser differences between varieties tend to increase into the greater differences between species. Now Fries has remarked in regard to plants. little groups of species are generally clustered like satellites around other species. they confirm the view. many of the species already manufactured still to a certain extent resemble varieties. they concur in this view. MANY OF THE SPECIES INCLUDED WITHIN THE LARGER GENERA RESEMBLE VARIETIES IN BEING VERY CLOSELY. often have much restricted ranges. more than do the species of the smaller genera.

for two forms. the forms of life throughout the universe become divided into groups subordinate to groups. and varieties. the species to which these varieties belong range over 14. In large genera the species are apt to be closely but unequally allied together. notwithstanding that they cannot be closely connected. the larger genera also tend to break up into smaller genera. by the discovery of intermediate linking forms. as have the closely allied forms. are generally ranked as varieties. Species very closely allied to other species apparently have restricted ranges. Its bearing on natural selection--The term used in a wide sense--Geometrical ratio of increase--Rapid increase of naturalised animals and plants--Nature of the checks to increase--Competition universal--Effects of climate--Protection from the number of individuals--Complex relations of all animals and plants throughout nature--Struggle for life most severe between individuals and varieties of the same species: often severe between species of the same genus--The relation of organism to organism the most important of all relations. and throughout nature the forms of life which are now dominant tend to become still more dominant by leaving many modified and dominant descendants. It has been seen in the last chapter that among organic beings in a state of nature there is some individual variability: indeed I am not aware that this has ever been disputed. but the amount of difference considered necessary to give to any two forms the rank of species cannot be defined. if differing very little. STRUGGLE FOR EXISTENCE. And thus. fifty-three acknowledged varieties are recorded. So that the acknowledged varieties have very nearly the same restricted average range. varieties cannot be distinguished from species--except. by steps hereafter to be explained.3 provinces. secondly. In all these respects the species of large genera present a strong analogy with varieties. the species of these genera have more than the average number of varieties.7 provinces. CHAPTER III. Thus the larger genera tend to become larger. and thus originated. the two or three hundred doubtful forms of British plants are entitled to hold. and these range over 7. forming little clusters round other species. And we can clearly understand these analogies. Finally. . whereas. as we shall hereafter see.same catalogue. if species once existed as varieties. but which are almost universally ranked by British botanists as good and true species. SUMMARY. and. whereas. marked for me by Mr. We have also seen that it is the most flourishing or dominant species of the larger genera within each class which on an average yield the greatest number of varieties. It is immaterial for us whether a multitude of doubtful forms be called species or sub-species or varieties. by a certain indefinite amount of difference between them. for instance. But. Watson as doubtful species. first. tend to become converted into new and distinct species. these analogies are utterly inexplicable if species are independent creations. what rank. In genera having more than the average number of species in any country. Before entering on the subject of this chapter I must make a few preliminary remarks to show how the struggle for existence bears on natural selection.

is preserved. extinction. are destroyed by birds and beasts of prey. we shall hereafter see. as we shall more fully see in the next chapter. arise? All these results. The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition.if the existence of any well-marked varieties be admitted. in the structure of the beetle which dives through the water. though food may be now superabundant. But the mere existence of individual variability and of some few well-marked varieties. is more accurate. But the expression often used by Mr. In regard to plants. also. or their nestlings. Yet unless it be thoroughly engrained in the mind. but a small number can survive. Owing to this struggle. with every fact on distribution. evidently the result of his great horticultural knowledge. through the accumulation of slight but useful variations. and can adapt organic beings to his own uses. become ultimately converted into good and distinct species. and is sometimes equally convenient. follow from the struggle for life. at greater length. We have seen that man by selection can certainly produce great results. helps us but little in understanding how species arise in nature. We will now discuss in a little more detail the struggle for existence. and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird. if useful. if they be in any degree profitable to the individuals of a species. which constitute what are called distinct genera and which differ from each other more than do the species of the same genus. and will generally be inherited by the offspring. Herbert. for. that. Herbert Spencer. of the many individuals of any species which are periodically born. How have all those exquisite adaptations of one part of the organisation to another part. In my future work this subject will be treated. and is as immeasurably superior to man's feeble efforts. The offspring. how is it that varieties. and are thus constantly destroying life. and variation. however slight and from whatever cause proceeding. which I have called incipient species. will thus have a better chance of surviving. . no one has treated this subject with more spirit and ability than W. by the term natural selection. it is not so at all seasons of each recurring year. as the works of Nature are to those of Art. Again. rarity. it may be asked. or more difficult--at least I found it so--than constantly to bear this conclusion in mind. I have called this principle. Dean of Manchester. or we forget how largely these songsters. though necessary as the foundation for the work. given to him by the hand of Nature. in the plumed seed which is wafted by the gentlest breeze. variations. We behold the face of nature bright with gladness. But Natural Selection. in their infinitely complex relations to other organic beings and to their physical conditions of life. abundance. we see beautiful adaptations everywhere and in every part of the organic world. in short. is a power incessantly ready for action. in order to mark its relation to man's power of selection. and to the conditions of life and of one organic being to another being. the whole economy of nature. or their eggs. Nothing is easier than to admit in words the truth of the universal struggle for life. will be dimly seen or quite misunderstood. we often see superabundance of food. been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and the mistletoe. as it well deserves. of the Survival of the Fittest. will tend to the preservation of such individuals. we do not always bear in mind. we do not see or we forget that the birds which are idly singing round us mostly live on insects or seeds. which in most cases obviously differ from each other far more than do the varieties of the same species? How do those groups of species. by which each slight variation.

which during its natural lifetime produces several eggs or seeds. for.THE TERM. otherwise. The mistletoe is dependent on the apple and a few other trees. or with the individuals of distinct species. growing close together on the same branch. in tempting the birds to devour and thus disseminate its seeds. and during some season or occasional year. Linnaeus has calculated that if an annual plant produced only two seeds--and there is no plant so unproductive as this--and their seedlings next year produced two. bringing forth six young in the interval. The elephant is reckoned the slowest breeder of all known animals. its existence depends on them. Two canine animals. But a plant on the edge of a desert is said to struggle for life against the drought. A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase. in numbers. but can only in a far-fetched sense be said to struggle with these trees. must suffer destruction during some period of its life. including dependence of one being on another. for in this case there can be no artificial increase of food. more or less rapidly. . and it may metaphorically be said to struggle with other fruit-bearing plants. the earth would soon be covered by the progeny of a single pair. and so on. then in twenty years there would be a million plants. after a period of from 740 to 750 years there would be nearly nineteen million elephants alive descended from the first pair. all cannot do so. if this be so. that. In these several senses. and at this rate. and I have taken some pains to estimate its probable minimum rate of natural increase. A plant which annually produces a thousand seeds. and goes on breeding till ninety years old. I should premise that I use this term in a large and metaphorical sense. Although some species may be now increasing. STRUGGLE FOR EXISTENCE. of which only one of an average comes to maturity. or with the physical conditions of life. and including (which is more important) not only the life of the individual. which pass into each other. It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms. may be more truly said to struggle with the plants of the same and other kinds which already clothe the ground. Hence. on the principle of geometrical increase. as more individuals are produced than can possibly survive. for the world would not hold them. As the mistletoe is disseminated by birds. its numbers would quickly become so inordinately great that no country could support the product. there would literally not be standing room for his progeny. may more truly be said to struggle with each other. But several seedling mistletoes. in less than a thousand years. There is no exception to the rule that every organic being naturally increases at so high a rate. may be truly said to struggle with each other which shall get food and live. in a time of dearth. it will be safest to assume that it begins breeding when thirty years old. there must in every case be a struggle for existence. but success in leaving progeny. either one individual with another of the same species. though more properly it should be said to be dependent on the moisture. I use for convenience sake the general term of Struggle for Existence. and surviving till one hundred years old. if too many of these parasites grow on the same tree. USED IN A LARGE SENSE. GEOMETRICAL RATIO OF INCREASE. and no prudential restraint from marriage. Every being. Even slow-breeding man has doubled in twenty-five years. it languishes and dies. if not destroyed.

simply explains their extraordinarily rapid increase and wide diffusion in their new homes. So it is with plants. Our familiarity with the larger domestic animals tends. and among animals there are very few which do not annually pair. If an animal can in any way protect its own eggs or young. many must be produced or the species will become extinct.But we have better evidence on this subject than mere theoretical calculations. and another. cases could be given of introduced plants which have become common throughout whole islands in a period of less than ten years. to mislead us. The only difference between organisms which annually produce eggs or seeds by the thousand. The Fulmar petrel lays but one egg. Several of the plants. Hence we may confidently assert that all plants and animals are tending to increase at a geometrical ratio--that all would rapidly stock every station in which they could any how exist. and that this geometrical tendency to increase must be checked by destruction at some period of life. and those which produce extremely few. that the slow breeders would require a few more years to people. Still more striking is the evidence from our domestic animals of many kinds which have run wild in several parts of the world. and yet in the same country the condor may be the more numerous of the two. supposing that this seed were never destroyed and could be ensured to germinate in a fitting place. the numerous recorded cases of the astonishingly rapid increase of various animals in a state of nature. One fly deposits hundreds of eggs. such as the cardoon and a tall thistle. when circumstances have been favourable to them during two or three following seasons. clothing square leagues of surface almost to the exclusion of every other plant. Their geometrical ratio of increase. they would have been incredible. from Cape Comorin to the Himalaya. I think. and this period in the great majority of cases is an early one. which have been imported from America since its discovery. like the hippobosca. the . But the real importance of a large number of eggs or seeds is to make up for much destruction at some period of life. and endless others could be given. a single one. In a state of nature almost every full-grown plant annually produces seed. and that in a state of nature an equal number would have somehow to be disposed of. It would suffice to keep up the full number of a tree. so that. a whole district. if a single seed were produced once in a thousand years. and we do not keep in mind that thousands are annually slaughtered for food. Falconer. had not been well authenticated. which lived on an average for a thousand years. is. in all cases. for it allows them rapidly to increase in number. The condor lays a couple of eggs and the ostrich a score. and there are plants which now range in India. But this difference does not determine how many individuals of the two species can be supported in a district. under favourable conditions. namely. we see no great destruction falling on them. The obvious explanation is that the conditions of life have been highly favourable. and latterly in Australia. let it be ever so large. as I hear from Dr. the result of which never fails to be surprising. and that there has consequently been less destruction of the old and young and that nearly all the young have been enabled to breed. A large number of eggs is of some importance to those species which depend on a fluctuating amount of food. no one supposes that the fertility of the animals or plants has been suddenly and temporarily increased in any sensible degree. a small number may be produced. if the statements of the rate of increase of slow-breeding cattle and horses in South America. have been introduced from Europe. yet it is believed to be the most numerous bird in the world. which are now the commonest over the wide plains of La Plata. but if many eggs or young are destroyed. In such cases. and yet the average stock be fully kept up.

which determines the average number of a species. The amount of food for each species. and hares on any large estate depends chiefly on the destruction of vermin. although so incomparably better known than any other animal. If not one head of game were shot during the next twenty years in England. but the serving as prey to other animals. grouse. and this is a tremendous destruction. and the case would be the same with turf closely browsed by quadrupeds. We know not exactly what the checks are even in a single instance. I marked all the seedlings of our native weeds as they came up. On the other hand. gives the extreme limit to which each can increase. for even the tiger in India most rarely dares to attack a young elephant protected by its dam. at the same time.average number of any animal or plant depends only indirectly on the number of its eggs or seeds. also. and the number of the species will almost instantaneously increase to any amount. though fully grown plants. Eggs or very young animals seem generally to suffer most. chiefly by slugs and insects. although hundreds of thousands of game animals are now annually shot. from the other species being allowed to grow up freely. for instance. With plants there is a vast destruction of seeds. there seems to be little doubt that the stock of partridges. but this is not invariably the case. The causes which check the natural tendency of each species to increase are most obscure. when we remember that ten per cent. in all probability. that heavy destruction inevitably falls either on the young or old during each generation or at recurrent intervals. dug and cleared. Look at the most vigorous species. mitigate the destruction ever so little. In looking at Nature. none are destroyed by beasts of prey. thus out of twenty species grown on a little plot of mown turf (three feet by four) nine species perished. by so much will it tend to increase still further. just to recall to the reader's mind some of the chief points. are destroyed in vast numbers by various enemies. it is most necessary to keep the foregoing considerations always in mind--never to forget that every single organic being may be said to be striving to the utmost to increase in numbers. If turf which has long been mown. is an . as with the elephant. on a piece of ground three feet long and two wide. more especially in regard to the feral animals of South America. of course. be less game than at present. This subject of the checks to increase has been ably treated by several authors. Seedlings. and out of 357 no less than 295 were destroyed. there would. be let to grow. and. I estimated (chiefly from the greatly reduced numbers of nests in the spring) that the winter of 1854-5 destroyed four-fifths of the birds in my own grounds. and periodical seasons of extreme cold or drought seem to be the most effective of all checks. NATURE OF THE CHECKS TO INCREASE. Climate plays an important part in determining the average numbers of a species. Nor will this surprise any one who reflects how ignorant we are on this head. if no vermin were destroyed. Here I will make only a few remarks. but from some observations which I have made it appears that the seedlings suffer most from germinating in ground already thickly stocked with other plants. and I hope in a future work to discuss it at considerable length. Thus. by as much as it swarms in numbers. Lighten any check. in some cases. that each lives by a struggle at some period of its life. and where there could be no choking from other plants. even in regard to mankind. the more vigorous plants gradually kill the less vigorous. but very frequently it is not the obtaining food.

or in ascending a mountain. as their numbers are checked during the winter. On the other hand. that is abounding in individuals. That climate acts in main part indirectly by favouring other species we clearly see in the prodigious number of plants which in our gardens can perfectly well endure our climate. we forget that each species. the other species must decrease. but in so far as climate chiefly acts in reducing food. increases inordinately in numbers in a small tract. some singular facts in nature such as that of very rare plants being sometimes extremely abundant. we may feel sure that the cause lies quite as much in other species being favoured. than we do in proceeding southward or in descending a mountain. a large stock of individuals of the same species. For in such cases. but in a somewhat lesser degree. we may believe. relatively to the numbers of its enemies. the struggle for life is almost exclusively with the elements. or snow-capped summits. for the number of species of all kinds. The action of climate seems at first sight to be quite independent of the struggle for existence. When a species. owing to highly favourable circumstances. due to the DIRECTLY injurious action of climate. in many cases. which subsist on the same kind of food. it will be the least vigorous individuals. for they cannot compete with our native plants nor resist destruction by our native animals. even on the extreme verge of their range. in our fields. for instance. we invariably see some species gradually getting rarer and rarer. So it is when we travel northward. which will suffer the most. But even some of these so-called epidemics appear to be due to parasitic worms. This view of the necessity of a large stock of the same species for its preservation. or from a damp region to a dry. they will increase in numbers. and as each area is already fully stocked with inhabitants. When we travel southward and see a species decreasing in numbers. and the change of climate being conspicuous. though having a superabundance of food at this one season. from enemies or from competitors for the same place and food. because the seeds are in great excess compared with the number of birds which feed on them. possibly in part through facility of diffusion among the crowded animals. we are tempted to attribute the whole effect to its direct action. even where it most abounds. and here we have a limiting check independent of the struggle for life. Thus we can easily raise plenty of corn and rape-seed. or absolute deserts. acts directly. Even when climate. as in this one being hurt. but which never become naturalised. and therefore of competitors. When we travel from south to north. When we reach the Arctic regions. but any one who has tried knows how troublesome it is to get seed from a few wheat or other such plants in a garden. But this is a false view. nor can the birds. this seems generally to occur with our game animals--often ensue. hence in going northward. whether of the same or of distinct species. been disproportionally favoured: and here comes in a sort of struggle between the parasite and its prey. increase in number proportionally to the supply of seed. it brings on the most severe struggle between the individuals.. I believe. that a plant could exist only where . explains. and that of some social plants being social. we far oftener meet with stunted forms. is absolutely necessary for its preservation. which have from some cause. and finally disappearing. etc. epidemics--at least. or those which have got least food through the advancing winter. in the few spots where they do exist. I have in this case lost every single seed. and if these enemies or competitors be in the least degree favoured by any slight change of climate. is constantly suffering enormous destruction at some period of its life.extraordinarily severe mortality from epidemics with man. extreme cold. decreases northward.

though they swarm southward and northward in a feral state. so close together that all cannot live. and the heath was frequented by two or three distinct insectivorous birds. I should add that the good effects of intercrossing. Here there are extensive heaths. Here we see how potent has been the effect of the introduction of a single tree. Many cases are on record showing how complex and unexpected are the checks and relations between organic beings.the conditions of its life were so favourable that many could exist together. it became thickly clothed with vigorously growing young firs. which could not be found on the heath. as soon as the land was enclosed. for six insectivorous birds were very common in the plantations. I counted thirty-two little trees. and this would . But on looking closely between the stems of the heath. The change in the native vegetation of the planted part of the heath was most remarkable. must be habitually checked by some means. where I had ample means of investigation. which lays its eggs in the navels of these animals when first born. with twenty-six rings of growth. and self-sown firs are now springing up in multitudes. during many years tried to raise its head above the stems of the heath. in Surrey. more than is generally seen in passing from one quite different soil to another: not only the proportional numbers of the heath-plants were wholly changed. When I ascertained that these young trees had not been sown or planted I was so much surprised at their numbers that I went to several points of view. COMPLEX RELATIONS OF ALL ANIMALS AND PLANTS TO EACH OTHER IN THE STRUGGLE FOR EXISTENCE. for here neither cattle nor horses nor dogs have ever run wild. but I will not here enlarge on this subject. I found a multitude of seedlings and little trees. No wonder that. Yet the heath was so extremely barren and so extensive that no one would ever have imagined that cattle would have so closely and effectually searched it for food. and one of them. the parasitic insects would probably increase. The increase of these flies. but twelve species of plants (not counting grasses and carices) flourished in the plantations. so that cattle could not enter. probably by other parasitic insects. In one square yard. on the estate of a relation. I will give only a single instance. which. which were not to be seen on the heath. except the old planted clumps. if certain insectivorous birds were to decrease in Paraguay. which have to struggle together in the same country. and literally I could not see a single Scotch fir. which had never been touched by the hand of man. nothing whatever else having been done. with a few clumps of old Scotch firs on the distant hill-tops: within the last ten years large spaces have been enclosed. but several hundred acres of exactly the same nature had been enclosed twenty-five years previously and planted with Scotch fir. at a point some hundred yards distant from one of the old clumps. Perhaps Paraguay offers the most curious instance of this. no doubt come into play in many of these cases. with the exception of the land having been enclosed. whence I could examine hundreds of acres of the unenclosed heath. Here we see that cattle absolutely determine the existence of the Scotch fir. and the ill effects of close interbreeding. which had been perpetually browsed down by the cattle. But how important an element enclosure is. In Staffordshire. but in several parts of the world insects determine the existence of cattle. and Azara and Rengger have shown that this is caused by the greater number in Paraguay of a certain fly. The effect on the insects must have been still greater. Hence. I plainly saw near Farnham. though a simple one. interested me. numerous as they are. there was a large and extremely barren heath. and had failed. had. and thus save the species from utter destruction.

Battle within battle must be continually recurring with varying success. we invoke cataclysms to desolate the world. and yet in the long-run the forces are so nicely balanced that the face of nature remains for long periods of time uniform. never sets a seed. and so high our presumption. from its peculiar structure. the frequency of certain flowers in that district! In the case of every species. on the number of cats. which destroy their combs and nests. the heartsease and red clover would become very rare. When we look at the plants and bushes clothing an entangled bank. but twenty other heads. for instance twenty heads of Dutch clover (Trifolium repens) yielded 2. who has long attended to the habits of humble-bees. or even the existence of the species. Not that under nature the relations will ever be as simple as this. for other bees do not visit this flower. as we have just seen in Staffordshire. and as we do not see the cause. though assuredly the merest trifle would give the victory to one organic being over another. a very different vegetation springs . many different checks. produced not one. I shall hereafter have occasion to show that the exotic Lobelia fulgens is never visited in my garden by insects. we are tempted to attribute their proportional numbers and kinds to what we call chance. if the whole genus of humble-bees became extinct or very rare in England. Nearly all our orchidaceous plants absolutely require the visits of insects to remove their pollen-masses and thus to fertilise them. or wholly disappear. as other bees cannot reach the nectar. and Colonel Newman says. "Near villages and small towns I have found the nests of humble-bees more numerous than elsewhere." Now the number of mice is largely dependent. so profound is our ignorance. some one check or some few being generally the most potent. the insectivorous birds. remote in the scale of nature. I find from experiments that humble-bees are almost indispensable to the fertilisation of the heartsease (Viola tricolor). and consequently. are bound together by a web of complex relations.290 seeds. but all will concur in determining the average number. and Colonel Newman. I have also found that the visits of bees are necessary for the fertilisation of some kinds of clover. and this would certainly greatly alter (as indeed I have observed in parts of South America) the vegetation: this again would largely affect the insects. that we marvel when we hear of the extinction of an organic being. but I doubt whether they could do so in the case of the red clover. In some cases it can be shown that widely-different checks act on the same species in different districts. and during different seasons or years. probably come into play. 100 heads of red clover (T. and so onwards in ever-increasing circles of complexity. through the intervention first of mice and then of bees. or invent laws on the duration of the forms of life! I am tempted to give one more instance showing how plants and animals. But how false a view is this! Every one has heard that when an American forest is cut down. Nevertheless. protected from bees. Humble bees alone visit red clover.700 seeds. Again. Hence we may infer as highly probable that. which I attribute to the number of cats that destroy the mice. as every one knows. believes that "more than two-thirds of them are thus destroyed all over England.lessen the number of the navel-frequenting flies--then cattle and horses would become feral. but the same number of protected heads produced not a single seed. pratense) produced 2. The number of humble-bees in any district depends in a great measure upon the number of field-mice. and this. acting at different periods of life. from their weight not being sufficient to depress the wing petals. It has been suggested that moths may fertilise the clovers." Hence it is quite credible that the presence of a feline animal in large numbers in a district might determine.

and the seed then mixed in due proportion. To keep up a mixed stock of even such extremely close varieties as the variously coloured sweet-peas. in the same manner as beings in a state of nature. but how simple is the problem where each shall fall compared to that of the action and reaction of the innumerable plants and animals which have determined. or on the trees. or are naturally the most fertile. The same result has followed from keeping together different varieties of the medicinal leech. and the mixed seed be resown. habits. so that they cannot be kept together. now display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests.up. the struggle will generally be almost equally severe. what war between insect and insect--between insects. that the original proportions of a mixed stock (crossing being prevented) could be kept up for half-a-dozen generations. Throw up a handful of feathers. require the same food. if they were allowed to struggle together. lies generally between beings remote in the scale of nature. the proportional numbers and kinds of trees now growing on the old Indian ruins! The dependency of one organic being on another. their seeds and seedlings. and if the seed or young were not annually preserved in due proportion. and will consequently in a few years supplant the other varieties. as of a parasite on its prey. each annually scattering its seeds by the thousand. and constitution. In the case of varieties of the same species. they must be each year harvested separately. and are exposed to the same dangers. much similarity in habits and constitution. In Australia the imported hive-bee is rapidly exterminating the small. snails. otherwise the weaker kinds will steadily decrease in number and disappear. and we sometimes see the contest soon decided: for instance. for they frequent the same districts. will beat the others and so yield more seed. if they come into competition with each other. But the struggle will almost invariably be most severe between the individuals of the same species. but it has been observed that ancient Indian ruins in the Southern United States. The recent increase of the missel-thrush in parts of Scotland has caused the decrease of the song-thrush. We see this in the recent extension over parts of the United States of one species of swallow having caused the decrease of another species. and all fall to the ground according to definite laws. stingless native . in the course of centuries. or on the other plants which first clothed the ground and thus checked the growth of the trees. the struggle will generally be more severe between them. which must formerly have been cleared of trees. As the species of the same genus usually have. some of the varieties which best suit the soil or climate. STRUGGLE FOR LIFE MOST SEVERE BETWEEN INDIVIDUALS AND VARIETIES OF THE SAME SPECIES. It may even be doubted whether the varieties of any of our domestic plants or animals have so exactly the same strength. though by no means invariably. and other animals with birds and beasts of prey--all striving to increase. What a struggle must have gone on during long centuries between the several kinds of trees. than between the species of distinct genera. How frequently we hear of one species of rat taking the place of another species under the most different climates! In Russia the small Asiatic cockroach has everywhere driven before it its great congener. if several varieties of wheat be sown together. and always in structure. This is likewise sometimes the case with those which may strictly be said to struggle with each other for existence. all feeding on each other. as in the case of locusts and grass-feeding quadrupeds. So again with the varieties of sheep: it has been asserted that certain mountain-varieties will starve out other mountain-varieties.

In this case we can clearly see that if we wish in imagination to give the plant the power of increasing in numbers. will competition cease. so that the seeds may be widely distributed and fall on unoccupied ground. This ought to convince us of our ignorance on the mutual . Hence we can see that when a plant or animal is placed in a new country. that the structure of every organic being is related. which fill nearly the same place in the economy of nature. The land may be extremely cold or dry. but we have reason to believe that only a few plants or animals range so far. On the confines of its geographical range. we should have to modify it in a different way to what we should have had to do in its native country. We can dimly see why the competition should be most severe between allied forms. In the water-beetle. A corollary of the highest importance may be deduced from the foregoing remarks. or over the animals which prey on it. when sown in the midst of long grass. The store of nutriment laid up within the seeds of many plants seems at first sight to have no sort of relation to other plants. But from the strong growth of young plants produced from such seeds. This is obvious in the structure of the teeth and talons of the tiger. It is good thus to try in imagination to give any one species an advantage over another. but probably in no one case could we precisely say why one species has been victorious over another in the great battle of life. damper or drier districts. or on which it preys. the relation seems at first confined to the elements of air and water. Yet the advantage of the plumed seeds no doubt stands in the closest relation to the land being already thickly clothed with other plants. allows it to compete with other aquatic insects. and in the flattened and fringed legs of the water-beetle. for we should have to give it some advantage over a different set of competitors or enemies. a change of constitution with respect to climate would clearly be an advantage to our plant. that they are destroyed exclusively by the rigour of the climate. we should have to give it some advantage over its competitors. it may be suspected that the chief use of the nutriment in the seed is to favour the growth of the seedlings. the conditions of its life will generally be changed in an essential manner. among new competitors. whilst struggling with other plants growing vigorously all around. yet there will be competition between some few species. for elsewhere it ranges into slightly hotter or colder. and so in other cases. One species of charlock has been known to supplant another species. dampness or dryness. Probably in no single instance should we know what to do. although the climate may be exactly the same as in its former home. with which it comes into competition for food or residence. If its average numbers are to increase in its new home. the structure of its legs. in the most essential yet often hidden manner. to that of all other organic beings. to hunt for its own prey.bee. or between the individuals of the same species. or from which it has to escape. and in that of the legs and claws of the parasite which clings to the hair on the tiger's body. Not until we reach the extreme confines of life. in the Arctic regions or on the borders of an utter desert. Look at a plant in the midst of its range! Why does it not double or quadruple its numbers? We know that it can perfectly well withstand a little more heat or cold. as peas and beans. But in the beautifully plumed seed of the dandelion. namely. and to escape serving as prey to other animals. for the warmest or dampest spots. so well adapted for diving.

apply under nature? I think we shall see that it can act most efficiently. in those under nature. by man. briefly discussed in the last chapter. 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. CHAPTER IV. and variability ensues. intercrossing. or at intervals.relations of all organic beings. but similar changes of conditions might and do occur under nature. can we doubt (remembering that many more individuals are born than can possibly survive) that individuals having any advantage. NATURAL SELECTION. When we reflect on this struggle we may console ourselves with the full belief that the war of nature is not incessant. be borne in mind. This preservation of favourable individual differences and variations. number of individuals--Slow action--Extinction caused by Natural Selection--Divergence of Character. But the variability. and the happy survive and multiply. OR THE SURVIVAL OF THE FITTEST. that death is generally prompt. isolation. over others. I have called Natural Selection. the healthy. that each. he can only preserve and accumulate such as do occur. at some period of its life. during some season of the year. How will the struggle for existence. a conviction as necessary. and that the vigorous. that no fear is felt. however slight. related to the diversity of inhabitants of any small area and to naturalisation--Action of Natural Selection. should occur in the course of many successive generations? If such do occur. Let the endless number of slight variations and individual differences occurring in our domestic productions. during each generation. as it is difficult to acquire. All that we can do is to keep steadily in mind that each organic being is striving to increase in a geometrical ratio. 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. has to struggle for life and to suffer great destruction. and. and the destruction of those which are injurious. act in regard to variation? Can the principle of selection. Can it then be thought improbable. he can neither originate varieties nor prevent their occurrence. or the Survival of the Fittest. it may truly be said that the whole organisation becomes in some degree plastic. which we have seen is so potent in the hands of man. Variations . on the descendants from a common parent--Explains the Grouping of all organic beings--Advance in organisation--Low forms preserved--Convergence of character--Indefinite multiplication of species--Summary. in a lesser degree. through Divergence of Character and Extinction. which we almost universally meet with in our domestic productions is not directly produced. as well as the strength of the hereditary tendency. Let it also be borne in mind how infinitely complex and close-fitting are the mutual relations of all organic beings to each other and to their physical conditions of life. Under domestication. we may feel sure that any variation in the least degree injurious would be rigidly destroyed. would have the best chance of surviving and procreating their kind? On the other hand. as Hooker and Asa Gray have well remarked. Unintentionally he exposes organic beings to new and changing conditions of life. and consequently what infinitely varied diversities of structure might be of use to each being under changing conditions of life.

natural selection is a false term. We have good reason to believe. natural selection is not applicable to them! In the literal sense of the word. and in the foregoing cases the conditions the changed. or would ultimately become fixed. 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. In such cases. as perhaps we see in certain polymorphic species. Some have even imagined that natural selection induces variability. independently of the change of climate itself. but I mean by nature. would tend to be preserved. that changes in the conditions of life give a tendency to increased variability. from what we have seen of the intimate and complex manner in which the inhabitants of each country are bound together. into which new and better adapted forms could not freely enter. would seriously affect the others. by affording a better chance of the occurrence of profitable variations. and this would likewise seriously disturb the relations of some of the former inhabitants. Under the term of "variations. only the aggregate action and product of many natural laws. and some species will probably become extinct. No one objects to agriculturists speaking of the potent effects of man's selection. and they are almost necessary for brevity. With a little familiarity such superficial objections will be forgotten. The proportional numbers of its inhabitants will almost immediately undergo a change. must of necessity first occur. We shall best understand the probable course of natural selection by taking the case of a country undergoing some slight physical change. which in any way favoured the individuals of any species. natural selection can do nothing. new forms would certainly immigrate. As man can produce a great result with his domestic animals . of climate. and natural selection would have free scope for the work of improvement. slight modifications. and it has even been urged that. So again it is difficult to avoid personifying the word Nature. Several writers have misapprehended or objected to the term Natural Selection." it must never be forgotten that mere individual differences are included. 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. and would be left either a fluctuating element. Unless such occur.neither useful nor injurious would not be affected by natural selection. had the area been open to immigration. 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. We may conclude. or of a country partly surrounded by barriers. by better adapting them to their altered conditions. 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. Others have objected that the term selection implies conscious choice in the animals which become modified. that any change in the numerical proportions of the inhabitants. for. and in this case the individual differences given by nature. whereas it implies only the preservation of such variations as arise and are beneficial to the being under its conditions of life. and by laws the sequence of events as ascertained by us. If the country were open on its borders. which man for some object selects. as plants have no volition. these same places would have been seized on by intruders. as shown in the first chapter. for instance. no doubt. and this would manifestly be favourable to natural selection. owing to the nature of the organism and the nature of the conditions.

for in all countries. he exposes sheep with long and short wool to the same climate. cares nothing for appearances. preserving and adding up all that are good. We see nothing of these slow changes in progress. a great result by his methodical and unconscious means of selection. WHENEVER AND WHEREVER OPPORTUNITY OFFERS. so could natural selection. and should plainly bear the stamp of far higher workmanship? It may metaphorically be said that natural selection is daily and hourly scrutinising. that none of them could be still better adapted or improved. For as all the inhabitants of each country are struggling together with nicely balanced forces. then. throughout the world. on every shade of constitutional difference. but far more easily from having incomparably longer time for action. or any unusual degree of isolation. that they should be infinitely better adapted to the most complex conditions of life. we may safely conclude that the natives might have been modified with advantage. he feeds a long and a short-beaked pigeon on the same food. Man keeps the natives of many climates in the same country. as far as lies in his power. compared with those accumulated by Nature during whole geological periods! Can we wonder. that Nature's productions should be far "truer" in character than man's productions. and so be preserved. He seldom exercises each selected character in some peculiar and fitting manner. Nor do I believe that any great physical change. he does not rigidly destroy all inferior animals. Under nature. rejecting those that are bad. until the hand of time has marked the long lapse of ages.and plants by adding up in any given direction individual differences. 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. extremely slight modifications in the structure or habits of one species would often give it an advantage over others. as is implied by the fact of their selection. so as to have better resisted the intruders. except in so far as they are useful to any being. does not allow the most vigorous males to struggle for the females. and then so imperfect is our view into long-past geological ages that we see . Every selected character is fully exercised by her. he does not exercise a long-backed or long-legged quadruped in any peculiar manner. as long as the species continued under the same conditions of life and profited by similar means of subsistence and defence. the slightest variations. No country can be named in which all the native inhabitants are now so perfectly adapted to each other and to the physical conditions under which they live. As man can produce. Nature only for that of the being which she tends. all his productions. on the whole machinery of life. and still further modifications of the same kind would often still further increase the advantage. but protects during each varying season. at the improvement of each organic being in relation to its organic and inorganic conditions of life. if I may be allowed to personify the natural preservation or survival of the fittest. as of climate. Man selects only for his own good. And as foreigners have thus in every country beaten some of the natives. the natives have been so far conquered by naturalised productions that they have allowed some foreigners to take firm possession of the land. the slightest differences of structure or constitution may well turn the nicely-balanced scale in the struggle for life. what may not natural selection effect? Man can act only on external and visible characters: Nature. or at least by some modification prominent enough to catch the eye or to be plainly useful to him. silently and insensibly working. and consequently how poor will be his results. He often begins his selection by some half-monstrous form. to check immigration. and certainly has produced. She can act on every internal organ. How fleeting are the wishes and efforts of man! How short his time.

we must not forget that climate. the down on the fruit and the colour of the flesh are considered by botanists as characters of the most trifling importance. may thus be acted on. whether a smooth or downy. determines whether they shall live or die. the red-grouse the colour of heather. etc. assuredly. when once acquired. which feed on the "paint-root" in Virginia. is likewise a simple assumption. whereas another disease attacks yellow-fleshed peaches far more than those with other coloured flesh. in the . we must believe that these tints are of service to these birds and insects in preserving them from danger. But whether it is true. we can judge only by seeing how far the hypothesis accords with and explains the general phenomena of nature. It is also necessary to bear in mind that. a yellow or a purple-fleshed fruit. When we see leaf-eating insects green. such differences would effectually settle which variety. Although natural selection can act only through and for the good of each being. that purple plums suffer far more from a certain disease than yellow plums. In looking at many small points of difference between species. which we are apt to consider as of very trifling importance. size and flavour of the seeds of the many varieties of our culinary and agricultural plants. with all the aids of art. and hawks are guided by eyesight to their prey. yet we hear from an excellent horticulturist. and these must again be preserved. true and constant. the alpine ptarmigan white in winter. step by step. vary or present individual differences of the same favourable nature as before. a variety. As we see that those variations which.. this can hardly be considered as an unwarrantable assumption. in the shape. other modifications. in the caterpillar and cocoon stages of the varieties of the silkworm. have no doubt produced some direct effect. Grouse. in a state of nature. the ordinary belief that the amount of possible variation is a strictly limited quantity. We have seen how the colour of hogs. when one part varies and the variations are accumulated through natural selection. as far as our ignorance permits us to judge. In plants. In order that any great amount of modification should be effected in a species. seem quite unimportant. where the trees would have to struggle with other trees and with a host of enemies.--so much so that on parts of the continent persons are warned not to keep white pigeons. food. and in keeping that colour. If. they are known to suffer largely from birds of prey. when once formed must again. tend to reappear in the offspring at the same period. we should remember how essential it is in a flock of white sheep to destroy a lamb with the faintest trace of black. under domestication. should succeed. Hence natural selection might be effective in giving the proper colour to each kind of grouse. than those with down. which. and bark-feeders mottled-grey. that in the United States smooth-skinned fruits suffer far more from a beetle. as being the most liable to destruction. often of the most unexpected nature. will ensue.only that the forms of life are now different from what they formerly were. owing to the law of correlation. perhaps after a long interval of time. a Curculio. if not destroyed at some period of their lives. yet characters and structures. Downing. Seeing that individual differences of the same kind perpetually recur. On the other hand. and so onward. Nor ought we to think that the occasional destruction of an animal of any particular colour would produce little effect. these slight differences make a great difference in cultivating the several varieties. would increase in countless numbers. appear at any particular period of life. for instance.

in the horns of our sheep and cattle when nearly adult. if of high importance to it. the process of modification would be very slow. conversely. so in a state of nature natural selection will be enabled to act on and modify organic beings at any age. If it profit a plant to have its seeds more and more widely disseminated by the wind. the species would become extinct. will tend to propagate their kind in larger numbers than the less well adapted. than in the cotton-planter increasing and improving by selection the down in the pods on his cotton-trees. which had the most powerful and hardest beaks. must be annually destroyed by accidental causes. But let the destruction of the adults be ever so heavy. So again a vast number of mature animals and plants. the structure of the adult. more delicate and more easily broken shells might be selected. if the community profits by the selected change. the best adapted individuals. might be modified to any extent by natural selection. If the numbers be wholly kept down by the causes just . so that fanciers assist in the act of hatching. What natural selection cannot do. It may be well here to remark that with all beings there must be much fortuitous destruction. 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. for all with weak beaks would inevitably perish: or. used for breaking the eggs. and in the colour of the down of their chickens. the great jaws possessed by certain insects. is to modify the structure of one species. whether or not they be the best adapted to their conditions. and by their inheritance at a corresponding age. if nature had to make the beak of a full-grown pigeon very short for the bird's own advantage. without giving it any advantage. A structure used only once in an animal's life. For instance.eggs of poultry. Natural selection may modify and adapt the larva of an insect to a score of contingencies. for instance. through correlation. used exclusively for opening the cocoon--or the hard tip to the beak of unhatched birds. and there would be simultaneously the most rigorous selection of all the young birds within the egg. by the accumulation of variations profitable at that age. I can see no greater difficulty in this being effected through natural selection. supposing that there is any variability in a favourable direction. which can have little or no influence on the course of natural selection. if not destroyed. I cannot find one case which will bear investigation. and these modifications may affect. for the good of another species. Now. and these could be modified through natural selection only if they varied in some manner which protected them from their enemies. 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. a vast number of eggs or seeds are annually devoured. have yielded individuals better adapted to their conditions of life than any of those which happened to survive. Yet many of these eggs or seeds would perhaps. modifications in the adult may affect the structure of the larva. So. It has been asserted. and though statements to this effect may be found in works of natural history. but in all cases natural selection will ensure that they shall not be injurious: for if they were so. that of the best short-beaked tumbler-pigeons a greater number perish in the egg than are able to get out of it. the thickness of the shell being known to vary like every other structure. wholly different from those which concern the mature insect. 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.

congregate. 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. Generally. by singing. The war is. as the mane of the lion. special means of defence may be given through means of sexual selection. believe that there is the severest rivalry between the males of many species to attract. This form of selection depends. generally the males. confined to the male sex. How low in the scale of nature the law of battle descends I know not. severest between the males of polygamous animals. birds of paradise. The rock-thrush of Guiana. and show off in the best manner. the contest is often of a more peaceful character. fighting for a particular female who sits by. and then retires with the conqueror. The result is not death to the unsuccessful competitor. male salmons have been observed fighting all day long. might surely give indomitable courage. but this is no valid objection to its efficiency at other times and in other ways. standing by as spectators. according to his standard of beauty. in nearly the same manner as does the brutal cockfighter by the careful selection of his best cocks. Heron has described how a pied peacock was eminently attractive to all his hen birds. the most melodious . but on having special weapons. I cannot here enter on the necessary details. for the shield may be as important for victory as the sword or spear. natural selection will be powerless in certain beneficial directions. their gorgeous plumage. A hornless stag or spurless cock would have a poor chance of leaving numerous offspring. but if man can in a short time give beauty and an elegant carriage to his bantams. will leave most progeny. male stag-beetles sometimes bear wounds from the huge mandibles of other males. an apparently unconcerned beholder of the struggle. during thousands of generations. the females. and some others.indicated. for the possession of the other sex. Among birds. I can see no good reason to doubt that female birds. they likewise perform strange antics before the females. not on a struggle for existence in relation to other organic beings or to external conditions. Sexual selection is. and these seem oftenest provided with special weapons. All those who have attended to the subject. length of spur. but few or no offspring. therefore. at last choose the most attractive partner. and strength to the wing to strike in the spurred leg. and the hooked jaw to the male salmon. like Indians in a war-dance. by always allowing the victor to breed. and whirling round. those which are best fitted for their places in nature. the males of certain hymenopterous insects have been frequently seen by that inimitable observer M. Those who have closely attended to birds in confinement well know that they often take individual preferences and dislikes: thus Sir R. as will often have been the case. and successive males display with the most elaborate care. Fabre. But in many cases victory depends not so much on general vigour. but on a struggle between the individuals of one sex. perhaps. or for one sex to be modified in relation to the other sex. by selecting. SEXUAL SELECTION. Sexual selection. for the possession of the females. though to them and to others. This leads me to say a few words on what I have called sexual selection. less rigorous than natural selection. male alligators have been described as fighting. so no doubt it will be under nature. which. bellowing. as commonly occurs. Thus it is rendered possible for the two sexes to be modified through natural selection in relation to different habits of life. The males of carnivorous animals are already well armed. as is sometimes the case. Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex. the most vigorous males.

and transmitted to the males alone or to both sexes at corresponding ages. Thus it is. OR THE SURVIVAL OF THE FITTEST. taking to catch rats. Under such circumstances the swiftest and slimmest wolves have the best chance of surviving. with shorter legs. and the other more bulky. Some of its young would probably inherit the same habits or structure. but I have not space here to enter on this subject. which more frequently attacks the shepherd's flocks. another mice.or beautiful males. I must beg permission to give one or two imaginary illustrations. according to Mr. some slight advantage over other males. according to Mr. or charms. and so be preserved or selected. that when the males and females of any animal have the same general habits of life. for we often observe great differences in the natural tendencies of our domestic animals. Pierce. indeed. it would have the best chance of surviving and of leaving offspring. another hares or rabbits. might produce a marked effect. 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. one with a light greyhound-like form. I can see no more reason to doubt that this would be the result. 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. in their weapons. which apparently have not been augmented through selection by man. or ornament. according to their standard of beauty. bringing home winged game. in the United States. Yet. during that season of the year when the wolf was hardest pressed for food. and another hunting on marshy ground and almost nightly catching woodcocks or snipes. had from any change in the country increased in numbers. which they have transmitted to their male offspring alone. John. provided always that they retained strength to master their prey at this or some other period of the year. can partly be explained through the action of sexual selection on variations occurring at different ages. I may add that. colour. In order to make it clear how. a cub might be born with an innate tendency to pursue certain kinds of prey. with respect to the plumage of male and female birds. and some by fleetness. there are two varieties of the wolf inhabiting the Catskill Mountains. when they were compelled to prey on other animals. securing some by craft. and by the repetition of this process. a deer for instance. means of defence. Let us take the case of a wolf. for instance. or that other prey had decreased in numbers. by individual males having had. one cat. in comparison with the plumage of the young. St. Even without any change in the proportional numbers of the animals on which our wolf preyed. but differ in structure. and let us suppose that the fleetest prey. I would not wish to attribute all sexual differences to this agency: for we see in our domestic animals peculiarities arising and becoming attached to the male sex. if any slight innate change of habit or of structure benefited an individual wolf. a new variety might be formed which would . in successive generations. Now. which preys on various animals. which pursues deer. such differences have been mainly caused by sexual selection: that is. natural selection acts. one cat. than that man should be able to improve the fleetness of his greyhounds by careful and methodical selection. as I believe. Some well-known laws. as I believe. The tendency to catch rats rather than mice is known to be inherited. Nor can this be thought very improbable. some by strength. had the tuft appeared under domestication it would have been called a monstrosity. ILLUSTRATIONS OF THE ACTION OF NATURAL SELECTION.

and this chance would go on decreasing in the succeeding generations. and that. which pursues deer. that this result would follow from the preservation during many generations of a large number of individuals with more or less strongly curved beaks. would be a rare event. until reading an able and valuable article in the "North British Review" (1867). as the Reviewer goes onto show. yet the chances would be strongly against its survival. with shorter legs. but to this subject of intercrossing we shall soon have to return. would naturally be forced to hunt different prey. again. of which. and this led me fully to discuss the results of unconscious selection by man. as long as the existing conditions remained the same. I saw. It should be observed that in the above illustration. and if one were born with its beak strongly curved. whether slight or strongly marked. giving it twice as good a chance of life as that of the other individuals. and on the destruction of the worst. a bird of some kind could procure its food more easily by having its beak curved. The author takes the case of a pair of animals. I think. Supposing it to survive and to breed. It should not. be disputed. however. Or. if the varying individual did not actually transmit to its offspring its newly-acquired character. If. nevertheless there would be a very poor chance of this one individual perpetuating its kind to the exclusion of the common form. and from the continued preservation of the individuals best fitted for the two sites. be overlooked that certain rather strongly marked variations. which varied in some manner. one with a light greyhound-like form. Nevertheless. judging by what we see taking place under domestication. and that half its young inherited the favourable variation. only two on an average survive to pro-create their kind. and the other more bulky. I speak of the slimmest individual wolves. a still stronger tendency to vary in the same manner. which depends on the preservation of all the more or less valuable individuals. such as a monstrosity. there are two varieties of the wolf inhabiting the Catskill Mountains in the United States. In former editions of this work I sometimes spoke as if this latter alternative had frequently occurred. the wolves inhabiting a mountainous district. 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 . The justice of these remarks cannot. the young would have only a slightly better chance of surviving and breeding. which no one would rank as mere individual differences. according to Mr. it would generally be lost by subsequent intercrossing with ordinary individuals. if at first preserved. I may add. which more frequently attacks the shepherd's flocks. and which consequently flourished. still. I saw the great importance of individual differences. He then shows that if a single individual were born. two varieties might slowly be formed. and those frequenting the lowlands. but there can hardly be a doubt. but by no means so for many of the lower organisms. and from the destruction of a still larger number with the straightest beaks. These varieties would cross and blend where they met. for instance. In such cases. frequently recur owing to a similar organisation being similarly acted on--of which fact numerous instances could be given with our domestic productions. This is rather an extreme estimate for most of the higher animals. that. Pierce. it would undoubtedly transmit to them. from various causes of destruction. producing during their lifetime two hundred offspring. that the preservation in a state of nature of any occasional deviation of structure.either supplant or coexist with the parent-form of wolf. could be perpetuated. I did not appreciate how rarely single variations. also. and not of any single strongly marked variation having been preserved.

and would often breed together. it would slowly spread from a central district. the original form would soon be supplanted by the modified form. other holly-trees bear only female flowers. which have four stamens producing a rather small quantity of pollen. and the act of crossing. and a rudimentary pistil. and would oftenest be crossed. In cases of this kind. 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. or tenth part of the individuals may have been thus affected. would be selected. and would often transport it from one flower to another. yet if a little pollen were carried. is greedily sought by insects. of which fact several instances could be given. by glands at the base of the stipules in some Leguminosae. Consequently each newly-formed variety would generally be at first local. and that they do this effectually I could easily show by many striking facts. which consequently would have the best chance of flourishing and surviving. through the survival of the fittest. had been rendered highly attractive to insects. and at the backs of the leaves of the common laurel. we see this even with migratory birds. by the pollen-devouring insects from flower to flower. excreting most nectar. and do not needlessly wander about. Or only a third. fifth. these have a full-sized . which almost always return to the same spot. as can be fully proved. and as pollen is formed for the sole purpose of fertilisation. would oftenest be visited by insects. though small in quantity. To the effects of intercrossing in eliminating variations of all kinds.selection. Certain plants excrete sweet juice. but their visits do not in any way benefit the plant. so that similarly modified individuals would soon exist in a small body together. When our plant. Now. I shall have to recur. for instance. at first occasionally and then habitually. regularly carry pollen from flower to flower. and the individuals which produced more and more pollen. so as to favour in any degree the transportal of the pollen. apparently for the sake of eliminating something injurious from the sap: this is effected. as seems to be the common rule with varieties in a state of nature. and a cross thus effected. also. in relation to the size and habits of the particular insect which visited them. We might have taken the case of insects visiting flowers for the sake of collecting pollen instead of nectar. Some holly-trees bear only male flowers. but it may be here remarked that most animals and plants keep to their proper homes. The flowers. that it was formerly ranked as a distinct species under the name of Uria lacrymans. its destruction appears to be a simple loss to the plant. I will give only one. would likewise be favoured. It may be worth while to give another and more complex illustration of the action of natural selection. and had larger anthers. competing with and conquering the unchanged individuals on the margins of an ever-increasing circle. This juice. if the variation were of a beneficial nature. by the above process long continued. Thus Graba estimates that about one-fifth of the guillemots in the Faroe Islands consist of a variety so well marked. which had their stamens and pistils placed. gives rise to vigorous seedlings. as likewise illustrating one step in the separation of the sexes of plants. unintentionally on their part. The plants which produced flowers with the largest glands or nectaries. and so in the long-run would gain the upper hand and form a local variety. The flowers of two distinct individuals of the same species would thus get crossed. although nine-tenths of the pollen were destroyed it might still be a great gain to the plant to be thus robbed. If the new variety were successful in its battle for life. they would. Insects in seeking the nectar would get dusted with pollen.

but not out of the common red clover. under the microscope. must be very trifling. by which the separation of the sexes in plants of various kinds is apparently now in progress. but only in the autumn. without exception. according to Asa Gray. in an exactly intermediate condition. and that certain insects depended in main part on its nectar for food. taken from different branches. it may be believed that under certain circumstances individual differences in the curvature or length of the proboscis. As the wind had set for several days from the female to the male tree. as pollen is already carried regularly from flower to flower. so that whole fields of the red clover offer in vain an abundant supply of precious nectar to the hive-bee. I could give many facts showing how anxious bees are to save time: for instance. nor whether another published statement can be trusted. The difference in the length of the corolla in the two kinds of clover. and as a more complete separation of the sexes of our plant would be advantageous on the principle of the division of labour. another process might commence. the pollen could not thus have been carried. or. so that certain individuals would be able to obtain their food more quickly than others. then. The weather had been cold and boisterous and therefore not favourable to bees. there were a few pollen-grains. the flowers of the second crop are somewhat smaller. and on some a profusion. The tubes of the corolla of the common red or incarnate clovers (Trifolium pratense and incarnatum) do not on a hasty glance appear to differ in length. that the Ligurian . which had flown from tree to tree in search of nectar." hence we may believe that it would be advantageous to a plant to produce stamens alone in one flower or on one whole plant. It would take up too much space to show the various steps. their habit of cutting holes and sucking the nectar at the bases of certain flowers. in which not a grain of pollen can be detected. and pistils alone in another flower or on another plant.. for I have repeatedly seen. sometimes the male organs and sometimes the female organs become more or less impotent. namely. I do not know whether this statement is accurate. and that these are visited by many hive-bees. until at last a complete separation of the sexes might be effected. to be a common plant. Having found a female tree exactly sixty yards from a male tree. as he expresses it. too slight to be appreciated by us. No naturalist doubts the advantage of what has been called the "physiological division of labour. individuals with this tendency more and more increased. as soon as the plant had been rendered so highly attractive to insects that pollen was regularly carried from flower to flower. Bearing such facts in mind. we may suppose the plant of which we have been slowly increasing the nectar by continued selection. now if we suppose this to occur in ever so slight a degree under nature. but I may add that some of the species of holly in North America are. and on all. would be continually favoured or selected. are more or less dioeciously polygamous. which determines the visits of the hive-bee. I put the stigmas of twenty flowers. In plants under culture and placed under new conditions of life. Let us now turn to the nectar-feeding insects. which is visited by humble-bees alone. But to return to our imaginary case. which with a very little more trouble they can enter by the mouth. yet the hive-bee can easily suck the nectar out of the incarnate clover. for I have been assured that when red clover has been mown. That this nectar is much liked by the hive-bee is certain. and thus the communities to which they belonged would flourish and throw off many swarms inheriting the same peculiarities. many hive-bees sucking the flowers through holes bitten in the base of the tube by humble bees.pistil. through dimorphism and other means. might profit a bee or other insect. nevertheless every female flower which I examined had been effectually fertilised by the bees. etc. and four stamens with shrivelled anthers.

or of any great and sudden modification in their structure. and a vast majority of plants are hermaphrodites. I must trust to some general considerations alone. What reason. In the case of animals and plants with separated sexes. is there for supposing in these cases that two individuals ever concur in reproduction? As it is impossible here to enter on details. in accordance with the almost universal belief of breeders.bee. Thus. This view was long ago doubtfully suggested by Sprengel. two individuals regularly unite for reproduction. I must here introduce a short digression. it is of course obvious that two individuals must always (with the exception of the curious and not well understood cases of parthenogenesis) unite for each birth." but we now seldom hear the agencies which we see still at work. pair for each birth. All vertebrate animals. if humble-bees were to become rare in any country. as the fertility of this clover absolutely depends on bees visiting the flowers. Thus I can understand how a flower and a bee might slowly become. and on the other hand. that these facts alone incline me to believe that it is a general law of nature that no organic being fertilises itself for a perpetuity . it might be a great advantage to the plant to have a shorter or more deeply divided corolla. it might be a great advantage to the hive-bee to have a slightly longer or differently constructed proboscis. though I have the materials prepared for an ample discussion. is able to reach and suck the nectar of the red clover. that CLOSE interbreeding diminishes vigour and fertility. ON THE INTERCROSSING OF INDIVIDUALS. but in the case of hermaphrodites this is far from obvious. in a country where this kind of clover abounded. I have collected so large a body of facts. either occasionally or habitually. by the continued preservation of all the individuals which presented slight deviations of structure mutually favourable to each other. so will natural selection banish the belief of the continued creation of new organic beings. when used in explaining the excavation of the deepest valleys or the formation of long lines of inland cliffs. and which freely crosses with it. concur for the reproduction of their kind. and as modern geology has almost banished such views as the excavation of a great valley by a single diluvial wave. which is all that concerns us. spoken of as trifling and insignificant. modified and adapted to each other in the most perfect manner. but I must here treat the subject with extreme brevity. I am well aware that this doctrine of natural selection. and made so many experiments. Natural selection acts only by the preservation and accumulation of small inherited modifications. as illustrative of geology. or between individuals of the same variety but of another strain. Nevertheless there is reason to believe that with all hermaphrodites two individuals. all insects and some other large groups of animals. On the other hand. that is. In the first place. Modern research has much diminished the number of supposed hermaphrodites and of real hermaphrodites a large number pair. We shall presently see its importance. it may be asked. Knight and Kolreuter. which is generally considered a mere variety of the common hive-bee. each profitable to the preserved being. showing. either simultaneously or one after the other. that with animals and plants a cross between different varieties. But still there are many hermaphrodite animals which certainly do not habitually pair. so that the hive-bees should be enabled to suck its flowers. exemplified in the above imaginary instances. gives vigour and fertility to the offspring. is open to the same objections which were first urged against Sir Charles Lyell's noble views on "the modern changes of the earth.

How strange are these facts! How strange that the pollen and stigmatic surface of the same flower. and it is sufficient. Every hybridizer knows how unfavourable exposure to wet is to the fertilisation of a flower. on the other hand. have their organs of fructification closely enclosed. to ensure fertilisation. which is visited by bees. the influence of the foreign pollen. the contrivance seems adapted solely to ensure self-fertilisation. yet. So it is with the reciprocally dimorphic and trimorphic plants previously alluded to. but it must not be supposed that bees would thus produce a multitude of hybrids between distinct species. before the stigma of that individual flower is ready to receive them. as Sprengel. I think. and not to carry pollen from one to the other. In very many other cases. or the stigma is ready before the pollen of that flower is ready. and in this very genus. yet what a multitude of flowers have their anthers and stigmas fully exposed to the weather! If an occasional cross be indispensable. though there is no special mechanical contrivance to prevent the stigma receiving pollen from the same flower. in Lobelia fulgens. though by placing pollen from one flower on the stigma of another. so largely do they naturally cross. and no doubt it is useful for this end: but the agency of insects is often required to cause the stamens to spring forward. though placed so close together. the fullest freedom for the entrance of pollen from another individual will explain the above state of exposure of the organs. When the stamens of a flower suddenly spring towards the pistil. and as this flower is never visited. by insects. for if a plant's own pollen and that from another species are placed on the same stigma. and as I can confirm. as well as from my own observations: for instance. far from self-fertilisation being favoured. which on any other view are inexplicable. as in the great papilionaceous or pea-family. Now. it is hardly possible to raise pure seedlings. as has been shown by Gartner. at least in my garden.of generations. which seems to have a special contrivance for self-fertilisation. such as the following. So necessary are the visits of bees to many papilionaceous flowers. and more recently Hildebrand and others have shown. to the great good of the plant. understand several large classes of facts. we can. the former is so prepotent that it invariably and completely destroys. On the belief that this is a law of nature. In numerous other cases. there is a really beautiful and elaborate contrivance by which all the infinitely numerous pollen-granules are swept out of the conjoined anthers of each flower. it is scarcely possible for insects to fly from flower to flower. as if for the very purpose of . but that a cross with another individual is occasionally--perhaps at long intervals of time--indispensable. it never sets a seed. Another species of Lobelia. Many flowers. if closely-allied forms or varieties are planted near each other. that their fertility is greatly diminished if these visits be prevented. or slowly move one after the other towards it. I raise plenty of seedlings. there are special contrivances which effectually prevent the stigma receiving pollen from its own flower. as I could show from the works of Sprengel and others. but these almost invariably present beautiful and curious adaptations in relation to the visits of insects. just to touch with the same brush the anthers of one flower and then the stigma of another. Insects act like a camel-hair pencil. as Kolreuter has shown to be the case with the barberry. notwithstanding that the plant's own anthers and pistil stand so near each other as almost to ensure self-fertilisation. so that these so-named dichogamous plants have in fact separated sexes. and must habitually be crossed. either the anthers burst before the stigma is ready for fertilisation. seeds freely in my garden. it is well known that.

for owing to the nature of the fertilising element there are no means. and the pollen of each flower readily gets on its stigma without insect agency. Hooker tabulated the trees of New Zealand. Cirripedes long appeared to me to present. When the sexes are separated. and the occasional influence of a distinct individual. and the result was as I anticipated. but that nature has largely provided against it by giving to trees a strong tendency to bear flowers with separated sexes. such as the land-mollusca and earth-worms. I find to be the case in this country. mongrels: for instance. Yet the pistil of each cabbage-flower is surrounded not only by its own six stamens but by those of the many other flowers on the same plant. When distinct SPECIES are crossed the case is reversed. although the male and female flowers may be produced on the same tree. radish. can be shown to be physically impossible. then. In the case of a large tree covered with innumerable flowers. which offers so strong a contrast with terrestrial plants. On the other hand. it may be objected that pollen could seldom be carried from tree to tree. and some even of these were not perfectly true. comes it that such a vast number of the seedlings are mongrelized? It must arise from the pollen of a distinct VARIETY having a prepotent effect over the flower's own pollen. as I found. and of these only 78 were true to their kind. I raised 233 seedling cabbages from some plants of different varieties growing near each other. and that this is part of the general law of good being derived from the intercrossing of distinct individuals of the same species. and this will give a better chance of pollen being occasionally carried from tree to tree. and Dr. and at most only from flower to flower on the same tree. be allowed to seed near each other. As yet I have not found a single terrestrial animal which can fertilise itself. a large majority of the seedlings thus raised turn out. I have made these few remarks on trees simply to call attention to the subject. is intelligible on the view of an occasional cross being indispensable. but here the currents of water offer an obvious means for an occasional cross. Professor Huxley. and at my request Dr. and of some other plants. namely. and flowers on the same tree can be considered as distinct individuals only in a limited sense. but these all pair. to discover a single hermaphrodite animal with the organs of reproduction so perfectly enclosed that access from without. Dr. should be in so many cases mutually useless to each other! How simply are these facts explained on the view of an occasional cross with a distinct individual being advantageous or indispensable! If several varieties of the cabbage. I have as yet failed. How. Turning for a brief space to animals: various terrestrial species are hermaphrodites. for I have found that plants carefully protected from insects produce the full number of pods.self-fertilisation. This remarkable fact. there are many self-fertilising hermaphrodites. under this point of view. Of aquatic animals. analogous to the action of insects and of the wind with plants. Hooker informs me that the rule does not hold good in Australia: but if most of the Australian trees are dichogamous. As in the case of flowers. but to this subject we shall return in a future chapter. but I have been . for a plant's own pollen is always prepotent over foreign pollen. a case of great difficulty. I believe this objection to be valid. That trees belonging to all orders have their sexes more often separated than other plants. after consultation with one of the highest authorities. the same result would follow as if they bore flowers with separated sexes. pollen must be regularly carried from flower to flower. by which an occasional cross could be effected with terrestrial animals without the concurrence of two individuals. Asa Gray those of the United States. onion.

inhabiting intermediate districts. it appears that with animals and plants an occasional intercross between distinct individuals is a very general. will evidently be favourable. all hermaphrodites do occasionally intercross. if not universal. From these several considerations and from the many special facts which I have collected. With hermaphrodite organisms which cross only occasionally. why should it prevail against natural selection? In the case of methodical selection. for within a confined area. to prove that two individuals. with some place in the natural polity not perfectly occupied. and which do not breed at a very quick rate. under which term individual differences are always included. and this I find to be the case. some species of the same family and even of the same genus. the difference between them and unisexual species is. Thus it will be under nature. The tendency to reversion may often check or prevent the work. varieties will generally be confined to separated countries. A large number of individuals. and is. but which wander little and can increase at a rapid rate. a highly important element of success. and might there . for as all organic beings are striving to seize on each place in the economy of nature. and if the individuals be allowed freely to intercross. will compensate for a lesser amount of variability in each individual. have a nearly common standard of perfection. if the same species undergoes modification in different districts. A great amount of variability. both with animals and plants. and likewise with animals which unite for each birth. by giving a better chance within any given period for the appearance of profitable variations. But if. will in the long run generally be supplanted by one of the adjoining varieties. but as this tendency has not prevented man from forming by selection numerous domestic races. notwithstanding that there is no separation of selected individuals.enabled. But if the area be large. though both are self-fertilising hermaphrodites. though agreeing closely with each other in their whole organisation. Though nature grants long periods of time for the work of natural selection. Hence with animals of this nature. though in different degrees. are hermaphrodites. I believe. will tend to be preserved. its several districts will almost certainly present different conditions of life. But when many men. she does not grant an indefinite period. and some unisexual. by a fortunate chance. very small. without intending to alter the breed. as far as function is concerned. law of nature. This is an extremely intricate subject. and then. a new and improved variety might be quickly formed on any one spot. Unless favourable variations be inherited by some at least of the offspring. improvement surely but slowly follows from this unconscious process of selection. nothing can be effected by natural selection. Intercrossing will chiefly affect those animals which unite for each birth and wander much. It must have struck most naturalists as a strange anomaly that. CIRCUMSTANCES FAVOURABLE FOR THE PRODUCTION OF NEW FORMS THROUGH NATURAL SELECTION. But we shall see in the sixth chapter that intermediate varieties. the newly formed varieties will intercross on the confines of each. all the individuals varying in the right direction. but which I am unable here to give. if any one species does not become modified and improved in a corresponding degree with its competitors it will be exterminated. in fact. do sometimes cross. his work will completely fail. a breeder selects for some definite object. and all try to procure and breed from the best animals. for instance birds.

The importance of isolation is likewise great in preventing. isolation will give time for a new variety to be improved at a slow rate. after any physical change in the conditions. If the conditions of life change and the form undergoes modification. either from being surrounded by barriers. we must not assume that free intercrossing would always eliminate the effects of natural selection. the young thus produced will gain so much in vigour and fertility over the offspring from long-continued self-fertilisation. which do not propagate sexually. It will obviously thus act far more efficiently with those animals which unite for each birth. and thus in the long run the influence of crosses. however. In a confined or isolated area. Intercrossing plays a very important part in nature by keeping the individuals of the same species. uniformity of character can be given to the modified offspring. from breeding at slightly different seasons. Isolation also is an important element in the modification of species through natural selection. etc. and which do not propagate rapidly. either for or against natural selection. only through the principle of inheritance. and this will retard the production of new species through natural selection. that they will have a better chance of surviving and propagating their kind. the total number of the inhabitants will be small. uniformity of character can be retained by them under the same conditions of life. as the chance of intercrossing is thus lessened. Lastly. an isolated area be very small. and has shown that the service rendered by isolation in preventing crosses between newly-formed varieties is probably greater even than I supposed. With respect to organic beings extremely low in the scale. for I can bring forward a considerable body of facts showing that within the same area two varieties of the same animal may long remain distinct.maintain itself in a body and afterward spread. so that the individuals of the new variety would chiefly cross together. from haunting different stations. But from reasons already assigned I can by no means agree with this naturalist. will be great. On this principle nurserymen always prefer saving seed from a large body of plants. and through natural selection which will destroy any individuals departing from the proper type. Even if these take place only at long intervals of time. and which cannot possibly intercross. the organic and inorganic conditions of life will generally be almost uniform. such as of climate. by decreasing the chances of favourable variations arising. we have reason to believe that occasional intercrosses take place with all animals and plants. nor conjugate. If. elevation of the land. if not very large. or of the same variety.. that migration and isolation are necessary elements for the formation of new species. or from the individuals of each variety preferring to pair together. even at rare intervals. and thus new places in the natural economy of the district will be left open to be filled up by the modification of the old inhabitants. I state this because it has been erroneously asserted . The mere lapse of time by itself does nothing. Intercrossing with the inhabitants of the surrounding districts. solely by natural selection preserving similar favourable variations. and this may sometimes be of much importance. the immigration of better adapted organisms. will also be thus prevented. so that natural selection will tend to modify all the varying individuals of the same species in the same manner. as already stated. true and uniform in character. or from having very peculiar physical conditions. Even with animals which unite for each birth. Moritz Wagner has lately published an interesting essay on this subject. but.

we can understand how it is that the flora of Madeira. In accordance with this view. They will thus play a more important part in the changing history of the organic world. the competition between fresh water productions will have been less severe than elsewhere. that it gives a better chance of beneficial variations arising and of their being selected. owing to former oscillations of level. and fixed. and its importance in this respect is great. It likewise tends to increase the direct action of the physical conditions of life. and will give rise to the greatest number of new varieties and species. arising from the large number of individuals of the same species there supported. especially for the production of species which shall prove capable of enduring for a long period. to a certain extent. we ought to make the comparison within equal times. understand some facts which will be again alluded to in our chapter on Geographical Distribution. although small isolated areas have been in some respects highly favourable for the production of new species. resembles to a certain extent the extinct tertiary flora of Europe. also. the fact of the productions of the smaller continent of Australia now yielding before those of the larger Europaeo-Asiatic area. which already have been victorious over many competitors. so that the good effects of isolation will generally. or they will be exterminated. and will thus come into competition with many other forms. Thus. have concurred. yet that the course of modification will generally have been more rapid on large areas. on the whole I am inclined to believe that largeness of area is still more important. On a small island. and if some of these many species become modified and improved.--that is. and of spreading widely. Consequently. and there will have been less modification and less extermination. has been most favourable for the production of new organic forms. or a large open area like a continent. accumulated. Although isolation is of great importance in the production of new species. as we shall see in our chapter on Geographical Distribution. taken together. Each new form. according to Oswald Heer. If we turn to nature to test the truth of these remarks.that the element of time has been assumed by me to play an all-important part in modifying species. not only will there be a better chance of favourable variations. But we may thus deceive ourselves. and look at any small isolated area. such as an oceanic island. will be able to spread over the open and continuous area. for instance. the race for life will have been less severe. although the number of the species inhabiting it is small. perhaps. Lapse of time is only so far important. others will have to be improved in a corresponding degree. as soon as it has been much improved. have been produced there and nowhere else in the world. Moreover. also. it is that continental productions have everywhere become so largely naturalised on islands. great areas. Hence an oceanic island at first sight seems to have been highly favourable for the production of new species. though now continuous. I conclude that. we can. have existed in a broken condition. yet of these species a very large proportion are endemic. will often. for to ascertain whether a small isolated area. Hence. and this we are incapable of doing. All fresh water basins. as if all the forms of life were necessarily undergoing change through some innate law. that the new forms produced on large areas. and what is more important. Throughout a great and open area. make a small area compared with that of the sea or of the land. and old . new forms will have been more slowly produced. but the conditions of life are much more complex from the large number of already existing species. in relation to the constitution of each organism. Finally. will be those that will spread most widely.

intermittent results accord well with what geology tells us of the rate and manner at which the inhabitants of the world have changed. When. and only on a few of the inhabitants of the same region. to the beauty and complexity of the coadaptations between all organic beings. and will have been subjected to severe competition. competition. and time will have been allowed for the varieties in each to become well modified and perfected. and this will create new places. and thus to produce new species. I further believe that these slow. . as the Ornithorhynchus and Lepidosiren. and again there will have been a fair field for natural selection to improve still further the inhabitants. And it is in fresh water basins that we find seven genera of Ganoid fishes. While the area existed as a continent the inhabitants will have been numerous in individuals and kinds.forms more slowly exterminated. so that new places in the polity of each island will have had to be filled up by the modification of the old inhabitants. Many will exclaim that these several causes are amply sufficient to neutralise the power of natural selection. they have endured to the present day. only at long intervals of time. will have been the most favourable for the production of many new forms of life. These anomalous forms may be called living fossils. which has undergone many oscillations of level. like fossils. Slow though the process of selection may be. The occurrence of such places will often depend on physical changes. there will again have been very severe competition. one with another and with their physical conditions of life. To sum up. but all this will take place very slowly. which. and on the immigration of better adapted forms being prevented. it would often be long before differences of the right nature in various parts of the organisation might occur. as far as the extreme intricacy of the subject permits. connect to a certain extent orders at present widely separated in the natural scale. That natural selection generally act with extreme slowness I fully admit. and therefore less severe. if feeble man can do much by artificial selection. from having inhabited a confined area. The result would often be greatly retarded by free intercrossing. I do not believe so. fitted to endure for a long time and to spread widely. But I do believe that natural selection will generally act very slowly. I conclude that for terrestrial productions a large continental area. which generally take place very slowly. the most favoured or improved varieties will have been enabled to spread. there will have been much extinction of the less improved forms. remnants of a once preponderant order: and in fresh water we find some of the most anomalous forms now known in the world. When converted by subsidence into large separate islands there will still have existed many individuals of the same species on each island: intercrossing on the confines of the range of each new species will have been checked: after physical changes of any kind immigration will have been prevented. ready to be filled up by better adapted forms. It can act only when there are places in the natural polity of a district which can be better occupied by the modification of some of its existing inhabitants. and from having been exposed to less varied. As some few of the old inhabitants become modified the mutual relations of others will often be disturbed. the islands were reconverted into a continental area. and the relative proportional numbers of the various inhabitants of the reunited continent will again have been changed. by renewed elevation. Although all the individuals of the same species differ in some slight degree from each other. I can see no limit to the amount of change. the circumstances favourable and unfavourable for the production of new species through natural selection.

The principle. several important facts. In Yorkshire. We have evidence of this. But we may go further than this. and explains. geology plainly tells us. will generally press hardest on its nearest kindred. others will become rarer and rarer. during the progress of its formation. take the place of older and inferior kinds. and we shall presently attempt to show why it is that the number of species throughout the world has not become immeasurably great. which I have designated by this term.which may have been effected in the long course of time through nature's power of selection. The forms which stand in closest competition with those undergoing modification and improvement. during great fluctuations in the nature or the seasons. which consequently endure. and it follows from this. Rarity.--which. EXTINCTION CAUSED BY NATURAL SELECTION. is the precursor to extinction. Owing to the high geometrical rate of increase of all organic beings. through the selection of improved forms by man. that as new species in the course of time are formed through natural selection. Many curious instances could be given showing how quickly new breeds of cattle. And we have seen in the chapter on the Struggle for Existence that it is the most closely-allied forms. Hence. in the facts stated in the second chapter. sheep and other animals. many old forms must become extinct. and finally extinct. though having somewhat of the character of species--as is shown by the hopeless doubts . That the number of specific forms has not indefinitely increased. that is by the survival of the fittest. rare species will be less quickly modified or improved within any given period. so. In the first place. and that these "were swept away by the short-horns" (I quote the words of an agricultural writer) "as if by some murderous pestilence. We see the same process of extermination among our domesticated productions. varieties. We can see that any form which is represented by few individuals will run a good chance of utter extinction. and tend to exterminate them. from having nearly the same structure. generally. each area is already fully stocked with inhabitants. Consequently. but it must here be alluded to from being intimately connected with natural selection. will the less favoured decrease and become rare. and varieties of flowers. showing that it is the common and diffused or dominant species which offer the greatest number of recorded varieties. or from a temporary increase in the number of its enemies. as geology tells us.--varieties of the same species. as I believe. We have seen that the species which are most numerous in individuals have the best chance of producing favourable variations within any given period. is of high importance. will naturally suffer most. and species of the same genus or related genera. Natural selection acts solely through the preservation of variations in some way advantageous. it is historically known that the ancient black cattle were displaced by the long-horns. they will consequently be beaten in the race for life by the modified and improved descendants of the commoner species. This subject will be more fully discussed in our chapter on Geology. constitution and habits. unless we admit that specific forms can go on indefinitely increasing in number. even strongly-marked ones." DIVERGENCE OF CHARACTER. From these several considerations I think it inevitably follows. for as new forms are produced. that as the favoured forms increase in number. each new variety or species. generally come into the severest competition with each other.

" they both go on (as has actually occurred with the sub-breeds of the tumbler-pigeon) choosing and breeding from birds with longer and longer beaks. and on the acknowledged principle that "fanciers do not and will not admire a medium standard. it may be asked. both from each other and from their common parent. for instance. being neither very swift nor very strong. If its natural power of increase be allowed to act. In practice. We shall here find something analogous. Nevertheless according to my view.in many cases how to rank them--yet certainly differ far less from each other than do good and distinct species. race and cart horses. How. varieties are species in the process of formation. the differences would become greater. or with shorter and shorter beaks. Take the case of a carnivorous quadruped. the several breeds of pigeons. The early differences would be very slight. at first barely appreciable. we see in man's productions the action of what may be called the principle of divergence. I have sought light on this head from our domestic productions. Again. Mere chance. the supposed prototypes and parents of future well-marked species. these sub-breeds would become converted into two well-established and distinct breeds. the inferior animals with intermediate characters. As has always been my practice. would not have been used for breeding. can any analogous principle apply in nature? I believe it can and does apply most efficiently (though it was a long time before I saw how). but like extremes. then. we may suppose that at an early period of history. and would be noted as forming two sub-breeds. we must infer from most of the innumerable species throughout nature presenting well-marked differences. etc.. and will thus have tended to disappear. and the offspring of this variety again to differ from its parent in the very same character and in a greater degree. struck by a pigeon having a slightly shorter beak. by so much will they be better enabled to seize on many and widely diversified places in the polity of nature. constitution. does the lesser difference between varieties become augmented into the greater difference between species? That this does habitually happen. while those of another required stronger and bulkier horses. the men of one nation or district required swifter horses. for instance. and of stronger ones in the other. but this alone would never account for so habitual and large a degree of difference as that between the species of the same genus. might cause one variety to differ in some character from its parents. as we may call it. whereas varieties. Here. and the breeds to diverge in character. But how. causing differences. in the course of time. and habits. a fancier is. As the differences became greater. steadily to increase. incipient species. Ultimately after the lapse of centuries. as I have called them. It will be admitted that the production of races so different as short-horn and Hereford cattle. and so be enabled to increase in numbers. of which the number that can be supported in any country has long ago arrived at its full average. present slight and ill-defined differences. could never have been effected by the mere chance accumulation of similar variations during many successive generations. but. from the simple circumstance that the more diversified the descendants from any one species become in structure. from the continued selection of swifter horses in the one case. another fancier is struck by a pigeon having a rather longer beak. it can succeed in increasing (the country not undergoing any change in conditions) only by its varying descendants seizing on places at present occupied by other animals: some of them. We can clearly discern this in the case of animals with simple habits. then. being enabled to feed on new kinds . or are.

a greater number of plants and a greater weight of dry herbage can be raised in the latter than in the former case. some inhabiting new stations. de Candolle has well remarked. with the accompanying differences of habit and constitution. perhaps. But the case is very different. take the rank of species.of prey. So it will be with plants. To give a single . supported twenty species of plants. though in a very slight degree. it is seen. in his great and admirable work. which shows how much these plants differed from each other. For instance. is seen under many natural circumstances. either dead or alive. So it is with the plants and insects on small and uniform islets: also in small ponds of fresh water. when rendered very distinct from each other. Hence. that floras gain by naturalisation. The truth of the principle that the greatest amount of life can be supported by great diversification of structure. and where the contest between individual and individual must be very severe. Consequently. The more diversified in habits and structure the descendants of our carnivorous animals become. would succeed in living on the same piece of ground. which had been exposed for many years to exactly the same conditions. the advantages of diversification of structure. It has been experimentally proved. the more places they will be enabled to occupy. three feet by four in size. proportionally with the number of the native genera and species. and varieties. the most distinct varieties of any one species of grass would have the best chance of succeeding and of increasing in numbers. and the varieties were continually selected which differed from each other in the same manner. if any one species of grass were to go on varying. frequenting water. which thus jostle each other most closely. The same has been found to hold good when one variety and several mixed varieties of wheat have been sown on equal spaces of ground. determine that the inhabitants. a greater number of individual plants of this species. I found that a piece of turf. as do the distinct species and genera of grasses. as it may be said. and thus of supplanting the less distinct varieties. have been expected that naturalised plants would have belonged to a few groups more especially adapted to certain stations in their new homes. including its modified descendants. could live on it (supposing its nature not to be in any way peculiar). Farmers find that they can raise more food by a rotation of plants belonging to the most different orders: nature follows what may be called a simultaneous rotation. if they vary--for otherwise natural selection can effect nothing. far more in new genera than in new species. that where they come into the closest competition. and a similar plot be sown with several distinct genera of grasses. and these belonged to eighteen genera and to eight orders. and Alph. especially if freely open to immigration. and some perhaps becoming less carnivorous. in the course of many thousand generations. Most of the animals and plants which live close round any small piece of ground. but. climbing trees. What applies to one animal will apply throughout all time to all animals--that is. that if a plot of ground be sown with one species of grass. It might also. to the utmost to increase in number. and may be said to be striving to the utmost to live there. The same principle is seen in the naturalisation of plants through man's agency in foreign lands. we always find great diversity in its inhabitants. and is thus striving. for these are commonly looked at as specially created and adapted for their own country. And we know that each species and each variety of grass is annually sowing almost countless seeds. as a general rule. belong to what we call different genera and orders. shall. It might have been expected that the plants which would succeed in becoming naturalised in any land would generally have been closely allied to the indigenes. In an extremely small area.

and rodent mammals. ON THE DESCENDANTS OF A COMMON ANCESTOR. so will a greater number of individuals be capable of there supporting themselves. Let A to L represent the species of a genus large in its own country. on an average more species vary in large genera than in small genera. with their organisation but little diversified. our carnivorous. may . The branching and diverging dotted lines of unequal lengths proceeding from (A). and have there become naturalised. could hardly compete with a set more perfectly diversified in structure. widely-diffused. these species are supposed to resemble each other in unequal degrees." 260 naturalised plants are enumerated. and feebly representing. whether the Australian marsupials. for instance. and these belong to 162 genera. tends to act. No physiologist doubts that a stomach by being adapted to digest vegetable matter alone. Asa Gray's "Manual of the Flora of the Northern United States. we may gain some crude idea in what manner some of the natives would have had to be modified in order to gain an advantage over their compatriots. We have. We thus see that these naturalised plants are of a highly diversified nature. no less than 100 genera are not there indigenous. we see the process of diversification in an early and incomplete stage of development. vary more than do the rare and restricted species. we may assume that the modified descendants of any one species will succeed so much the better as they become more diversified in structure. the same as that of the physiological division of labour in the organs of the same individual body--a subject so well elucidated by Milne Edwards. also. as is so generally the case in nature. as Mr. could successfully compete with these well-developed orders. In the Australian mammals. It may be doubted. They differ. which are the commonest and most widely-diffused. ruminant. for out of the 162 naturalised genera. The accompanying diagram will aid us in understanding this rather perplexing subject. to a large extent. So in the general economy of any land. the more widely and perfectly the animals and plants are diversified for different habits of life. and are thus enabled to encroach on places occupied by other beings. and thus a large proportional addition is made to the genera now living in the United States. Let (A) be a common. After the foregoing discussion. A set of animals. and as is represented in the diagram by the letters standing at unequal distances. which has been much compressed. amounting to new generic differences. and we may at least infer that diversification of structure. The advantage of diversification of structure in the inhabitants of the same region is. or flesh alone. draws most nutriment from these substances. Waterhouse and others have remarked. which are divided into groups differing but little from each other. THE PROBABLE EFFECTS OF THE ACTION OF NATURAL SELECTION THROUGH DIVERGENCE OF CHARACTER AND EXTINCTION.instance: in the last edition of Dr. in fact. seen that the species. would be profitable to them. because as we saw in the second chapter. belonging to a genus large in its own country. I have said a large genus. Now let us see how this principle of benefit being derived from divergence of character. and the varying species of the large genera present a greater number of varieties. and varying species. By considering the nature of the plants or animals which have in any country struggled successfully with the indigenes. moreover. combined with the principles of natural selection and of extinction. from the indigenes.

nor that it goes on continuously. and commonly in nearly the same manner as did their parents. In the diagram the process is represented up to the ten-thousandth generation. they will also partake of those more general advantages which made the genus to which the parent-species belonged. the more places they will be enabled to seize on. the more diversified in structure the descendants from any one species can be rendered. And here the importance of the principle of benefit derived from divergence of character comes in.represent its varying offspring. for natural selection will always act according to the nature of the places which are either unoccupied or not perfectly occupied by other beings. And after this interval. some of the varieties. and the more their modified progeny will increase. Moreover. nor are they all supposed to endure for equal periods. producing only a single variety. and under a condensed and simplified form up to the fourteen-thousandth generation. And all these circumstances are favourable to the production of new varieties. and some failing to produce any. In our diagram . will generally go on increasing in number and diverging in character. it is far more probable that each form remains for long periods unaltered. But as a general rule. When a dotted line reaches one of the horizontal lines. differing from each other. such as would be thought worthy of record in a systematic work. but of the most diversified nature. and may or may not produce more than one modified descendant. Only those variations which are in some way profitable will be preserved or naturally selected. then. a large genus in its own country. they are not supposed all to appear simultaneously. for this will generally lead to the most different or divergent variations (represented by the outer dotted lines) being preserved and accumulated by natural selection. namely a1 and m1. namely m2 and s2. and this will depend on infinitely complex relations. some producing two or three varieties. and then again undergoes modification. Nor do I suppose that the most divergent varieties are invariably preserved: a medium form may often long endure. and is there marked by a small numbered letter. but often after long intervals of time. the most divergent of their variations will generally be preserved during the next thousand generations. will tend to inherit those advantages which made their parent (A) more numerous than most of the other inhabitants of the same country. and the tendency to variability is in itself hereditary. which will. The variations are supposed to be extremely slight. We may continue the process by similar steps for any length of time. The intervals between the horizontal lines in the diagram. Variety m1 is supposed to have produced two varieties. after each thousand generations. a sufficient amount of variation is supposed to have been accumulated to form it into a fairly well-marked variety. may represent each a thousand or more generations. owing to the principle of divergence. After a thousand generations. But I must here remark that I do not suppose that the process ever goes on so regularly as is represented in the diagram. these two varieties be variable. these two varieties. though in itself made somewhat irregular. and more considerably from their common parent (A). These two varieties will generally still be exposed to the same conditions which made their parents variable. If. variety a1 is supposed in the diagram to have produced variety a2. but in a more and more modified condition. being only slightly modified forms. consequently they will likewise tend to vary. Thus the varieties or modified descendants of the common parent (A). species (A) is supposed to have produced two fairly well-marked varieties. differ more from (A) than did variety a1.

the line of succession is broken at regular intervals by small numbered letters marking the successive forms which have become sufficiently distinct to be recorded as varieties. marked by the letters n14 to z14. In any genus. species (A) is supposed to have produced three forms. The other nine species (marked by capital letters) of our original genus. the species which are already very different in character from each other. which. will have come to differ largely. and might have been inserted anywhere. according to the amount of change supposed to be represented between the horizontal lines. it is probable. will tend to partake of the same advantages which made their parent successful in life. and the nearly extreme species (I). all descended from (A). six new species. and the number of modified descendants will not be increased. The modified offspring from the later and more highly improved branches in the lines of descent. and m10. to convert these three forms into doubtful or at least into well-defined species: thus the diagram illustrates the steps by which the small differences distinguishing varieties are increased into the larger differences distinguishing species. without either having given off any fresh branches or races. In a large genus it is probable that more than one species would vary. we get eight species. By continuing the same process for a greater number of generations (as shown in the diagram in a condensed and simplified manner). often take the place of. either two well-marked varieties (w10 and z10) or two species. belonging to a large genus. if all the lines proceeding from (A) were removed. will. In the diagram I have assumed that a second species (I) has produced. and so destroy. and this is shown in the diagram by the dotted lines unequally prolonged upwards. they will generally go on multiplying in number as well as diverging in character: this is represented in the diagram by the several divergent branches proceeding from (A). will generally tend to produce the greatest number of modified descendants. This case would be represented in the diagram. But these breaks are imaginary. from having diverged in character during the successive generations. In the same way the English racehorse and English pointer have apparently both gone on slowly diverging in character from their original stocks. species are multiplied and genera are formed. a10. after intervals long enough to allow the accumulation of a considerable amount of divergent variation. from each other and from their common parent. these three forms may still be only well-marked varieties. After ten thousand generations. marked by the letters between a14 and m14. as those which have largely varied. although the amount of divergent modification may have been augmented. After fourteen thousand generations. as I believe. by analogous steps. Thus. f10. but perhaps unequally. may for long but unequal periods continue to transmit unaltered descendants. and have given rise to new varieties and species. . for these will have the best chance of seizing on new and widely different places in the polity of nature: hence in the diagram I have chosen the extreme species (A). are supposed to have been produced. If we suppose the amount of change between each horizontal line in our diagram to be excessively small. after ten thousand generations. excepting that from a1 to a10. but we have only to suppose the steps in the process of modification to be more numerous or greater in amount. In some cases no doubt the process of modification will be confined to a single line of descent. As all the modified descendants from a common and widely-diffused species. the earlier and less improved branches: this is represented in the diagram by some of the lower branches not reaching to the upper horizontal lines.

If. extremely probable that they will have taken the places of. the extreme amount of difference in character between species a14 and z14 will be much greater than that between the most distinct of the original eleven species. H. will probably have inherited some of the same advantages: they have also been modified and improved in a diversified manner at each stage of descent. e14. The new species in our diagram. moreover. and thus exterminated. from having diverged at the first commencement of the process of modification.But during the process of modification. The new species. These two species (A and I). but. will now be fifteen in number. For it should be remembered that the competition will generally be most severe between those forms which are most nearly related to each other in habits. If. b14 and f14. Owing to the divergent tendency of natural selection. Hence very few of the original species will have transmitted offspring to the fourteen-thousandth generation. then. The original species of our genus were supposed to resemble each other in unequal degrees. constitution and structure. will be nearly related one to the other. represented in the diagram. the modified offspring of a species get into some distinct country. and species (I) will be replaced by six (n14 to z14) new species. It seems. C. or become quickly adapted to some quite new station. will have played an important part. not only their parents (A) and (I). from having diverged at an earlier period from a5. and species (I) more to G. but likewise some of the original species which were most nearly related to their parents. there will be a constant tendency in the improved descendants of any one species to supplant and exterminate in each stage of descent their predecessors and their original progenitor. Of the eight descendants from (A) the three marked a14. descended from the original eleven species. than to the others. As in each fully stocked country natural selection necessarily acts by the selected form having some advantage in the struggle for life over other forms. our diagram be assumed to represent a considerable amount of modification. therefore. L. Hence all the intermediate forms between the earlier and later states. will generally tend to become extinct. species (A) being more nearly related to B. fourteen in number at the fourteen-thousandth generation. q14. p14. both may continue to exist. o14. and D than to the other species. will be allied to each other in a widely different manner. another of our principles. We may suppose that only one (F) of the two species (E and F) which were least closely related to the other nine original species. So it probably will be with many whole collateral lines of descent. as is so generally the case in nature. But we may go further than this. . will be in some degree distinct from the three first-named species. which will be conquered by later and improved lines. so as to have become adapted to many related places in the natural economy of their country. being replaced by eight new species (a14 to m14). has transmitted descendants to this late stage of descent. that is between the less and more improved states of a the same species. so that they must originally have had some advantage over most of the other species of the genus. K. will be widely different from the other five species. and m14. and lastly. Their modified descendants. will be nearly related from having recently branched off from a10. in which offspring and progenitor do not come into competition. and may constitute a sub-genus or a distinct genus. however. as well as the original parent-species itself. namely that of extinction. species (A) and all the earlier varieties will have become extinct. were also supposed to be very common and widely diffused species.

intermediate in character between existing groups. but to have retained the form of (F). as now explained. In this case its affinities to the other fourteen new species will be of a curious and circuitous nature. for the extinct species lived at various remote epochs when the branching lines of descent had diverged less. will form three very distinct genera. yet are often. But as the original species (I) differed largely from (A). or orders. but each may represent a million or more generations. We shall also have two very distinct genera descended from (I). which. are supposed to have gone on diverging in different directions. from two or more species of the same genus. and we can understand this fact. in some degree. according to the amount of divergent modification supposed to be represented in the diagram. the supposed progenitor of our several new sub-genera and genera. or genera. we suppose the amount of change represented by each successive group of diverging dotted lines to be great. I see no reason to limit the process of modification. but rather between types of the two groups. extinct. also (and this is a very important consideration). owing to inheritance alone. and I think we shall then see that the diagram throws light on the affinities of extinct beings. We shall. it may also represent a section of the successive strata of the earth's crust including extinct remains. the forms marked a14 to p14. the two groups. the six descendants from (I) will. now supposed to be extinct and unknown. though generally belonging to the same orders. And the two or more parent-species are supposed to be descended from some one species of an earlier genus. and the eight descendants from (A). These two groups of genera will thus form two distinct families. in the diagram. and every naturalist will be able to call such cases before his mind. In the diagram each horizontal line has hitherto been supposed to represent a thousand generations.The six descendants from (I) will form two sub-genera or genera. with those now living. and have left no descendants. that two or more genera are produced by descent with modification. differing widely from the descendants of (A). And the two new families. But as these two groups have gone on diverging in character from the type of their parents. as I believe. and these are supposed to be descended from some still more ancient and unknown form. converging in sub-branches downwards towards a single point. The intermediate species. Hence the six new species descended from (I). differ considerably from the eight descendants from (A). standing nearly at the extreme end of the original genus. moreover. have to refer again to this subject. to the formation of genera alone. In our diagram this is indicated by the broken lines beneath the capital letters. . If. or even as distinct sub-families. Being descended from a form that stood between the parent-species (A) and (I). have all become. and those marked o14 to m14. will have to be ranked as very distinct genera. the new species (F14) will not be directly intermediate between them. when we come to our chapter on geology. or orders. except (F). which connected the original species (A) and (I). either unaltered or altered only in a slight degree. families. which is supposed not to have diverged much in character. are descended from two species of the original genus. It is worth while to reflect for a moment on the character of the new species F14. this point represents a species. Thus it is. those marked b14 and f14. it will be in some degree intermediate in character between the two groups descended from these two species.

reduce its number. of the species living at any one period. This. for a long period. we can predict that the groups of organic beings which are now large and triumphant. Looking still more remotely to the future. But here we enter on a very intricate subject. a multitude of smaller groups will become utterly extinct. at remote geological periods. yet. families. from branching out and seizing on many new places in the polity of nature. so that the mature animal cannot be called higher than its larva.We have seen that in each country it is the species belonging to the larger genera which oftenest present varieties or incipient species. continue to increase. But which groups will ultimately prevail. the struggle for the production of new and modified descendants will mainly lie between the larger groups. I shall have to return to this subject in the chapter on classification. ON THE DEGREE TO WHICH ORGANISATION TENDS TO ADVANCE. have now become extinct. extremely few of the more ancient species have transmitted descendants to the present day. the later and more highly perfected sub-groups. Looking to the future. as all the descendants of the same species form a class. that is. which have as yet suffered least extinction. as at the present day. and consequently that. as with certain parasitic crustaceans. Among the vertebrata the degree of intellect and an approach in structure to man clearly come into play. in the adult state. namely. One large group will slowly conquer another large group. and. might have been expected. the earth may have been almost as well peopled with species of many genera. and their specialisation for different functions. formerly most extensively developed. Small and broken groups and sub-groups will finally disappear. no man can predict. and the largeness of any group shows that its species have inherited from a common ancestor some advantage in common. we can understand how it is that there exist so few classes in each main division of the animal and vegetable kingdoms. The ultimate result is that each creature tends to become more and more improved in relation to its conditions. it will chiefly act on those which already have some advantage. Hence. which are all trying to increase in number. It might be thought that the amount of change which the various parts and organs pass through in their development from embryo to maturity would suffice as a standard of comparison. according to this view. as I should be inclined to add. but I may add that as. the . as Milne Edwards would express it. will. indeed. Although few of the most ancient species have left modified descendants. for naturalists have not defined to each other's satisfaction what is meant by an advance in organisation. in which several parts of the structure become less perfect. and which are least broken up. This improvement inevitably leads to the gradual advancement of the organisation of the greater number of living beings throughout the world. owing to the continued and steady increase of the larger groups. the amount of differentiation of the parts of the same organic being. Within the same large group. for we know that many groups. will constantly tend to supplant and destroy the earlier and less improved sub-groups. orders and classes. which are beneficial under the organic and inorganic conditions to which each creature is exposed at all periods of life. and thus lessen its chance of further variation and improvement. Natural selection acts exclusively by the preservation and accumulation of variations. and leave no modified descendants. for as natural selection acts through one form having some advantage over other forms in the struggle for existence. we may predict that. but there are cases. Von Baer's standard seems the most widely applicable and the best. extremely few will transmit descendants to a remote futurity. or.

And geology tells us that some of the lowest forms. petals. and here some botanists rank those plants as highest which have every organ. for every naturalist who has dissected some of the beings now ranked as very low in the scale. fully developed in each flower. stamens and pistils. look at the plants which have their several organs much modified and reduced in number as the highest. or the survival of the fittest. among which some naturalists rank those as highest which. whatever the future may reveal. to be highly organised. On the other hand. But to suppose that most of the many now existing low forms have not in the least advanced since the first dawn of life would be extremely rash. Science has not as yet proved the truth of this belief. as the infusoria and rhizopods. And it may be asked what advantage. to fishes. we can see. natural selection clearly leads towards this standard: for all physiologists admit that the specialisation of organs. the amount of differentiation and specialisation of the several organs in each being when adult (and this will include the advancement of the brain for intellectual purposes). Whether organisation on the whole has actually advanced from the remotest geological periods to the present day will be more conveniently discussed in our chapter on Geological Succession. like the sharks. for natural selection. must have been struck with their really wondrous and beautiful organisation. and differ most from the other vertebrate classes. how is it that throughout the world a multitude of the lowest forms still exist. as far as we can see. and hence the accumulation of variations tending towards specialisation is within the scope of natural selection. who believed in an innate and inevitable tendency towards perfection in all organic beings. does not necessarily include progressive development--it only takes advantage of such variations as arise and are beneficial to each creature under its complex relations of life. and might remain for indefinite ages in their present lowly condition. probably with more truth. have remained for an enormous period in nearly their present state. We see still more plainly the obscurity of the subject by turning to plants. whereas other botanists. But it may be objected that if all organic beings thus tend to rise in the scale. But we shall see how obscure this subject is if we look. as sepals. would it be to an infusorian animalcule--to an intestinal worm--or even to an earth-worm. and how is it that in each great class some forms are far more highly developed than others? Why have not the more highly developed forms every where supplanted and exterminated the lower? Lamarck. inasmuch as they are most strictly fish-like. for instance. seems to have felt this difficulty so strongly that he was led to suppose that new and simple forms are continually being produced by spontaneous generation. unimproved or but little improved. If it were no advantage. .completeness of the division of physiological labour. by natural selection. is an advantage to each being. while other naturalists rank the common bony or teleostean fishes as the highest. On our theory the continued existence of lowly organisms offers no difficulty. approach nearest to amphibians. bearing in mind that all organic beings are striving to increase at a high ratio and to seize on every unoccupied or less well occupied place in the economy of nature. that it is quite possible for natural selection gradually to fit a being to a situation in which several organs would be superfluous or useless: in such cases there would be retrogression in the scale of organisation. among which the standard of intellect is of course quite excluded. inasmuch as in this state they perform their functions better. these forms would be left. If we take as the standard of high organisation.

Nearly the same remarks are applicable. marsupials. yet the scale will always present many degrees of perfection. as we shall hereafter see. an anomalous annelid. could the first step in the advancement or differentiation of parts have arisen? Mr. has time sufficed for the utmost possible amount of development. so that warm-blooded mammals when inhabiting the water lie under a disadvantage in having to come continually to the surface to breathe. or of certain members in this class. The three lowest orders of mammals. where they have been subjected to less severe competition. on the whole. co-exist in South America in the same region with numerous monkeys. In some few cases there has been what we must call retrogression or organisation. I believe that many lowly organised forms now exist throughout the world. In some cases variations or individual differences of a favourable nature may never have arisen for natural selection to act on and accumulate. lowly organised forms appear to have been preserved to the present day. speculation on the subject is almost useless. or became attached to any supporting surface. and probably interfere little with each other. Physiologists believe that the brain must be bathed by warm blood to be highly active. to the co-existence of mammals and fish--among mammalia. presented the simplest structure. as I hear from Fritz Muller. in the vertebrata. may have advanced and be still advancing throughout the world. has as sole companion and competitor on the barren sandy shore of South Brazil. if we look to the different grades of organisation within the same great group. the advancement of the whole class of mammals. In no case. and more liable to be put out of order and injured. from various causes. With fishes. to the co-existence of the shark and the lancelet (Amphioxus). an error to suppose that there would be no struggle for existence. Herbert Spencer would probably answer that. no natural selection. until many forms had been produced: variations in a single species inhabiting an isolated station might be beneficial. to the highest grade would not lead to their taking the place of fishes. no one ought to feel surprise at much remaining as yet unexplained on the origin of species. it has been asked. and rodents. for instance. But. It is. members of the shark family would not tend to supplant the lancelet. for the high advancement of certain whole classes. Although organisation. consequently. But mammals and fish hardly come into competition with each other. and. edentata. or two distinct forms might arise. But the main cause lies in the fact that under very simple conditions of life a high organisation would be of no service--possibly would be of actual disservice. or of certain members of each class. Finally. and where their scanty numbers have retarded the chance of favourable variations arising. namely. when all organic beings. if we make due allowance for our profound ignorance on the mutual relations of the inhabitants of the world at the present . to the co-existence of man and the ornithorhynchus--among fishes. In some cases. probably. however. But as we have no facts to guide us. for the lancelet. how. Looking to the first dawn of life. as we may believe. and thus the whole mass of individuals might be modified. as being of a more delicate nature. and this requires aerial respiration. which latter fish in the extreme simplicity of its structure approaches the invertebrate classes. as I remarked towards the close of the introduction. his law "that homologous units of any order become differentiated in proportion as their relations to incident forces become different" would come into action. as soon as simple unicellular organisms came by growth or division to be compounded of several cells. from inhabiting confined or peculiar stations. does not at all necessarily lead to the extinction of those groups with which they do not enter into close competition.

on inheritance (in itself a fluctuating element) from innumerable progenitors. etc. Imagine the extreme case of as many species as individuals in England. that from an early part of the tertiary period the number of species of shells. and it is not surprising that dissimilar substances should sometimes assume the same form. and in a still higher degree on the surrounding organisms with which each being has come into competition--and lastly. What then checks an indefinite increase in the number of species? The amount of life (I do not mean the number of specific forms) supported on an area must have a limit. The shape of a crystal is determined solely by the molecular forces. and still more so during past ages. If two species belonging to two distinct though allied genera. and the . and that convergence. together with divergence of character. as it may be called. depending so largely as it does on physical conditions. and thus the descendants of two distinct genera would converge into one. whereas the production of new species must always be slow. should ever afterwards converge so closely as to lead to a near approach to identity throughout their whole organisation. We do not know that even the most prolific area is fully stocked with specific forms: at the Cape of Good Hope and in Australia. As far as mere inorganic conditions are concerned. Mr. Consequently there seems at first no limit to the amount of profitable diversification of structure.time. however. but I fully admit that the mutual relations of organic beings are more important.C. Watson thinks that I have overrated the importance of divergence of character (in which. would tend to make an indefinite number of specific forms. The process of extermination in such cases would be rapid. and therefore no limit to the number of species which might be produced. recurring in widely separated geological formations. which support such an astonishing number of species. all of which have had their forms determined through equally complex relations. he apparently believes).. Mr. and that from the middle part of this same period. CONVERGENCE OF CHARACTER. many European plants have become naturalised. and this depends on the surrounding physical conditions. but with organic beings we should bear in mind that the form of each depends on an infinitude of complex relations. If this had occurred. if an area be inhabited by very many species. But geology shows us. and the balance of evidence is opposed to any such an admission. which had originally differed in a marked manner. But it would in most cases be extremely rash to attribute to convergence a close and general similarity of structure in the modified descendants of widely distinct forms. moisture. and as the number of species in any country goes on increasing. we should meet with the same form. the number of mammals has not greatly or at all increased. namely on the variations which have arisen. has likewise played a part. these being due to causes far too intricate to be followed out--on the nature of the variations which have been preserved or selected. each or nearly each species will be represented by few individuals. had both produced a large number of new and divergent forms. and such species will be liable to extermination from accidental fluctuations in the nature of the seasons or in the number of their enemies. independently of genetic connection. it seems probable that a sufficient number of species would soon become adapted to all considerable diversities of heat. it is conceivable that these might approach each other so closely that they would have all to be classed under the same genus. It is incredible that the descendants of two organisms. Watson has also objected that the continued action of natural selection. the organic conditions of life must become more and more complex. therefore. H.

it would be a most extraordinary fact if no variations had ever occurred useful to each being's own welfare. authors have thought that this comes into play in accounting for the deterioration of the aurochs in Lithuania. I have called natural selection. constitution. When any species becomes very rare. and consequently. and how largely extinction has acted in the world's history. causing an infinite diversity in structure. there are many invaders from different quarters of the globe. and this I am inclined to think is the most important element. season or year. Hooker has recently shown that in the southeast corner of Australia. present within a given period few favourable variations. Sexual selection will also give characters useful to the males alone in their struggles or rivalry with other males. assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life. seed. Among many animals sexual selection will have given its aid to ordinary selection by assuring to the most vigorous and best adapted males the greatest number of offspring. owing to their geometrical rate of increase. and these characters will be transmitted to one sex or to both sexes. where. then. or young as easily as the adult. in most cases. Rare species. If under changing conditions of life organic beings present individual differences in almost every part of their structure. How much weight to attribute to these several considerations I will not pretend to say. will. Alph. apparently. and habits. It leads to the improvement of each creature in relation to its organic and inorganic conditions of life. the process of giving birth to new specific forms would thus be retarded. low and simple forms will long endure if well fitted for their simple conditions of life. on the principal often explained. to be advantageous to them. in the same manner as so many variations have occurred useful to man. consequently they will tend to supplant and exterminate several species in several areas. but conjointly they must limit in each country the tendency to an indefinite augmentation of specific forms. a severe struggle for life at some age. But if variations useful to any organic being ever do occur. according to the form of inheritance which prevails. But we have already seen how it entails extinction. Dr. SUMMARY OF CHAPTER. and thus check the inordinate increase of specific forms throughout the world. these will tend to produce offspring similarly characterised. a dominant species.first severe winter or very dry summer would exterminate thousands on thousands of species. on the principle of qualities being inherited at corresponding ages. and each species will become rare if the number of species in any country becomes indefinitely increased. close interbreeding will help to exterminate it. must be judged by the general tenour and balance of evidence given in the following chapters. of red deer in Scotland and of bears in Norway. which has already beaten many competitors in its own home. can modify the egg. and this certainly cannot be disputed. geology plainly . considering the infinite complexity of the relations of all organic beings to each other and to their conditions of life. and from the strong principle of inheritance. consequently. de Candolle has shown that those species which spread widely tend generally to spread VERY widely. Nevertheless. Natural selection. and this cannot be disputed. or the survival of the fittest. if there be. the endemic Australian species have been greatly reduced in number. etc. to what must be regarded as an advance in organisation. This principle of preservation. Whether natural selection has really thus acted in adapting the various forms of life to their several conditions and stations. Lastly. will tend to spread and supplant many others.

and which are known to us only in a fossil state. but it is explained through inheritance and the complex action of natural selection. leads to divergence of character and to much extinction of the less improved and intermediate forms of life.declares. as we have seen illustrated in the diagram. orders. and this connexion of the former and present buds by ramifying branches may well represent the classification of all extinct and living species in groups subordinate to groups. when the tree was young. as has just been remarked. were themselves once. and during the incessant struggle of all species to increase in numbers. steadily tend to increase. and to overtop and kill the surrounding twigs and branches. may be explained. now grown into great branches. and to the productions naturalised in foreign lands. and the generally well defined distinctions between the innumerable organic beings in each class throughout the world. also. which vary most. and these tend to transmit to their modified offspring that superiority which now makes them dominant in their own countries. I believe this simile largely speaks the truth. and these into lesser and lesser branches. families. Of the many twigs which flourished when the tree was a mere bush. of which we see proof by looking to the inhabitants of any small spot. The several subordinate groups in any class cannot be ranked in a single file. in the manner which we everywhere behold--namely. and classes. leads to divergence of character. species of distinct genera much less closely related. very few have left living and modified descendants. forming sub-families. and these round other points. and genera related in different degrees. in the same manner as species and groups of species have at all times overmastered other species in the great battle for life. sub-classes. by so much the more can a large number be supported on the area. the better will be their chance of success in the battle for life. Natural selection. but seem clustered round points. . belonging to the larger genera within each class. habits and constitution. varieties of the same species most closely related. As we here and there see a thin. Therefore. entailing extinction and divergence of character. the widely diffused. families. subordinate to groups. and widely ranging species. At each period of growth all the growing twigs have tried to branch out on all sides. The limbs divided into great branches. species of the same genus less closely and unequally related. Natural selection. only two or three. or even of distinct genera. till they equal the greater differences between species of the same genus. and so on in almost endless cycles. yet survive and bear the other branches. so with the species which lived during long-past geological periods. during the modification of the descendants of any one species. The green and budding twigs may represent existing species. the nature of the affinities. for the more organic beings diverge in structure. no explanation would have been possible of this kind of classification. From the first growth of the tree. It is a truly wonderful fact--the wonder of which we are apt to overlook from familiarity--that all animals and all plants throughout all time and space should be related to each other in groups. and these fallen branches of various sizes may represent those whole orders. straggling branch springing from a fork low down in a tree. and those produced during former years may represent the long succession of extinct species. and genera which have now no living representatives. If species had been independently created. forming sections and sub-genera. many a limb and branch has decayed and dropped off. the more diversified the descendants become. On these principles. budding twigs. Thus the small differences distinguishing varieties of the same species. The affinities of all the beings of the same class have sometimes been represented by a great tree. We have seen that it is the common.

which we see throughout nature between various organic beings. branch out and overtop on all sides many a feebler branch. and covers the surface with its ever-branching and beautiful ramifications.and which by some chance has been favoured and is still alive on its summit. rudimentary. There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence. lead to the conclusion that variability is generally related to the conditions of life to which each species has been exposed during several successive generations. and in a lesser degree with those under nature--were due to chance. so by generation I believe it has been with the great Tree of Life. or nearly all. Effects of changed conditions--Use and disuse. The direct action of changed conditions leads to definite or indefinite results. or slight deviations of structure. In all cases there are two factors.. and lowly organised structures variable--Parts developed in an unusual manner are highly variable: specific characters more variable than generic: secondary sexual characters variable--Species of the same genus vary in an analogous manner--Reversions to long-lost characters--Summary. of course is a wholly incorrect expression. LAWS OF VARIATION. CHAPTER V. the individuals become modified in the same way. and these. the nature of the organism. I have hitherto sometimes spoken as if the variations--so common and multiform with organic beings under domestication. which in some small degree connects by its affinities two large branches of life. and all. Forbes asserts that shells at their southern limit. This. and which has apparently been saved from fatal competition by having inhabited a protected station. but it serves to acknowledge plainly our ignorance of the cause of each particular variation. cannot be attributed simply to such action. which fills with its dead and broken branches the crust of the earth. Some authors believe it to be as much the function of the reproductive system to produce individual differences. if vigorous. such as of climate. and we have much fluctuating variability. have acted in a definite manner. In the former case the nature of the organism is such that it yields readily. and when living in shallow water. It is very difficult to decide how far changed conditions. But the fact of variations and monstrosities occurring much more frequently under domestication than under nature. food. As buds give rise by growth to fresh buds. which is much the most important of the two. and indirectly through the reproductive system. In the first chapter I attempted to show that changed conditions act in two ways. and the greater variability of species having wide ranges than of those with restricted ranges. so we occasionally see an animal like the Ornithorhynchus or Lepidosiren. In the latter case the organisation seems to become plastic. But we may safely conclude that the innumerable complex co-adaptations of structure. are more brightly coloured than those of the same species from further north or from a . organs of flight and of vision--Acclimatisation--Correlated variation--Compensation and economy of growth--False correlations--Multiple. as to make the child like its parents. when subjected to certain conditions. combined with natural selection. In the following cases the conditions seem to have produced some slight definite effect: E. etc. directly on the whole organisation or on certain parts alone. and the nature of the conditions.

Again. but who can tell how much of this difference may be due to the warmest-clad individuals having been favoured and preserved during many generations. But when man is the selecting agent. Instances could be given of similar varieties being produced from the same species under external conditions of life as different as can well be conceived. I think there can be no doubt that use in our domestic animals has strengthened and enlarged certain parts. but many animals possess structures which can be best explained by the effects of disuse. although living under the most opposite climates. but it is the will of man which accumulates the variations in certain direction. when growing near the sea-shore. for the conditions determine whether this or that variety shall survive. and how much to the definite action of the conditions of life. for we know not the parent-forms. we cannot tell how much to attribute to the accumulative action of natural selection. not only to cause variability. Thus. either directly or indirectly. we clearly see that the two elements of change are distinct. there is no greater anomaly in nature than a bird that cannot fly. and is exposed to danger from which it cannot escape by flight. The logger-headed duck of South America can only flap along the surface of the water. Such considerations as these incline me to lay less weight on the direct action of the surrounding conditions. of dissimilar varieties being produced under apparently the same external conditions. of species keeping true. it is probable that the nearly wingless condition of several birds. When a variation is of the slightest use to any being. but likewise to include natural selection. and it is this latter agency which answers to the survival of the fittest under nature. due to causes of which we are quite ignorant. can fly. In one sense the conditions of life may be said. EFFECTS OF THE INCREASED USE AND DISUSE OF PARTS. and Wollaston is convinced that residence near the sea affects the colours of insects. but it . while the adults have lost this power. and that such modifications are inherited. and. or not varying at all. innumerable instances are known to every naturalist. have their leaves in some degree fleshy. it is well known to furriers that animals of the same species have thicker and better fur the further north they live.greater depth. than when living near the coast or on islands. according to Mr. Under free nature we have no standard of comparison by which to judge of the effects of long-continued use or disuse. The ostrich indeed inhabits continents. though not elsewhere fleshy. As the larger ground-feeding birds seldom take flight except to escape danger. and disuse diminished them. As Professor Owen has remarked. variability is in some manner excited. but this certainly does not always hold good. AS CONTROLLED BY NATURAL SELECTION. now inhabiting or which lately inhabited several oceanic islands. From the facts alluded to in the first chapter. yet there are several in this state. than on a tendency to vary. Moquin-Tandon gives a list of plants which. and has its wings in nearly the same condition as the domestic Aylesbury duck: it is a remarkable fact that the young birds. Gould believes that birds of the same species are more brightly coloured under a clear atmosphere. Mr. and how much to the action of the severe climate? For it would appear that climate has some direct action on the hair of our domestic quadrupeds. has been caused by disuse. These slightly varying organisms are interesting in as far as they present characters analogous to those possessed by the species which are confined to similar conditions. tenanted by no beasts of prey. Cunningham. on the other hand.

and thus destroyed. their wings not at all reduced. namely. that the proportion of wingless beetles is larger on the exposed Desertas than in Madeira itself. but the remarkable cases observed by Brown-Sequard in guinea-pigs. therefore the tarsi cannot be of much importance or be much used by these insects. that certain large groups of beetles. and their rudimentary condition in some other genera. the tendency of natural selection to enlarge or to reduce the wings. that the beetles in Madeira. In some cases we might easily put down to disuse modifications of structure which are wholly. or by giving up the attempt and rarely or never flying. are here almost entirely absent. no less than twenty-three have all their species in this condition! Several facts. We may believe that the progenitor of the ostrich genus had habits like those of the bustard. Kirby has remarked (and I have observed the same fact) that the anterior tarsi. on the other hand. this must happen early in life. of the twenty-nine endemic genera. so strongly insisted on by Mr. which absolutely require the use of their wings. but as due to the effects of long-continued disuse. he examined seventeen specimens in his own collection. until they became incapable of flight. as efficiently as many quadrupeds. by kicking its enemies. have. elsewhere excessively numerous. The insects in Madeira which are not ground-feeders. but even enlarged. The evidence that accidental mutilations can be inherited is at present not decisive. and. should make us cautious in denying this tendency. In the Onites apelles the tarsi are so habitually lost that the insect has been described as not having them.can defend itself. Wollaston. For during many successive generations each individual beetle which flew least. those beetles which most readily took to flight would oftenest have been blown to sea. Wollaston. As with mariners shipwrecked near a coast. For when a new insect first arrived on the island. of the inherited effects of operations. This is quite compatible with the action of natural selection. its legs were used more and its wings less. it will perhaps be safest to look at the entire absence of the anterior tarsi in Ateuchus. These several considerations make me believe that the wingless condition of so many Madeira beetles is mainly due to the action of natural selection. and that. lie much concealed. and not one had even a relic left. and which. until the wind lulls and the sun shines. are so far deficient in wings that they cannot fly. they are totally deficient. for as many dung-feeding beetles are generally found with their tarsi lost. Wollaston suspects. whereas it would have been better for the bad swimmers if they had not been able to swim . or mainly due to natural selection. but in a rudimentary condition. Mr. In some other genera they are present. out of the 550 species (but more are now known) inhabiting Madeira. as the size and weight of its body were increased during successive generations. it would have been better for the good swimmers if they had been able to swim still further. as certain flower-feeding coleoptera and lepidoptera. will have had the best chance of surviving from not being blown out to sea. either from its wings having been ever so little less perfectly developed or from indolent habit. Wollaston has discovered the remarkable fact that 200 beetles. must habitually use their wings to gain their subsistence. as Mr. that beetles in many parts of the world are very frequently blown to sea and perish. or feet. combined probably with disuse. In the Ateuchus or sacred beetle of the Egyptians. would depend on whether a greater number of individuals were saved by successfully battling with the winds. as observed by Mr. of many male dung-feeding beetles are often broken off. not as cases of inherited mutilations. and that. Hence. and especially the extraordinary fact.

namely. the cave-rat (Neotoma). slowly migrated by successive generations from the outer world into the deeper and deeper recesses of the Kentucky caves. though the eye is gone. belonging to the most different classes. Animals not far remote from ordinary forms. and these animals. as I am informed by Professor Silliman. such as an . which inhabit the caves of Carniola and Kentucky. By the time that an animal had reached. very close similarity in their organisation and affinities might have been expected. is even more subterranean in its habits than the mole. though useless. but aided perhaps by natural selection. or Ctenomys. as Schiodte remarks: "We accordingly look upon the subterranean faunas as small ramifications which have penetrated into the earth from the geographically limited faunas of the adjacent tracts. having in most cases ordinary powers of vision. This is certainly not the case if we look at the two whole faunas. the cause. could be in any way injurious to animals living in darkness. disuse will on this view have more or less perfectly obliterated its eyes. It is difficult to imagine conditions of life more similar than deep limestone caverns under a nearly similar climate. for. the deepest recesses. and in some cases are quite covered by skin and fur. having been inflammation of the nictitating membrane. after having been exposed for about a month to a graduated light. In one of the blind animals. as did European animals into the caves of Europe. a reduction in their size. but to distinct species. and if so. who had often caught them. are blind. that they were frequently blind. Schiodte has remarked: "We are accordingly prevented from considering the entire phenomenon in any other light than something purely local. and the similarity which is exhibited in a few forms between the Mammoth Cave (in Kentucky) and the caves in Carniola. This state of the eyes is probably due to gradual reduction from disuse. As it is difficult to imagine that eyes. One which I kept alive was certainly in this condition. last of all. with the adhesion of the eyelids and growth of fur over them. the eyes were lustrous and of large size. In South America. the stand for the telescope is there. might in such case be an advantage." These remarks of Schiodte's it should be understood. We have some evidence of this gradation of habit. and. those destined for total darkness. a burrowing rodent. prepare the transition from light to darkness. and whose formation is quite peculiar. as they extended themselves into darkness. their loss may be attributed to disuse." On my view we must suppose that American animals. after numberless generations. the tuco-tuco. have been accommodated to surrounding circumstances. so that. in accordance with the old view of the blind animals having been separately created for the American and European caverns. and as eyes are certainly not necessary to animals having subterranean habits. It is well known that several animals. though the telescope with its glasses has been lost. In some of the crabs the foot-stalk for the eye remains. and therefore not in the profoundest depths. The eyes of moles and of some burrowing rodents are rudimentary in size. with respect to the insects alone. two of which were captured by Professor Silliman at above half a mile distance from the mouth of the cave. apply not to the same. As frequent inflammation of the eyes must be injurious to any animal. and natural selection will often have effected other changes.at all and had stuck to the wreck. and I was assured by a Spaniard. natural selection would aid the effects of disuse. acquired a dim perception of objects. as appeared on dissection. otherwise than as a very plain expression of that analogy which subsists generally between the fauna of Europe and of North America. and which. Next follow those that are constructed for twilight.

they become acclimatised: thus the pines and rhododendrons.. and from the number of plants and animals brought from different countries which are here perfectly healthy. analogous observations have been made by Mr.C. that the species. and then have become extinct. I am only surprised that more wrecks of ancient life have not been preserved. may formerly have ranged over both continents. Mr. as I hear from Professor Dana. and some of the European cave-insects are very closely allied to those of the surrounding country. and this leads me to say a few words on acclimatisation. yet those which inhabit the several caves of Europe and America are distinct. by adaptation to particular climates. Hooker from the same species growing at different heights on the Himalayas. etc. Thwaites informs me that he has observed similar facts in Ceylon. we might expect still to see in the cave-animals of America. we have evidence with some few plants. and as is the case with the blind Proteus. but it is possible that the progenitors of these several species. Watson on European species of plants brought . ACCLIMATISATION. in the time of sleep. if it be true that all the species of the same genus are descended from a single parent-form. to a certain extent. It would be difficult to give any rational explanation of the affinities of the blind cave-animals to the other inhabitants of the two continents on the ordinary view of their independent creation. As a blind species of Bathyscia is found in abundance on shady rocks far from caves. excepting in their present secluded abodes. as Mr. Murray observes. as in the period of flowering. for it is natural that an insect already deprived of vision should readily become adapted to dark caverns. raised from seed collected by Dr. as a compensation for blindness. many succulent plants cannot endure a damp climate. were found to possess in this country different constitutional powers of resisting cold. Notwithstanding such modifications. the loss of vision in the cave species of this one genus has probably had no relation to its dark habitation. We have reason to believe that species in a state of nature are closely limited in their ranges by the competition of other organic beings quite as much as. owing to the less severe competition to which the scanty inhabitants of these dark abodes will have been exposed. It is notorious that each species is adapted to the climate of its own home: species from an arctic or even from a temperate region cannot endure a tropical climate. or conversely.increase in the length of the antennae or palpi. as Agassiz has remarked in regard to the blind fish. That several of the inhabitants of the caves of the Old and New Worlds should be closely related. we might expect from the well-known relationship of most of their other productions. have not as yet been found anywhere except in caves. and in those of Europe to the inhabitants of the European continent. We may infer this from our frequent inability to predict whether or not an imported plant will endure our climate. Habit is hereditary with plants. while they were furnished with eyes. So again. with reference to the reptiles of Europe. But the degree of adaptation of species to the climates under which they live is often overrated. Another blind genus (Anophthalmus) offers this remarkable peculiarity. that is. Far from feeling surprise that some of the cave-animals should be very anomalous. And this is the case with some of the American cave-animals. of their becoming. naturally habituated to different temperatures. As it is extremely common for distinct species belonging to the same genus to inhabit hot and cold countries. in the amount of rain requisite for seeds to germinate. affinities to the other inhabitants of that continent. acclimatisation must be readily effected during a long course of descent. H. or more than. But whether or not this adaptation is in most cases very close. the Amblyopsis.

and not because they were subsequently found capable of far-extended transportation. and as most of these varieties are of recent origin. into action. new varieties have not been produced. That habit or custom has some influence. Hence adaptation to any special climate may be looked at as a quality readily grafted on an innate wide flexibility of constitution. which is never propagated in England by seed. the common and extraordinary capacity in our domestic animals of not only withstanding the most different climates. I think. be due to habit. even in the ancient Encyclopaedias of China. on account of the probable origin of some of our domestic animals from several wild stocks: the blood. to be very cautious in transporting animals from one district to another. their range from warmer to colder latitudes. In regard to animals. within historical times. and now have a far wider range than any other rodent. of the kidney-bean has been often cited for a similar purpose. for it is now as tender as ever it was! The case. during a score of generations. certain varieties are said to withstand certain climates better than others. however. common to most animals. as proving that acclimatisation cannot be effected. but we do not positively know that these animals were strictly adapted to their native climate. they cannot owe their constitutional differences to habit. and the fact of the extinct elephant and rhinoceros having formerly endured a glacial climate. of a tropical and arctic wolf may perhaps be mingled in our domestic breeds. On this view. On the other hand. natural selection would inevitably tend to preserve those individuals which were born with constitutions best adapted to any country which they inhabited. whereas the living species are now all tropical or sub-tropical in their habits. As we may infer that our domestic animals were originally chosen by uncivilised man because they were useful. but they have been transported by man to many parts of the world. the result must. and how much to the natural selection of varieties having different innate constitutions. both from analogy and from the incessant advice given in agricultural works. nor do we know that they have subsequently become specially acclimatised to their new homes. is an obscure question. several authentic instances could be adduced of species having largely extended. and on many an island in the torrid zones. though in all ordinary cases we assume such to be the case. I must believe. for instance. and of which. ought not to be looked at as anomalies. The case of the Jerusalem artichoke. but until some one will sow. in which certain varieties are habitually recommended for the northern and others for the southern states. We must not. and conversely. for they live under the cold climate of Faroe in the north and of the Falklands in the south. so as to be better fitted for them than they were at first. consequently.from the Azores to England. and because they bred readily under confinement. has even been advanced. his . push the foregoing argument too far. the capacity of enduring the most different climates by man himself and by his domestic animals. How much of the acclimatisation of species to any peculiar climate is due to mere habit. and how much to both means combined. under peculiar circumstances. may be used as an argument that a large proportion of other animals now in a state of nature could easily be brought to bear widely different climates. and with much greater weight. but of being perfectly fertile (a far severer test) under them. but as examples of a very common flexibility of constitution. The rat and mouse cannot be considered as domestic animals. also. In treatises on many kinds of cultivated plants. And as it is not likely that man should have succeeded in selecting so many breeds and sub-breeds with constitutions specially fitted for their own districts. this is strikingly shown in works on fruit-trees published in the United States. and I could give other cases. brought.

it is believed by some authors that with birds the diversity in the shape of the pelvis causes the remarkable diversity in the shape of the kidneys. Geoffroy St. Hard parts seem to affect the form of adjoining soft parts. played a considerable part in the modification of the constitution and structure. according to Schlegel. in some cases. that variations of structure arising in the young or larvae naturally tend to affect the structure of the mature animal. In snakes. at an early embryonic period. other parts become modified. most imperfectly understood. with the same precautions. I do not doubt. are identical in structure. or between the tortoise-shell colour and the female sex. Nor let it be supposed that differences in the constitution of seedling kidney-beans never appear. This is a very important subject. during its growth and development. On the whole. with care to prevent accidental crosses. Hilaire has forcibly remarked that certain malconformations frequently. One of the most obvious real cases is. it might probably have been rendered permanent by natural selection. though here no doubt homology . and that others rarely. and then collect seed from the few survivors. Homologous parts. as has been remarked by some authors. and which are necessarily exposed to similar conditions. coexist without our being able to assign any reason. I mean by this expression that the whole organisation is so tied together. and even in the jaws and limbs. or. have. and then again get seed from these seedlings. tend to cohere. the relation between the hair and the teeth in the naked Turkish dog. for the lower jaw is believed by some anatomists to be homologous with the limbs. or between the presence of more or less down on the young pigeon when first hatched. and if this had been of any great use to the breed. for an account has been published how much more hardy some seedlings are than others. The several parts which are homologous. in the front and hind legs. We shall presently see that simple inheritance often gives the false appearance of correlation. What can be more singular than the relation in cats between complete whiteness and blue eyes with deafness. but that the effects have often been largely combined with. and of this fact I have myself observed striking instances. may be mastered more or less completely by natural selection: thus a family of stags once existed with an antler only on one side. The nature of the bond is frequently quite obscure. the natural selection of innate variations. varying together. and sometimes overmastered by. and which. CORRELATED VARIATION. M. again. or in pigeons. and no doubt wholly different classes of facts may be here easily confounded together. with the future colour of its plumage. we may conclude that habit. These tendencies. or use and disuse. as in the union of the petals into a tube.kidney-beans so early that a very large proportion are destroyed by frost. seem eminently liable to vary in a like manner: we see this in the right and left sides of the body varying in the same manner. Is. the experiment cannot be said to have been even tried. the shape of the body and the manner of swallowing determine the position and form of several of the most important viscera. between their feathered feet and skin betwixt the outer toes. Others believe that the shape of the pelvis in the human mother influences by pressure the shape of the head of the child. this is often seen in monstrous plants: and nothing is more common than the union of homologous parts in normal structures. that when slight variations in any one part occur and are accumulated through natural selection.

by drawing nourishment from the reproductive organs causes their abortion. for in some Compositae the seeds of the outer and inner florets differ. that it has little value. and this difference is often accompanied with the partial or complete abortion of the reproductive organs. the daisy.). Mivart has remarked. after thousands of generations. For . Cetacea (whales) and Edentata (armadilloes. that in many pelargoniums the two upper petals in the central flower of the truss often lose their patches of darker colour. of the slightest service to the species. and which in truth are simply due to inheritance. as Mr. at least. for instance. scaly ant-eaters. or to their mutual pressure. may be wholly due to the laws of variation and correlation. and the shape of the seeds in the ray-florets of some Compositae countenances this idea. We know. It might have been thought that the development of the ray-petals. can be in any way beneficial. I may add. having been transmitted to a whole group of descendants with diverse habits. but this can hardly be the sole case. which are not always correlated with any difference in the corolla. When the colour is absent from only one of the two upper petals. as an instance of this fact. the central flower thus becoming peloric or regular. without being. than that of the difference between the outer and inner flowers in some Compositous and Umbelliferous plants. the nectary is not quite aborted but is much shortened. etc. would naturally be thought to be in some necessary manner correlated. it seems impossible that their differences in shape. that is to become abnormally symmetrical. yet in the Umbelliferae these differences are of such apparent importance--the seeds being sometimes orthospermous in the exterior flowers and coelospermous in the central flowers--that the elder De Candolle founded his main divisions in the order on such characters. and.. and as a striking case of correlation. Hence modifications of structure. without any difference in the corolla. Hooker informs me.comes into play? With respect to this latter case of correlation. for an ancient progenitor may have acquired through natural selection some one modification in structure. These differences have sometimes been attributed to the pressure of the involucra on the florets. and therefore of natural selection. independently of utility. We may often falsely attribute to correlated variation structures which are common to whole groups of species. that with irregular flowers those nearest to the axis are most subject to peloria. But in some of these plants the seeds also differ in shape and sculpture. and if so. and when this occurs. but there are so many exceptions to this rule. is highly probable. the adherent nectary is quite aborted. viz. But with respect to the seeds. the species with the densest heads which most frequently differ in their inner and outer flowers. Everyone is familiar with the difference between the ray and central florets of. but with the Umbelliferae it is by no means. Possibly these several differences may be connected with the different flow of nutriment towards the central and external flowers. as far as we can judge. I know of no case better adapted to show the importance of the laws of correlation and variation. Some other correlations are apparently due to the manner in which natural selection can alone act. some other and independent modification. I think it can hardly be accidental that the two orders of mammals which are most abnormal in their dermal covering. whose agency is highly advantageous. Sprengel's idea that the ray-florets serve to attract insects. are likewise on the whole the most abnormal in their teeth. and these two modifications. or necessary for the fertilisation of these plants. as Dr. viewed by systematists as of high value. With respect to the development of the corolla. natural selection may have come into play.

superfluous." I think this holds true to a certain extent with our domestic productions: if nourishment flows to one part or organ in excess. for in the struggle for life to which every animal is exposed.instance. the whole anterior part of the head is reduced to the merest rudiment attached to the bases of the prehensile antennae. "in order to spend on one side. and. but in the parasitic and protected Proteolepas. their law of compensation or balancement of growth. thus it is difficult to get a cow to give much milk and to fatten readily. to another part. may be merged under a more general principle. a large tuft of feathers on the head is generally accompanied by a diminished comb. the actual withdrawal of nutriment from one part owing to the excess of growth in another and adjoining part. And conversely. at least in excess. for I see hardly any way of distinguishing between the effects. when rendered superfluous. more especially botanists. The elder Geoffroy and Goethe propounded. becomes less useful. would be a decided advantage to each successive individual of the species. and a large beard by diminished wattles. by less nutriment being wasted. for in this case alone could the seeds. the fruit itself gains largely in size and quality. that some of the cases of compensation which have been advanced. here give any instances. however. that natural selection is continually trying to economise in every part of the organisation. each would have a better chance of supporting itself. for it will profit the individual not to have its nutriment wasted in building up a useless structure. at about the same time. and furnished with great nerves and muscles. and of which many other instances could be given: namely. as I believe. Alph. it loses more or less completely its own shell or carapace. namely. I will not. COMPENSATION AND ECONOMY OF GROWTH. gain an advantage over others less well fitted for wide dispersal. before useful. as Goethe expressed it. through changed habits. Thus. and in a truly extraordinary manner with the Proteolepas: for the carapace in all other cirripedes consists of the three highly important anterior segments of the head enormously developed. that when a cirripede is parasitic within another cirripede and is thus protected. on the one hand. as soon as it becomes. The same varieties of the cabbage do not yield abundant and nutritious foliage and a copious supply of oil-bearing seeds. without by any means causing some other part to be largely developed in a corresponding degree. De Candolle has remarked that winged seeds are never found in fruits which do not open. I should explain this rule by the impossibility of seeds gradually becoming winged through natural selection. Now the saving of a large and complex structure. on the other hand. and likewise some other facts. I suspect. which were a little better adapted to be wafted by the wind. With species in a state of nature it can hardly be maintained that the law is of universal application. that natural selection may perfectly well succeed in largely developing an organ without requiring as a necessary compensation the reduction of . In our poultry. natural selection will tend in the long run to reduce any part of the organisation. If under changed conditions of life a structure. it rarely flows. of a part being largely developed through natural selection and another and adjoining part being reduced by the same process or by disuse. its diminution will be favoured. but many good observers. or. also. When the seeds in our fruits become atrophied. This is the case with the male Ibla. nature is forced to economise on the other side. believe in its truth. unless the capsules were open. I can thus only understand a fact with which I was much struck when examining cirripedes.

RUDIMENTARY. as is generally admitted. seems to have come to a nearly similar conclusion. have further remarked that multiple parts are extremely liable to vary in structure. But . The rule being so plainly applicable in the case of secondary sexual characters. but more rarely to females. however unusually developed. both with varieties and species. I presume that lowness here means that the several parts of the organisation have been but little specialised for particular functions. It is hopeless to attempt to convince any one of the truth of the above proposition without giving the long array of facts which I have collected. but I hope that I have made due allowances for them. is a sign of low organisation. can act solely through and for the advantage of each being. and I will here only add that their variability seems to result from their uselessness. Waterhouse. may be due to the great variability of these characters. it would apply only if some one species had wings developed in a remarkable manner in comparison with the other species of the same genus. Several years ago I was much struck by a remark to the above effect made by Mr. It seems to be a rule. A PART DEVELOPED IN ANY SPECIES IN AN EXTRAORDINARY DEGREE OR MANNER. MULTIPLE. is constant." to use Professor Owen's expression. AND LOWLY-ORGANISED STRUCTURES ARE VARIABLE. As "vegetative repetition. Rudimentary parts. whereas the number of the same part or organ. Geoffroy St. but are not directly connected with the act of reproduction. when displayed in any unusual manner. that is. and consequently from natural selection having had no power to check deviations in their structure. but the rule would not apply here. it should never be forgotten. I can only state my conviction that it is a rule of high generality. that beings which stand low in the scale of nature are more variable than those which are higher. The same author as well as some botanists. The rule applies very strongly in the case of secondary sexual characters. as they seldom offer remarkable secondary sexual characters. Professor Owen. we can perhaps see why it should remain variable. whether or not displayed in any unusual manner--of which fact I think there can be little doubt. the foregoing statements accord with the common opinion of naturalists. In the same way that a knife which has to cut all sorts of things may be of almost any shape. unless it be unusually developed in one species or in a few species in comparison with the same part in many closely allied species. I am aware of several causes of error. Natural selection. relates to characters which are attached to one sex. and which cannot possibly be here introduced. IN COMPARISON WITH THE SAME PART IN ALLIED SPECIES. The term. also. are apt to be highly variable. The rule applies to males and females. Thus. whilst a tool for some particular purpose must be of some particular shape. that when any part or organ is repeated many times in the same individual (as the vertebrae in snakes. It should be understood that the rule by no means applies to any part. TENDS TO BE HIGHLY VARIABLE. secondary sexual characters. We shall have to recur to this subject. used by Hunter. as remarked by Is. why natural selection should not have preserved or rejected each little deviation of form so carefully as when the part has to serve for some one special purpose. because the whole group of bats possesses wings. and as long as the same part has to perform diversified work. and the stamens in polyandrous flowers) the number is variable. when it occurs in lesser numbers. the wing of the bat is a most abnormal structure in the class of mammals. Hilaire.some adjoining part.

give a list of all the more remarkable cases. in every sense of the word. and no selection be applied. the homologous valves in the different species being sometimes wholly unlike in shape. in the carriage and tail of fantails. many departing widely from the standard. I have particularly attended to them. had not the great variability in plants made it particularly difficult to compare their relative degrees of variability.that our rule is not confined to secondary sexual characters is clearly shown in the case of hermaphrodite cirripedes. are also eminently liable to variation. Pyrgoma. But what here more particularly concerns us is. If. Waterhouse's remark. etc. in a future work. than do the species belonging to other distinct genera. I think we can obtain some light. Look at the individuals of the same breed of the pigeon. on the one hand. which at the present time are undergoing rapid change by continued selection. the power of steady selection to keep the breed true. and have been modified through natural selection. much variability in the parts undergoing modification may always be expected. as in that of the short-faced tumbler. the tendency to reversion to a less perfect state. as well as an innate tendency to new variations. . and I am fully convinced that the rule almost always holds good. these being the points now mainly attended to by English fanciers. I particularly attended to Mr. There may truly be said to be a constant struggle going on between.. that those points in our domestic animals. vary extremely little. and see what a prodigious amount of difference there is in the beak of tumblers. with all its parts as we now see them. and the amount of variation in the individuals of the same species is so great that it is no exaggeration to state that the varieties of the same species differ more from each other in the characters derived from these important organs. and in those which have been but little specialised for any particular purpose. and perhaps in polymorphic groups. the comb in the Dorking fowl) or the whole breed will cease to have a uniform character: and the breed may be said to be degenerating. I can see no explanation. and we do not expect to fail so completely as to breed a bird as coarse as a common tumbler pigeon from a good short-faced strain. any part or the whole animal be neglected. I cannot make out that it applies to plants. But as long as selection is rapidly going on. very important structures. As with birds the individuals of the same species. and this would have seriously shaken my belief in its truth. But on the view that groups of species are descended from some other species. Even in the same sub-breed. it is notoriously difficult to breed nearly perfect birds. In the long run selection gains the day. When we see any part or organ developed in a remarkable degree or manner in a species. First let me make some preliminary remarks. on the other hand. these valves present a marvellous amount of diversification. inhabiting the same country. in our domestic animals. as it illustrates the rule in its largest application. in the beak and wattle of carriers. and they differ extremely little even in distinct genera. Why should this be so? On the view that each species has been independently created. we see a nearly parallel case. The opercular valves of sessile cirripedes (rock barnacles) are. I shall. that part (for instance. the fair presumption is that it is of high importance to that species: nevertheless it is in this case eminently liable to variation. and thus the organisation is left in a fluctuating condition. for in such cases natural selection either has not or cannot come into full play. I will here give only one. whilst investigating this order. and the rule certainly seems to hold good in this class. but in the several species of one genus. In rudimentary organs. and.

Something of the same kind applies to monstrosities: at least Is. An extraordinary amount of modification implies an unusually large and long-continued amount of variability. or conversely. For in this case the variability will seldom as yet have been fixed by the continued selection of the individuals varying in the required manner and degree. which is generally very constant throughout a large group of species. we may conclude that this part has undergone an extraordinary amount of modification since the period when the several species branched off from the common progenitor of the genus. I am convinced. which is generally of generic value. true. which has continually been accumulated by natural selection for the benefit of the species. for an immense period in nearly the same state. that when an author remarks with surprise that some important organ or part. because they are taken from parts of less physiological importance than those commonly used for classing genera. and the tendency to reversion and variability on the other hand. I see no reason to doubt. have to return to this point in the chapter on Classification. It would be almost superfluous to adduce evidence in support of the statement. it must have existed. and that the most abnormally developed organs may be made constant. that specific characters are more variable than generic. when an organ. To explain by a simple example what is meant: if in a large genus of plants some species had blue flowers and some had red. Hilaire apparently . Geoffroy St. however abnormal it may be. and thus it has come not to be more variable than any other structure. when it sinks in value and becomes only of specific value. namely. it is often VARIABLE in the individuals of the same species. It is only in those cases in which the modification has been comparatively recent and extraordinarily great that we ought to find the GENERATIVE VARIABILITY. And this. as it may be called. that ordinary specific characters are more variable than generic. and no one would be surprised at one of the blue species varying into red. often becomes variable. however. but if all the species had blue flowers. yet only indirectly. DIFFERS considerably in closely-allied species. I have chosen this example because the explanation which most naturalists would advance is not here applicable. It is notorious that specific characters are more variable than generic. But as the variability of the extraordinarily developed part or organ has been so great and long-continued within a period not excessively remote.Now let us turn to nature. That the struggle between natural selection on the one hand. This period will seldom be remote in any extreme degree. has been transmitted in approximately the same condition to many modified descendants. I have repeatedly noticed in works on natural history. and its variation would be a more unusual circumstance. I believe this explanation is partly. still expect to find more variability in such parts than in other parts of the organisation which have remained for a much longer period nearly constant. but with respect to important characters. Hence. still present in a high degree. we might. compared with the other species of the same genus. SPECIFIC CHARACTERS MORE VARIABLE THAN GENERIC CHARACTERS. according to our theory. the colour would become a generic character. the colour would be only a specific character. and by the continued rejection of those tending to revert to a former and less modified condition. though its physiological importance may remain the same. as in the case of the wing of the bat. will in the course of time cease. When a part has been developed in an extraordinary manner in any one species. The principle discussed under the last heading may be applied to our present subject. as species rarely endure for more than one geological period. is the case. as a general rule. And this fact shows that a character. I shall.

and subsequently have not varied or come to differ in any degree. but in the Engidae. than in other parts of their organisation. why should that part of the structure. which is less rigid in its action than ordinary selection. On the ordinary view of each species having been independently created. with the amount of difference between the females. we might expect often to find them still continuing to vary in those parts of their structure which have varied within a moderately recent period. that secondary sexual characters are highly variable. On the other hand. The cause of the original variability of these characters is not manifest. as Westwood has remarked. the more subject it is to anomalies in the individuals. sexual selection will have had a wide scope for action.entertains no doubt. The same number of joints in the tarsi is a character common to very large groups of beetles. and these characters may be attributed to inheritance from a common progenitor. and as these specific characters have varied and come to differ since the period when the species branched off from a common progenitor. the relation can hardly be accidental. that the secondary differences between the two sexes of the same species are generally displayed in the very same parts of the organisation in which the species of the same genus differ from each other. but only gives fewer offspring to the less favoured males. compare. for they are accumulated by sexual selection. But on the view that species are only strongly marked and fixed varieties. Or to state the case in another manner: the points in which all the species of a genus resemble each other. be more variable than those parts which are closely alike in the several species? I do not see that any explanation can be given. Whatever the cause may be of the variability of secondary sexual characters. but we can see why they should not have been rendered as constant and uniform as others. the amount of difference between the males of gallinaceous birds. the number varies greatly and the number likewise differs in . and in which they differ from allied genera. SECONDARY SEXUAL CHARACTERS VARIABLE. fitted to more or less widely different habits. It is a remarkable fact. without my entering on details. and which have thus come to differ. in which secondary sexual characters are strongly displayed. it is probable that they should still often be in some degree variable--at least more variable than those parts of the organisation which have for a very long period remained constant. and as the differences in these cases are of a very unusual nature. for instance. in exactly the same manner: and as these so-called generic characters have been inherited from before the period when the several species first branched off from their common progenitor. which differs from the same part in other independently created species of the same genus. as it does not entail death. or only in a slight degree. that the more an organ normally differs in the different species of the same group. Of this fact I will give in illustration the first two instances which happen to stand on my list. the points in which species differ from other species of the same genus are called specific characters. as they are highly variable. It will also be admitted that species of the same group differ from each other more widely in their secondary sexual characters. for it can rarely have happened that natural selection will have modified several distinct species. I think it will be admitted by naturalists. and may thus have succeeded in giving to the species of the same group a greater amount of difference in these than in other respects. are called generic characters. it is not probable that they should vary at the present day.

I conclude that the greater variability of specific characters. The most distinct breeds of the pigeon. and thus having been adapted for secondary sexual. be taken advantage of by natural and sexual selection. whatever part of the structure of the common progenitor. in countries widely apart. All being mainly due to the species of the same group being the descendants of a common progenitor. that several minute crustaceans offer excellent illustrations of this law. that the great variability of secondary sexual characters and their great difference in closely allied species. to natural selection having more or less completely. OR REVERTS TO SOME OF THE CHARACTERS OF AN EARLY PROGENITOR. In the vegetable kingdom we have a case of analogous variation. became variable. are all principles closely connected together. when acted on by similar unknown influences. DISTINCT SPECIES PRESENT ANALOGOUS VARIATIONS. These propositions will be most readily understood by looking to our domestic races. Consequently. overmastered the tendency to reversion and to further variability." This relation has a clear meaning on my view: I look at all the species of the same genus as having as certainly descended from the same progenitor. and with feathers on the feet. then. or those which distinguish species from species. because common to large groups. present sub-varieties with reversed feathers on the head. Lubbock has recently remarked. as have the two sexes of any one species. and likewise in the two sexes of the same species. Finally. these then are analogous variations in two or more distinct races. and to variations in the same parts having been accumulated by natural and sexual selection. or those which are possessed by all the species. the sexual characters are afforded mainly by the anterior antennae and by the fifth pair of legs: the specific differences also are principally given by these organs. that the frequent extreme variability of any part which is developed in a species in an extraordinary manner in comparison with the same part in its congeners. and the slight degree of variability in a part. it is highly probable. for instance. characters not possessed by the aboriginal rock-pigeon. from whom they have inherited much in common. to sexual selection being less rigid than ordinary selection. Again in the fossorial hymenoptera. and likewise to fit the two sexes of the same species to each other. variations of this part would. but in certain genera the neuration differs in the different species. the neuration of the wings is a character of the highest importance. "In Pontella. plants which several botanists rank as varieties produced by cultivation from a common parent: if this . in the enlarged stems. if it be common to a whole group of species. I presume that no one will doubt that all such analogous variations are due to the several races of the pigeon having inherited from a common parent the same constitution and tendency to variation. in order to fit the several places in the economy of nature. to parts which have recently and largely varied being more likely still to go on varying than parts which have long been inherited and have not varied. or as commonly called roots.the two sexes of the same species. than of generic characters. or of its early descendants. Sir J. and for ordinary purposes. or to fit the males to struggle with other males for the possession of the females. SO THAT A VARIETY OF ONE SPECIES OFTEN ASSUMES A CHARACTER PROPER TO AN ALLIED SPECIES. the fantail. however extraordinarily it may be developed. that secondary sexual and ordinary specific differences are generally displayed in the same parts of the organisation. The frequent presence of fourteen or even sixteen tail-feathers in the pouter may be considered as a variation representing the normal structure of another race. according to the lapse of time. of the Swedish turnip and ruta-baga.

be not so. whether strong or weak. beyond the influence of the mere act of crossing on the laws of inheritance. of slaty-blue birds with two black bars on the wings. the most probable hypothesis is. the case will then be one of analogous variation in two so-called distinct species. With pigeons. A mere tendency to produce a rudiment is indeed sometimes thus inherited. Walsh. namely. As all the species of the same genus are supposed to be descended from a common progenitor. to reproduce the lost character might. is developed. it might be expected that they would occasionally vary in an analogous manner. In a breed which has not been crossed. so that the varieties of two or more species would resemble each other. to use a common expression. not that one individual suddenly takes after an ancestor removed by some hundred generations. as was formerly remarked. under unknown favourable conditions. is only 1 in 2048. because. the offspring occasionally show for many generations a tendency to revert in character to the foreign breed--some say. we have another case. which very rarely produces a blue bird. Similar cases occurring with insects under natural conditions have lately been discussed with much ability by Mr. be transmitted for almost any number of generations. the occasional appearance in all the breeds. I think. or that a variety of one species would resemble in certain characters another and distinct species. and in this case there is nothing in the external conditions of life to cause the reappearance of the slaty-blue. only a well-marked and permanent variety. I presume that no one will doubt that this is a case of reversion. for all that we can see to the contrary. from one ancestor. But characters exclusively due to analogous variation . we should have to attribute this similarity in the enlarged stems of these three plants. reappears after a great number of generations. No doubt it is a very surprising fact that characters should reappear after having been lost for many. and to these a third may be added. namely. these coloured marks are eminently liable to appear in the crossed offspring of two distinct and differently coloured breeds. but in which BOTH parents have lost some character which their progenitor possessed. We may. and yet. With the barb-pigeon. however. and at last. this other species being. as we see. Many similar cases of analogous variation have been observed by Naudin in the great gourd family. The abstract improbability of such a tendency being transmitted through a vast number of generations. But when a breed has been crossed only once by some other breed. probably for hundreds of generations. the proportion of blood. the common turnip. confidently come to this conclusion. according to our view. it is generally believed that a tendency to reversion is retained by this remnant of foreign blood. with the outer feathers externally edged near their bases with white. According to the ordinary view of each species having been independently created. but to three separate yet closely related acts of creation. and not of a new yet analogous variation appearing in the several breeds. a bar at the end of the tail. who has grouped them under his law of equable variability. for a dozen or even a score of generations. As all these marks are characteristic of the parent rock-pigeon. but that in each successive generation the character in question has been lying latent. for instance. white loins. and a consequent tendency to vary in a like manner. as we have seen. is not greater than that of quite useless or rudimentary organs being similarly transmitted. and by various authors in our cereals. with the several marks. After twelve generations. it is probable that there is a latent tendency in each generation to produce blue plumage. not to the vera causa of community of descent. When a character which has been lost in a breed. the tendency.

mouse-duns. other species of the same genus. and in one instance in a chestnut. and from inquiries which I have made. has been described without either spinal or shoulder stripe. but NOT an albino. however. More especially we might have inferred this from the blue colour and the several marks so often appearing when differently coloured breeds are crossed. we do not know the common ancestor of any natural group. we could not have told. It is a case almost certainly of reversion. and of ALL colours. but Dr. but from occurring in several species of the same genus. transverse bars on the legs are not rare in duns. and what are new but analogous variations. But the best evidence of analogous variations is afforded by parts or organs which are generally constant in character. and seem to me very remarkable. a faint shoulder-stripe may sometimes be seen . for the preservation of all functionally important characters will have been determined through natural selection. The ass sometimes has very distinct transverse bars on its legs. sometimes to find the varying offspring of a species assuming characters which are already present in other members of the same group. The quagga. I have collected a long list of such cases. which themselves can only doubtfully be ranked as species. Mr. we cannot distinguish between reversionary and analogous characters. although under nature it must generally be left doubtful. on our theory. give one curious and complex case. as it were. and this shows. also. but which occasionally vary so as to resemble. A white ass. not indeed as affecting any important character. we did not know that the parent rock-pigeon was not feather-footed or turn-crowned. though so plainly barred like a zebra over the body. I have collected cases in England of the spinal stripe in horses of the most distinct breeds. If. as before. in accordance with the different habits of the species. the same part or organ in an allied species. for instance. whether such characters in our domestic breeds were reversions or only analogous variations. I lie under the great disadvantage of not being able to give them. in some degree. As. It might further be expected that the species of the same genus would occasionally exhibit reversions to long-lost characters. Gray has figured one specimen with very distinct zebra-like bars on the hocks. though it properly has none. The difficulty in distinguishing variable species is largely due to the varieties mocking. which are correlated with this tint. however. but we might have inferred that the blue colour was a case of reversion from the number of the markings. It has been asserted that these are plainest in the foal. I can only repeat that such cases certainly occur. and I have been informed by Colonel Poole that foals of this species are generally striped on the legs and faintly on the shoulder. that they have in varying assumed some of the characters of the others.would probably be of an unimportant nature. I will. Hence. what cases are reversions to formerly existing characters. in dark-coloured asses. The stripe on the shoulder is sometimes double. like those on the legs of a zebra. And this undoubtedly is the case. and these stripes are sometimes very obscure. I believe this to be true. and which would not probably have all appeared together from simple variation. The koulan of Pallas is said to have been seen with a double shoulder-stripe. yet we ought. but here. Blyth has seen a specimen of the hemionus with a distinct shoulder-stripe. unless all these closely allied forms be considered as independently created species. With respect to the horse. and is very variable in length and outline. A considerable catalogue. or actually quite lost. could be given of forms intermediate between two other forms. is without bars on the legs. partly under domestication and partly under nature.

. has given a figure of a similar mule. a hybrid has been figured by Dr. the dun. was striped. according to Mr. Martin. Rollin asserts that the common mule from the ass and horse is particularly apt to have bars on its legs. when a week old. who has written on this subject. W. that with the English race-horse the spinal stripe is much commoner in the foal than in the full-grown animal. This foal. as I hear from Colonel Poole. Norwegian cobs. the side of the face. Lastly. inhabiting the most distant parts of the world. and the shoulder-stripe. from information given me by Mr. etc. the legs are generally barred. should have all have been crossed with one supposed aboriginal stock. both with THREE parallel stripes on each shoulder. which I have seen. and a small dun Welsh pony has been carefully described to me. In four coloured drawings. the lanky Kattywar race. a horse without stripes is not considered as purely bred. I am aware that Colonel Hamilton Smith. and that the above-described appearances are all due to ancient crosses with the dun stock. the legs were much more plainly barred than the rest of the body. believes that the several breeds of the horse are descended from several aboriginal species. Gray (and he informs me that he knows of a second case) from the ass and the hemionus. though the ass only occasionally has stripes on his legs and the hemionus has none and has not even a shoulder-stripe. and sometimes quite disappear in old horses. I once saw a mule with its legs so much striped that any one might have thought that it was a hybrid zebra. from one between brown and black to a close approach to cream colour. All the stripes soon disappeared completely. in certain parts of the United States. moreover. in his excellent treatise on the horse. of hybrids between the ass and zebra. I have myself recently bred a foal from a bay mare (offspring of a Turkoman horse and a Flemish mare) by a bay English race-horse. who examined this breed for the Indian Government. The spine is always striped. which is sometimes double and sometimes treble. one of which. Gosse. Without here entering on further details I may state that I have collected cases of leg and shoulder stripes in horses of very different breeds in various countries from Britain to Eastern China. But this view may be safely rejected. for it is highly improbable that the heavy Belgian cart-horse. W. and in one of them there was a double shoulder-stripe. is common. is sometimes striped. and had three short . Welsh ponies. and Mr. was marked on its hinder quarters and on its forehead with numerous very narrow. In Lord Morton's famous hybrid. I have myself seen a dun Devonshire pony. I have also reason to suspect. from a chestnut mare and male quagga. In all parts of the world these stripes occur far oftenest in duns and mouse-duns. were much more plainly barred across the legs than is even the pure quagga. that. My son made a careful examination and sketch for me of a dun Belgian cart-horse with a double stripe on each shoulder and with leg-stripes. Now let us turn to the effects of crossing the several species of the horse genus. zebra-like bars. by the term dun a large range of colour is included.C. and this is another most remarkable case.W. dark. and from Norway in the north to the Malay Archipelago in the south. The stripes are often plainest in the foal. and I have seen a trace in a bay horse. Colonel Poole has seen both gray and bay Kattywar horses striped when first foaled. and this hybrid. nevertheless had all four legs barred. and its legs were feebly striped. In the northwest part of India the Kattywar breed of horses is so generally striped.in duns. Edwards. about nine out of ten mules have striped legs. the hybrid and even the pure offspring subsequently produced from the same mare by a black Arabian sire.

we see a strong tendency for the blue tint and bars and marks to reappear in the mongrels. When the oldest and truest breeds of various colours are crossed. With respect to this last fact. I have stated that the most probable hypothesis to account for the reappearance of very ancient characters. I presume. Call the breeds of pigeons. but perhaps otherwise very differently constructed. that I was led solely from the occurrence of the face-stripes on this hybrid from the ass and hemionus to ask Colonel Poole whether such face-stripes ever occurred in the eminently striped Kattywar breed of horses. I venture confidently to look back thousands on thousands of generations. Not in one case out of a hundred can we pretend to assign any reason why this or that part has varied. And we have just seen that in several species of the horse genus the stripes are either plainer or appear more commonly in the young than in the old. and how exactly parallel is the case with that of the species of the horse genus! For myself. What now are we to say to these several facts? We see several distinct species of the horse genus becoming. some of which have bred true for centuries. and the greater differences . to reject a real for an unreal. quagga. will. both under nature and under domestication. these bars and other marks invariably reappear. so as often to become striped like the other species of the genus. But whenever we have the means of instituting a comparison. To admit this view is. but other species of the genus. when crossed with species inhabiting distant quarters of the world. not their own parents. We see this tendency to become striped most strongly displayed in hybrids from between several of the most distinct species. and zebra. to produce hybrids resembling in their stripes.shoulder-stripes. Now observe the case of the several breeds of pigeons: they are descended from a pigeon (including two or three sub-species or geographical races) of a bluish colour. and I see an animal striped like a zebra. The appearance of the stripes is not accompanied by any change of form. species. the same laws appear to have acted in producing the lesser differences between varieties of the same species. like those on the dun Devonshire and Welsh ponies. by simple variation. that fossil shells had never lived. I would almost as soon believe with the old and ignorant cosmogonists. assert that each species has been created with a tendency to vary. as we have seen. and even had some zebra-like stripes on the sides of its face. striped on the legs like a zebra. It makes the works of God a mere mockery and deception. is--that there is a TENDENCY in the young of each successive generation to produce the long-lost character. the common parent of our domestic horse (whether or not it be descended from one or more wild stocks) of the ass. but without any other change of form or character. In the horse we see this tendency strong whenever a dun tint appears--a tint which approaches to that of the general colouring of the other species of the genus. answered in the affirmative. or at least for an unknown cause. but had been created in stone so as to mock the shells now living on the sea-shore. SUMMARY. as it seems to me. or striped on the shoulders like an ass. and that this tendency. or by any other new character. Our ignorance of the laws of variation is profound. sometimes prevails. from unknown causes. I was so convinced that not even a stripe of colour appears from what is commonly called chance. and was. in this particular manner. the hemionus. and when any breed assumes by simple variation a bluish tint. He who believes that each equine species was independently created. with certain bars and other marks. and that each has been created with a strong tendency.

Whatever the cause may be of each slight difference between the . but sometimes they cause direct and definite effects. and such characters differ much in the species of the same group. from being useless. and specific differences to the several species of the same genus. or these same species may occasionally revert to some of the characters of their ancient progenitors. the characters which have come to differ since the several species of the same genus branched off from a common parent--are more variable than generic characters. appear in many cases to have been potent in their effects. undoubtedly occur. Changes of structure at an early age may affect parts subsequently developed. must have gone through an extraordinary amount of modification since the genus arose. most varieties. Changed conditions generally induce mere fluctuating variability. and every part of the structure which can be saved without detriment will be saved. but we have also seen in the second chapter that the same principle applies to the whole individual. and exposed to similar influences. Although new and important modifications may not arise from reversion and analogous variation. or where the manufactory of new specific forms has been actively at work--in that district and among these species. and these may become strongly marked in the course of time. Specific characters--that is. Secondary sexual characters are highly variable. It follows probably from this same cause. though we have not sufficient evidence on this head. such modifications will add to the beautiful and harmonious diversity of nature. When one part is largely developed. and many cases of correlated variation. perhaps it tends to draw nourishment from the adjoining parts. are not regulated by natural selection. Variability in the same parts of the organisation has generally been taken advantage of in giving secondary sexual differences to the two sexes of the same species. requiring a long lapse of time--in this case. and thus we can understand why it should often still be variable in a much higher degree than other parts. natural selection has succeeded in giving a fixed character to the organ. because they have recently varied and thus come to differ. and hence are variable. Habit in producing constitutional peculiarities. in comparison with the same part or organ in the allied species. where there has been much former variation and differentiation. perhaps arising from such parts not having been closely specialised for any particular function. and have not differed within this same period. Homologous parts tend to vary in the same manner. But when a species with an extraordinarily developed organ has become the parent of many modified descendants--which on our view must be a very slow process. that organic beings low in the scale are more variable than those standing higher in the scale. and disuse in weakening and diminishing organs. and homologous parts tend to cohere. so that their modifications have not been closely checked by natural selection. or those which have long been inherited. Species inheriting nearly the same constitution from a common parent. on an average. and natural selection will in such cases not as yet have had time to overcome the tendency to further variability and to reversion to a less modified state. Rudimentary organs. in however extraordinary a manner it may have been developed. naturally tend to present analogous variations. In these remarks we have referred to special parts or organs being still variable. for in a district where many species of a genus are found--that is. Any part or organ developed to an extraordinary size or in an extraordinary manner. for variation is a long-continued and slow process. and which have their whole organisation more specialised. the nature of which we are unable to understand. Modifications in hard parts and in external parts sometimes affect softer and internal parts. we now find. and use in strengthening. Multiple parts are variable in number and in structure.between species of the same genus.

each new form will tend in a fully-stocked country to take the place of. I think. on the other hand. the structure and habits of a bat. whereas. but. being sterile and producing sterile offspring. for instance. is it possible that an animal having. and. Some of them are so serious that to this day I can hardly reflect on them without being in some degree staggered. such as the tail of a giraffe. Instinct and Hybridism in the two succeeding chapters. and those that are real are not. its own less improved parent-form and other less-favoured forms with which it comes into competition. the greater number are only apparent. which serves as a fly-flapper. Long before the reader has arrived at this part of my work. why. do we not everywhere see innumerable transitional forms? Why is not all nature in confusion. can instincts be acquired and modified through natural selection? What shall we say to the instinct which leads the bee to make cells. their fertility is unimpaired? The two first heads will be here discussed. fatal to the theory.offspring and their parents--and a cause for each must exist--we have reason to believe that it is the steady accumulation of beneficial differences which has given rise to all the more important modifications of structure in relation to the habits of each species. an organ of trifling importance. to the best of my judgment. if species have descended from other species by fine gradations. well defined? Secondly. could have been formed by the modification of some other animal with widely different habits and structure? Can we believe that natural selection could produce. some miscellaneous objections in the following chapter. Difficulties of the theory of descent with modification--Absence or rarity of transitional varieties--Transitions in habits of life--Diversified habits in the same species--Species with habits widely different from those of their allies--Organs of extreme perfection--Modes of transition--Cases of difficulty--Natura non facit saltum--Organs of small importance--Organs not in all cases absolutely perfect--The law of Unity of Type and of the Conditions of Existence embraced by the theory of Natural Selection. CHAPTER VI. As natural selection acts solely by the preservation of profitable modifications. and finally to exterminate. and which has practically anticipated the discoveries of profound mathematicians? Fourthly. on the one hand. an organ so wonderful as the eye? Thirdly. instead of the species being. Thus extinction and natural selection go hand in hand. when varieties are crossed. when crossed. a crowd of difficulties will have occurred to him. These difficulties and objections may be classed under the following heads: First. ON THE ABSENCE OR RARITY OF TRANSITIONAL VARIETIES. as we see them. DIFFICULTIES OF THE THEORY. . how can we account for species.

why do we not now find closely-linking intermediate varieties? This difficulty for a long time quite confounded me. But if we compare these species where they intermingle. for I believe that many perfectly defined species have been formed on strictly continuous areas. why do we not find them embedded in countless numbers in the crust of the earth? It will be more convenient to discuss this question in the chapter on the imperfection of the geological record. and has supplanted and exterminated its original parent-form and all the transitional varieties between its past and present states. But in the intermediate region. till the one replaces the other. and only at long intervals of time. De Candolle has observed. But it may be urged that when several closely allied species inhabit the same territory. that it has been continuous during a long period.Hence. Let us take a simple case: in travelling from north to south over a continent. and may be embedded there in a fossil condition. These representative species often meet and interlock. In looking at species as they are now distributed over a wide area. Hence the neutral territory between two representative species is generally narrow in comparison with the territory proper to each. marine areas now continuous must often have existed within recent times in a far less continuous and uniform condition than at present. though I do not doubt that the formerly broken condition of areas now continuous. more especially with freely-crossing and wandering animals. and as the one becomes rarer and rarer. as Alph. But I think it can be in large part explained. but the natural collections have been imperfectly made. The crust of the earth is a vast museum. We see the same fact in ascending mountains. the other becomes more and more frequent. But. then becoming somewhat abruptly rarer and rarer on the confines. having intermediate conditions of life. we generally meet at successive intervals with closely allied or representative species. and I will here only state that I believe the answer mainly lies in the record being incomparably less perfect than is generally supposed. . though they must have existed there. evidently filling nearly the same place in the natural economy of the land. The same fact has been noticed by E. we generally find them tolerably numerous over a large territory. because an area is now continuous. and sometimes it is quite remarkable how abruptly. as by this theory innumerable transitional forms must have existed. a common alpine species disappears. if we look at each species as descended from some unknown form. Hence we ought not to expect at the present time to meet with numerous transitional varieties in each region. both the parent and all the transitional varieties will generally have been exterminated by the very process of the formation and perfection of the new form. By my theory these allied species are descended from a common parent. But I will pass over this way of escaping from the difficulty. and in such islands distinct species might have been separately formed without the possibility of intermediate varieties existing in the intermediate zones. and during the process of modification. we surely ought to find at the present time many transitional forms. Geology would lead us to believe that most continents have been broken up into islands even during the later tertiary periods. each has become adapted to the conditions of life of its own region. has played an important part in the formation of new species. By changes in the form of the land and of climate. In the first place we should be extremely cautious in inferring. and finally disappearing. they are generally as absolutely distinct from each other in every detail of structure as are specimens taken from the metropolis inhabited by each.

Wollaston. sooner than the forms which they originally linked together. this rule holds good with varieties in a state of nature. in the race for life. . not blending one into another by insensible gradations. Now. as far as I can make out. during fluctuations in the number of its enemies or of its prey. each species on the confines of its range. Watson. and a third variety to a narrow intermediate zone. depending as it does on the range of others. But when we bear in mind that almost every species.Forbes in sounding the depths of the sea with the dredge. run a greater chance of being exterminated than one existing in large numbers. then. For any form existing in lesser numbers would. and in this particular case the intermediate form would be eminently liable to the inroads of closely allied forms existing on both sides of it. or in the nature of the seasons. that during the process of further modification. that each organic being is either directly or indirectly related in the most important manner to other organic beings--we see that the range of the inhabitants of any country by no means exclusively depends on insensibly changing physical conditions. as climate and height or depth graduate away insensibly. we shall have to adapt two varieties to two large areas. for these will be more slowly modified and improved. Asa Gray. be extremely liable to utter extermination. where it exists in lessened numbers. Dr. As allied or representative species. with a comparatively narrow neutral territory between them. even in its metropolis. and as these species are already defined objects. when varieties intermediate between two other forms occur. but in large part on the presence of other species. The intermediate variety. are generally distributed in such a manner that each has a wide range. and practically. and if we take a varying species inhabiting a very large area. or by which it is destroyed. the range of any one species. they are much rarer numerically than the forms which they connect. the same rule will probably apply to both. or with which it comes into competition. To those who look at climate and the physical conditions of life as the all-important elements of distribution. when inhabiting a continuous area. will. than will the rarer forms which exist in lesser numbers. will tend to beat and supplant the less common forms. if we may trust these facts and inferences. that generally. then we can understand why intermediate varieties should not endure for very long periods: why. But it is a far more important consideration. And it would appear from information given me by Mr. on which it lives. they should be exterminated and disappear. will have a great advantage over the intermediate variety. as varieties do not essentially differ from species. from inhabiting larger areas. and Mr. Moreover. these facts ought to cause surprise. that nearly all either prey on or serve as prey for others. Hence. which exists in smaller numbers in a narrow and intermediate zone. I have met with striking instances of the rule in the case of varieties intermediate between well-marked varieties in the genus Balanus. and thus its geographical range will come to be still more sharply defined. were it not for other competing species. within any given period. will tend to be sharply defined. of presenting further favourable variations for natural selection to seize on. by which two varieties are supposed to be converted and perfected into two distinct species. would increase immensely in numbers. as a general rule. in which they become rather suddenly rarer and rarer. the more common forms. will exist in lesser numbers from inhabiting a narrow and lesser area. in short. and conclude that varieties linking two other varieties together generally have existed in lesser numbers than the forms which they connect. as already remarked. the two which exist in larger numbers. consequently. For forms existing in larger numbers will have a better chance.

because new varieties are very slowly formed. in the aggregate. for these. and natural selection can do nothing until favourable individual differences or variations occur. present more varieties. I believe that species come to be tolerably well-defined objects. numberless intermediate varieties. but these links during the process of natural selection will have been supplanted and exterminated. From this cause alone the intermediate varieties will be liable to accidental extermination. I may illustrate what I mean by supposing three varieties of sheep to be kept. if my theory be true. at first have been formed in the intermediate zones. For these intermediate varieties will. the chances in this case will be strongly in favour of the great holders on the mountains or on the plains improving their breeds more quickly than the small holders on the intermediate narrow. from existing in greater numbers will. or on the occasional immigration of new inhabitants. and likewise of acknowledged varieties). and that the inhabitants are all trying with equal steadiness and skill to improve their stocks by selection. hilly tract. they will almost certainly be beaten and supplanted by the forms which they connect. intermediate varieties between the several representative species and their common parent. in a still more important degree. more especially among the classes which unite for each birth and wander much.It is the same principle which. will come into close contact with each other. hilly tract. but at all time. with the new forms thus produced and the old ones acting and reacting on each other. looking not to any one time. And such new places will depend on slow changes of climate. linking closely together all . without the interposition of the supplanted. intermediate varieties will. and consequently the improved mountain or plain breed will soon take the place of the less improved hill breed. which originally existed in greater numbers. it is probable. so that they will no longer be found in a living state. and a third to the wide plains at the base. on some of the old inhabitants becoming slowly modified. we ought to see only a few species presenting slight modifications of structure in some degree permanent. Thirdly. as I believe. accounts for the common species in each country. a second to a comparatively narrow. and thus be further improved through natural selection and gain further advantages. presenting on an average a greater number of well-marked varieties than do the rarer species. and during the process of further modification through natural selection. when two or more varieties have been formed in different portions of a strictly continuous area. Secondly. in which many forms. as shown in the second chapter. To sum up. Lastly. probably. areas now continuous must often have existed within the recent period as isolated portions. for variation is a slow process. from reasons already assigned (namely from what we know of the actual distribution of closely allied or representative species. but they will generally have had a short duration. and thus the two breeds. and. must formerly have existed within each isolated portion of the land. one adapted to an extensive mountainous region. may have separately been rendered sufficiently distinct to rank as representative species. in any one region and at any one time. intermediate hill variety. and this assuredly we do see. exist in the intermediate zones in lesser numbers than the varieties which they tend to connect. and until a place in the natural polity of the country can be better filled by some modification of some one or more of its inhabitants. and do not at any one period present an inextricable chaos of varying and intermediate links: first. In this case. So that.

Now look at the Galeopithecus or so-called flying lemur. as Sir J. for instance. or. Look at the family of squirrels. each being propagated. at least. or to collect food more quickly. but during the long winter it leaves the frozen waters. We cannot doubt that each structure is of use to each kind of squirrel in its own country. it is clear that each must be well adapted to its place in nature. to lessen the danger from occasional falls. I lie under a heavy disadvantage. during summer this animal dives for and preys on fish. But it does not follow from this fact that the structure of each squirrel is the best that it is possible to conceive under all possible conditions. ON THE ORIGIN AND TRANSITION OF ORGANIC BEINGS WITH PECULIAR HABITS AND STRUCTURE. like other polecats on mice and land animals. and flying squirrels have their limbs and even the base of the tail united by a broad expanse of skin. in the same species. which serves as a parachute and allows them to glide through the air to an astonishing distance from tree to tree. and from others. here we have the finest gradation from animals with their tails only slightly flattened. each modification being useful. with the posterior part of their bodies rather wide and with the skin on their flanks rather full. and all analogy would lead us to believe that some. could a land carnivorous animal have been converted into one with aquatic habits. which are preserved. Here. Let the climate and vegetation change. must assuredly have existed. of the squirrels would decrease in numbers or become exterminated. short legs. and it had been asked how an insectivorous quadruped could possibly have been converted into a flying bat. It has been asked by the opponents of such views as I hold. which has webbed feet. by the accumulated effects of this process of natural selection. in an extremely imperfect and intermittent record. for how could the animal in its transitional state have subsisted? It would be easy to show that there now exist carnivorous animals presenting close intermediate grades from strictly terrestrial to aquatic habits. as there is reason to believe. which was . Look at the Mustela vison of North America. either constant or occasional. unless they also become modified and improved in structure in a corresponding manner. the question would have been far more difficult to answer. for. And it seems to me that nothing less than a long list of such cases is sufficient to lessen the difficulty in any particular case like that of the bat. If a different case had been taken. and form of tail. to exterminate the parent forms and the intermediate links. by enabling it to escape birds or beasts of prey. but the very process of natural selection constantly tends. how. more especially under changing conditions of life. and of diversified habits. Yet I think such difficulties have little weight. Consequently evidence of their former existence could be found only among fossil remains. to the so-called flying squirrels. out of the many striking cases which I have collected. until. Therefore. a perfect so-called flying squirrel was produced. and preys. and which resembles an otter in its fur. and as each exists by a struggle for life. I can give only one or two instances of transitional habits and structures in allied species. Richardson has remarked. as we shall attempt to show in a future chapter. as has been so often remarked. or old ones become modified.the species of the same group. as on other occasions. let other competing rodents or new beasts of prey immigrate. in the continued preservation of individuals with fuller and fuller flank-membranes. I can see no difficulty.

we perhaps see traces of an apparatus originally fitted for gliding through the air rather than for flight. Furthermore. as far as the organs of flight are concerned. and seeing that we have flying birds and mammals. we should bear in mind that animals displaying early transitional grades of the structure will seldom have survived to the present day. slightly rising and turning by the aid of their fluttering fins. so as to have given them a decided advantage over other animals in the battle for life. which perhaps may all be the result of disuse. might have been modified into perfectly winged animals. for each has to live by a struggle: but it is not necessarily the best possible under all possible conditions. Nor can I see any insuperable difficulty in further believing it possible that the membrane-connected fingers and fore-arm of the Galeopithecus might have been greatly lengthened by natural selection. It must not be inferred from these remarks that any of the grades of wing-structure here alluded to. it is conceivable that flying-fish. which now glide far through the air. who would have ventured to surmise that birds might have existed which used their wings solely as flappers. yet there is no difficulty in supposing that such links formerly existed. like the logger headed duck (Micropterus of Eyton). Although no graduated links of structure. as sails. it does not seem probable that fishes capable of true flight would have been developed under many subordinate forms. and this. now connect the Galeopithecus with the other Insectivora. which were gradually rendered more perfect through natural selection. indicate the steps by which birds actually acquired their perfect power of flight. An extremely wide flank-membrane stretches from the corners of the jaw to the tail. so far as we know. fitted for gliding through the air. and had used their incipient organs of flight exclusively. like the penguin. we may conclude that transitional states between structures fitted for very different habits of life will rarely have been developed at an early period in great numbers and under many subordinate forms.formerly ranked among bats. but they serve to show what diversified means of transition are at least possible. under the conditions of life to which it is exposed. as the wings of a bird for flight. each grade of structure having been useful to its possessor. on the land and in the water. but is now believed to belong to the Insectivora. who would have ever imagined that in an early transitional state they had been inhabitants of the open ocean. and functionally for no purpose. like the ostrich. Seeing that a few members of such water-breathing classes as the Crustacea and Mollusca are adapted to live on the land. If about a dozen genera of birds were to become extinct. would have converted the animal into a bat. In certain bats in which the wing-membrane extends from the top of the shoulder to the tail and includes the hind-legs. If this had been effected. until their organs of flight had come to a high stage of perfection. and includes the limbs with the elongated fingers. to escape being devoured by other fish? When we see any structure highly perfected for any particular habit. Thus. and that each was developed in the same manner as with the less perfectly gliding squirrels. like the apteryx? Yet the structure of each of these birds is good for it. flying insects of the most diversified types. This flank-membrane is furnished with an extensor muscle. and formerly had flying reptiles. to return to our imaginary illustration of the flying-fish. for they will have been supplanted by their successors. Hence the chance of discovering species with transitional grades of structure in a fossil condition will always be . for taking prey of many kinds in many ways. as fins in the water and as front legs on the land.

I will now give two or three instances. but with . but not so stiff as in the typical wood-peckers. and bores holes in the trunk for its nest. in its manner of swimming and of flying when made to take flight. or exclusively to one of its several habits. or exclusively on artificial substances. As we sometimes see individuals following habits different from those proper to their species and to the other species of the same genus. both probably often occurring almost simultaneously. than in the case of species with fully developed structures. and undulatory flight. it is essentially a petrel. having anomalous habits. and at other times standing stationary on the margin of water. Hence this Colaptes. insects in the water. Of diversified habits innumerable instances could be given: I have often watched a tyrant flycatcher (Saurophagus sulphuratus) in South America. Petrels are the most aerial and oceanic of birds. there is a woodpecker (Colaptes campestris) which has two toes before and two behind. On the plains of La Plata.less. where hardly a tree grows. in its astonishing power of diving. the Puffinuria berardi. both of diversified and of changed habits. and I have many times seen and heard it hammering the seeds of the yew on a branch. And such instances occur in nature. frequents trees. a long-pointed tongue. is not so straight or so strong as in the typical woodpeckers but it is strong enough to bore into wood. kills small birds by blows on the head. pointed tail-feathers. yet. is a woodpecker. as Mr. whether habits generally change first and structure afterwards. In North America the black bear was seen by Hearne swimming for hours with widely open mouth. and a straight. but from those of the accurate Azara. that a Mexican Colaptes has been described by De Saussure as boring holes into hard wood in order to lay up a store of acorns. thus catching. like a shrike. sufficiently stiff to support the bird in a vertical position on a post. in the individuals of the same species. almost like a whale. and with their structure either slightly or considerably modified from that of their type. not only from my own observations. from their having existed in lesser numbers. I may mention as another illustration of the varied habits of this genus. strong beak. as I can assert. difficult to decide and immaterial for us. like a kestrel. Of cases of changed habits it will suffice merely to allude to that of the many British insects which now feed on exotic plants. It is. it sometimes. nevertheless. or whether slight modifications of structure lead to changed habits. we might expect that such individuals would occasionally give rise to new species. in certain large districts it does not climb trees. hovering over one spot and then proceeding to another. however. in the quiet sounds of Tierra del Fuego. In either case it would be easy for natural selection to adapt the structure of the animal to its changed habits. in its general habits. In our own country the larger titmouse (Parus major) may be seen climbing branches. this same woodpecker. The beak. and then dashing into it like a kingfisher at a fish. but. and others with elongated wings which chase insects on the wing. and thus breaking them like a nuthatch. the harsh tone of the voice. and it makes its nest in holes in banks! In certain other districts. Even in such trifling characters as the colouring. in all the essential parts of its structure. however. Can a more striking instance of adaptation be given than that of a woodpecker for climbing trees and seizing insects in the chinks of the bark? Yet in North America there are woodpeckers which feed largely on fruit. would be mistaken by any one for an auk or a grebe. its close blood-relationship to our common woodpecker is plainly declared. Hudson states. almost like a creeper. however.

All the members of the great order of Hymenopterous insects are terrestrial. and thus gains an advantage over some other inhabitant of the same country. and for the correction of spherical and chromatic aberration. and remains as long as four hours beneath the surface. which Sir John Lubbock has discovered to be aquatic in its habits. will acknowledge that every organic being is constantly endeavouring to increase in numbers. for admitting different amounts of light. though not in structure. and petrels with the habits of auks. and that if any one being varies ever so little. What seems plainer than that the long toes. either in habits or structure. habits have changed without a corresponding change of structure. are formed for walking over swamps and floating plants. but this seems to me only restating the fact in dignified language. the deeply scooped membrane between the toes shows that structure has begun to change. that there should be long-toed corncrakes. The water-hen and landrail are members of this order.many parts of its organisation profoundly modified in relation to its new habits of life. The webbed feet of the upland goose may be said to have become almost rudimentary in function. has seen the frigate-bird. of the Grallatores. which is allied to the thrush family. and the second is nearly as terrestrial as the quail or partridge. Reason tells . but the old saying of Vox populi. living in meadows instead of in swamps. whereas the woodpecker of La Plata has had its structure only slightly modified. cannot be trusted in science. which has all its four toes webbed. it often enters the water and dives about by the use not of its legs but of its wings. and many others could be given. On the other hand. could have been formed by natural selection. When it was first said that the sun stood still and the world turned round. seems. that there should be diving thrushes and diving Hymenoptera. I freely confess. In the frigate-bird. absurd in the highest degree. living on the dry land and rarely alighting on the water. What can be plainer than that the webbed feet of ducks and geese are formed for swimming? Yet there are upland geese with webbed feet which rarely go near the water. as every philosopher knows. excepting the genus Proctotrupes. that in these cases it has pleased the Creator to cause a being of one type to take the place of one belonging to another type. grebes and coots are eminently aquatic. subsists by diving. In such cases. it will seize on the place of that inhabitant. In the case of the water-ouzel. by examining its dead body. alight on the surface of the ocean. and no one except Audubon. To suppose that the eye with all its inimitable contrivances for adjusting the focus to different distances. that there should be woodpeckers where hardly a tree grows. yet it exhibits no modification in structure in accordance with its abnormal habits. the common sense of mankind declared the doctrine false. although their toes are only bordered by membrane. the acutest observer.--using its wings under water and grasping stones with its feet. He who believes in the struggle for existence and in the principle of natural selection. He who believes that each being has been created as we now see it. must occasionally have felt surprise when he has met with an animal having habits and structure not in agreement. yet this bird. however different that may be from its own place. would never have suspected its sub-aquatic habits. ORGANS OF EXTREME PERFECTION AND COMPLICATION. He who believes in separate and innumerable acts of creation may say. Hence it will cause him no surprise that there should be geese and frigate-birds with webbed feet. not furnished with membrane. vox Dei. yet the first is nearly as aquatic as the coot.

but only to concentrate the luminous rays and render their perception more easy. and for the chance of some gradations having been transmitted in an unaltered or little altered condition. for we have only to place the naked extremity of the optic nerve. In searching for the gradations through which an organ in any species has been perfected. are capable of perceiving light. projecting with a convex surface. In certain star-fishes. the eye ever varies and the variations be inherited. but destitute of lens or other optical contrivance. with respect to the wide. according to M. which in some of the lower animals lies deeply buried in the body. with transparent gelatinous matter. but this is scarcely ever possible. as described by the author just quoted. that if numerous gradations from a simple and imperfect eye to one complex and perfect can be shown to exist. though insuperable by our imagination. the latter sometimes forming a sort of pupil. We may. and if such variations should be useful to any animal under changing conditions of life. and resting merely on sarcodic tissue. and in some near the surface. and serve only to distinguish light from darkness. But these organs in the Articulata are so much diversified that Muller formerly made three main classes with seven subdivisions. In the great class of the Articulata. as some of the lowest organisms in which nerves cannot be detected. hardly concerns us more than how life itself originated. and graduated range of structure in the eyes of the lower animals. however. and when we bear in mind how small the number of all living forms must be in comparison with those which have become extinct. besides a fourth main class of aggregated simple eyes. The simplest organ which can be called an eye consists of an optic nerve. How a nerve comes to be sensitive to light. surrounded by pigment-cells and covered by translucent skin. But the state of the same organ in distinct classes may incidentally throw light on the steps by which it has been perfected. descend even a step lower and find aggregates of pigment-cells. here given much too briefly. then the difficulty of believing that a perfect and complex eye could be formed by natural selection. and an image will be formed on it. picture-forming eye. the difficulty ceases to be very great in believing that natural selection may have converted the simple apparatus of an optic . small depressions in the layer of pigment which surrounds the nerve are filled. Jourdain. In this concentration of the rays we gain the first and by far the most important step towards the formation of a true. and we are forced to look to other species and genera of the same group. that is to the collateral descendants from the same parent-form. should not be considered as subversive of the theory. as is certainly the case. at the right distance from the concentrating apparatus. we may start from an optic nerve simply coated with pigment. as is likewise certainly the case. diversified. like the cornea in the higher animals. in order to see what gradations are possible. without any nerves. When we reflect on these facts.me. With insects it is now known that the numerous facets on the cornea of their great compound eyes form true lenses. we ought to look exclusively to its lineal progenitors. apparently serving as organs of vision. and that the cones include curiously modified nervous filaments. but I may remark that. Eyes of the above simple nature are not capable of distinct vision. He suggests that this serves not to form an image. it does not seem impossible that certain sensitive elements in their sarcode should become aggregated and developed into nerves. each grade being useful to its possessor. but without any lens or other refractive body. endowed with this special sensibility. if further.

and then suppose every part of this layer to be continually changing slowly in density. so as to separate into layers of different densities and thicknesses. it is assumed. He who will go thus far. placed at different distances from each other. then any increased regularity of curvature will be an improvement." Within the highest division of the animal kingdom. although in this case he does not know the transitional states. of a little sack of transparent skin. each to be preserved until a better is produced. under varied circumstances.nerve. into an optical instrument as perfect as is possessed by any member of the Articulata class. could not be done through natural selection. however. it is indispensable that the reason should conquer the imagination. at a just conclusion regarding the formation of the eye. that it consists. but I have felt the difficulty far to keenly to be surprised at others hesitating to extend the principle of natural selection to so startling a length. and then the old ones to be all . It is scarcely possible to avoid comparing the eye with a telescope. with all its marvellous yet not absolutely perfect characters. otherwise inexplicable. In fishes and reptiles. It has been objected that in order to modify the eye and still preserve it as a perfect instrument. and carefully preserving each which. the beautiful crystalline lens is formed in the embryo by an accumulation of epidermic cells. always intently watching each slight alteration in the transparent layers. in any way or degree. coated with pigment and invested by transparent membrane. tends to produce a distincter image. according to the high authority of Virchow. lying in a sack-like fold of the skin." It is a significant fact that even in man. as in the lancelet. and the vitreous body is formed from embryonic subcutaneous tissue. if he finds on finishing this volume that large bodies of facts. But may not this inference be presumptuous? Have we any right to assume that the Creator works by intellectual powers like those of man? If we must compare the eye to an optical instrument. and the rays do not converge to a point. "If a lens has too short or too long a focus. or an alteration of density. ought not to hesitate to go one step further. but as I have attempted to show in my work on the variation of domestic animals. many changes would have to be effected simultaneously. as Owen has remarked. which. if they were extremely slight and gradual. So the contraction of the iris and the muscular movements of the eye are neither of them essential to vision. and with the surfaces of each layer slowly changing in form. also. and we naturally infer that the eye has been formed by a somewhat analogous process. the Vertebrata. but destitute of any other apparatus. To arrive. "The range of gradation of dioptric structures is very great. furnished with a nerve and lined with pigment. represented by natural selection or the survival of the fittest. we can start from an eye so simple. if the curvature be irregular. but only improvements which might have been added and perfected at any stage of the construction of the instrument. Further we must suppose that there is a power. can be explained by the theory of modification through natural selection. it is not necessary to suppose that the modifications were all simultaneous. he ought to admit that a structure even as perfect as an eagle's eye might thus be formed. namely. and with a nerve sensitive to light beneath. we ought in imagination to take a thick layer of transparent tissue. Wallace has remarked. it may be amended either by an alteration of curvature. We know that this instrument has been perfected by the long-continued efforts of the highest human intellects. Different kinds of modification would. serve for the same general purpose: as Mr. We must suppose each new state of the instrument to be multiplied by the million. with spaces filled with fluid.

by spirally twining. slight modifications. and natural selection will pick out with unerring skill each improvement. Numerous cases could be given among the lower animals of the same organ performing at the same time wholly distinct functions. or the same organ under two very different forms. this latter organ being divided by highly vascular partitions and having a ductus pneumaticus for the supply of air. successive. combined in the same individual. a great change would be effected with comparative suddenness in the character of the species. Many plants are known which regularly produce at the same time differently constructed flowers. In such cases natural selection might specialise. It is. the animal may be turned inside out. In all such cases one of the two organs might readily be modified and perfected so as to perform all the work. we should have to look to very ancient ancestral forms. To give another instance from the vegetable kingdom: plants climb by three distinct means. Let this process go on for millions of years. or be wholly obliterated. for one function alone. and may we not believe that a living optical instrument might thus be formed as superior to one of glass. The illustration of the swim-bladder in fishes is a good one. at the same time that they breathe free air in their swim-bladders. and in order to discover the early transitional grades through which the organ has passed. however. Again. But I can find out no such case. and the exterior surface will then digest and the stomach respire. more especially if we look to much-isolated species. may simultaneously perform in the same individual the same function.destroyed. and during each year on millions of individuals of many kinds. there has been much extinction. generation will multiply them almost infinitely. which had previously performed two functions. for in this latter case the organ must have been originally formed at a remote period. according to the theory. and thus by insensible steps greatly change its nature. but some few species exhibit two of the means. and this is an extremely important means of transition: to give one instance--there are fish with gills or branchiae that breathe the air dissolved in the water. probable that the two sorts of flowers borne by the same plant were originally differentiated by finely graduated steps. since which all the many members of the class have been developed. these three means are usually found in distinct groups. and excretes. variation will cause the slight alteration. if we take an organ common to all the members of a class. No doubt many organs exist of which we do not know the transitional grades. If it could be demonstrated that any complex organ existed. being aided during the progress of modification by the other organ. my theory would absolutely break down. as the works of the Creator are to those of man? MODES Of TRANSITION. thus in the larva of the dragon-fly and in the fish Cobites the alimentary canal respires. the whole or part of an organ. and by the emission of aerial rootlets. and if such plants were to produce one kind alone. and then this other organ might be modified for some other and quite distinct purpose. Or again. digests. if any advantage were thus gained. long since become extinct. In the Hydra. In living bodies. around which. by clasping a support with their sensitive tendrils. which could not possibly have been formed by numerous. We should be extremely cautious in concluding that an organ could not have been formed by transitional gradations of some kind. because . which may still be followed in some few cases. or even all three. two distinct organs.

This has lately been insisted on by Professor Cope and others in the United States. It is now known that some animals are capable of reproduction at a very early age. The Balanidae or sessile cirripedes. These cirripedes have no branchiae. large. which originally served as ovigerous frena. it is therefore highly probable that in this great class organs which once served for respiration have been actually converted into organs for flight. indeed. also. or "ideally similar" in position and structure with the lungs of the higher vertebrate animals: hence there is no reason to doubt that the swim-bladder has actually been converted into lungs. or an organ used exclusively for respiration. Now I think no one will dispute that the ovigerous frena in the one family are strictly homologous with the branchiae of the other family. have been gradually converted by natural selection into branchiae. before they have acquired their perfect characters. been worked in as an accessory to the auditory organs of certain fishes. on the other hand. who would ever have imagined that the branchiae in this latter family had originally existed as organs for preventing the ova from being washed out of the sack? There is another possible mode of transition. together with the small frena. they graduate into each other. in the same relative position with the frena. We can thus. According to this view it may be inferred that all vertebrate animals with true lungs are descended by ordinary generation from an ancient and unknown prototype which was furnished with a floating apparatus or swim-bladder. it . But it is conceivable that the now utterly lost branchiae might have been gradually worked in by natural selection for some distinct purpose: for instance. likewise. Therefore it need not be doubted that the two little folds of skin. and if this power became thoroughly well developed in a species. but they have. which freely communicate with the circulatory lacunae of the sack and body. namely flotation. through the acceleration or retardation of the period of reproduction.it shows us clearly the highly important fact that an organ originally constructed for one purpose. serving for respiration. All physiologists admit that the swim-bladder is homologous. have no ovigerous frena. it is so important to bear in mind the probability of conversion from one function to another. If all pedunculated cirripedes had become extinct. as I infer from Professor Owen's interesting description of these parts. notwithstanding the beautiful contrivance by which the glottis is closed. to retain the eggs until they are hatched within the sack. the whole surface of the body and of the sack. within the well-enclosed shell. that I will give another instance. simply through an increase in their size and the obliteration of their adhesive glands. In the higher Vertebrata the branchiae have wholly disappeared--but in the embryo the slits on the sides of the neck and the loop-like course of the arteries still mark their former position. The swim-bladder has. and they have suffered far more extinction than have sessile cirripedes. but which. with some risk of falling into the lungs. much-folded membranes. and which have been considered by all naturalists to act as branchiae. which serve. In considering transitions of organs. namely. Landois has shown that the wings of insects are developed from the trachea. through the means of a sticky secretion. called by me the ovigerous frena. very slightly aided in the act of respiration. namely respiration. may be converted into one for a widely different purpose. Pedunculated cirripedes have two minute folds of skin. the eggs lying loose at the bottom of the sack. understand the strange fact that every particle of food and drink which we swallow has to pass over the orifice of the trachea.

it would be extremely bold to maintain that no serviceable transitions are possible by which these organs might have been gradually developed. Whether species have often or ever been modified through this comparatively sudden mode of transition. as observed by Matteucci. Professor Cope states that the teeth of certain lizards change much in shape with advancing years. and in this case. These organs appear at first to offer another and far more serious . With crustaceans not only many trivial.seems probable that the adult stage of development would sooner or later be lost. the form of the skull is often much altered with age. as Dr. yet in the ray. The electric organs of fishes offer another case of special difficulty. not known to be electrical. there would seem to be a charge in every respect like that which is met with in muscle and nerve during the rest. but as we know so little about the uses of these organs. R. yet undoubtedly serious cases of difficulty occur. McDonnell has shown. but if this has occurred. another organ near the head. there is. In all such cases--and many could be given--if the age for reproduction were retarded. after arriving at maturity. In the gymnotus and torpedo they no doubt serve as powerful means of defence. for instance. and between the mature and the old. as Dr. an analogous organ in the tail manifests but little electricity. at least in its adult state. but some important parts assume a new character. But this is not surprising. not a few animals. SPECIAL DIFFICULTIES OF THE THEORY OF NATURAL SELECTION. were primordially acquired by graduated steps. It should. "in the electrical apparatus of the torpedo during rest. the character of the species. It is generally admitted that there exists between these organs and ordinary muscle a close analogy. but which appears to be the real homologue of the electric battery in the torpedo. Again. Moreover. Radcliffe insists. also. which are often differently constructed from either the males or fertile females. for it is impossible to conceive by what steps these wondrous organs have been produced. One of the most serious is that of neuter insects. as recorded by Fritz Muller. transitional gradations. besides the organ just referred to. be especially observed that muscular contraction is accompanied by an electrical discharge. the character of the species would be greatly changed and degraded. and as we know nothing about the habits and structure of the progenitors of the existing electric fishes. Although we must be extremely cautious in concluding that any organ could not have been produced by successive. instead of being peculiar. for we do not even know of what use they are. in intimate structure. especially if the larva differed much from the mature form. With mammals. nor is it improbable that the previous and earlier stages of development would in some cases be hurried through and finally lost. small. and perhaps for securing prey. may be only another form of the discharge which attends upon the action of muscle and motor nerve. but this case will be treated of in the next chapter. even when the animal is greatly irritated. after maturity. so little that it can hardly be of any use for the above purposes. would be modified. and the discharge of the torpedo. I can form no opinion. in the ray. and in the manner in which they are acted on by various reagents. and the plumes of some birds become more finely developed. Every one knows how the horns of stags become more and more branched. of which Dr. Murie has given some striking instances with seals. and. go on changing in character during nearly their whole lives. it is probable that the differences between the young and the mature. in the distribution of the nerves." Beyond this we cannot at present go in the way of explanation. as they grow older.

Consequently there is no reason to suppose that they have been inherited from a common progenitor. the very curious contrivance of a mass of pollen-grains. disappears. apparently the same. a difficulty almost exactly parallel with that of the electric organs. Hence it is not a little difficult to decide how far even the same terms ought to be . is apparently the same in Orchis and Asclepias. and. and so in other points. especially if in members having very different habits of life. but only as analogous in function. When the same organ is found in several members of the same class. and must include some sort of lens for throwing an image at the back of a darkened chamber. of which several are widely remote in their affinities. by what graduated steps these organs have been developed in each separate group of fishes. that they differ in construction. In all cases of beings.difficulty. with an actual inversion of the elemental parts. for instance in plants. But when we look at the subject more closely. but this is far from the case. belonging to widely different families. according to Pacini. placed one behind the other like two lenses. genera almost as remote as is possible among flowering plants. An organ for vision must be formed of transparent tissue. we may generally attribute its presence to inheritance from a common ancestor. but here again the parts are not homologous. The relations of the muscles are as different as it is possible to conceive. and its absence in some of the members to loss through disuse or natural selection. we might have expected that all electric fishes would have been specially related to each other. for they occur in about a dozen kinds of fish. Beyond this superficial resemblance. and with a large nervous ganglion included within the membranes of the eye. it will be found that although the general appearance and function of the organs may be the same. Nor does geology at all lead to the belief that most fishes formerly possessed electric organs. The retina is wholly different. as in the arrangement of the plates. and in such widely sundered groups no part of this resemblance can be due to inheritance from a common progenitor. both having a very different structure and disposition to what occurs in the vertebrata. Other similar cases could be given. but I am unable to see the force of his argument. in the process or means by which the electricity is excited--and lastly. there is hardly any real similarity between the eyes of cuttle-fish and vertebrates. that these are situated in different parts of the body. if the electric organs had been inherited from some one ancient progenitor. The crystalline lens in the higher cuttle-fish consists of two parts. The luminous organs which occur in a few insects. yet fundamental differences between them can always be detected. Hence in the several fishes furnished with electric organs. Mivart has advanced this case as one of special difficulty. as may be seen by consulting Hensen's admirable memoir on these organs in the Cephalopoda. and which are situated in different parts of the body. It is impossible for me here to enter on details. Thus the difficulty of an organ. in being supplied with nerves proceeding from different sources. borne on a foot-stalk with an adhesive gland. leaving only the lesser yet still great difficulty: namely. these cannot be considered as homologous. arising in several remotely allied species. we find in the several fishes provided with electric organs. which are furnished with similar and peculiar organs. but I may specify a few of the points of difference. offer. and this is perhaps the most important of all the differences. So that. for had this been the case they would have closely resembled each other in all respects. the eyes of Cephalopods or cuttle-fish and of vertebrate animals appear wonderfully alike. under our present state of ignorance. far removed from each other in the scale of organisation. which their modified descendants have now lost. Mr. For instance.

on the supposition that species belonging to distinct families had slowly become adapted to live more and more out of water. as far as function is concerned. He shows that there are parasitic mites (Acaridae). and lastly in the whole structure of the water-breathing branchiae. and has arrived at the same result. Several families of crustaceans include a few species. of course. which were more especially examined by Muller. have been made to differ. and taking advantage of all favourable variations. would have differed to a certain extent. in the manner in which they are opened and closed. identical. in accordance with this view of their manner of formation. Now such differences are intelligible. Fritz Muller argues that this close similarity in so many points of structure must. But as the vast majority of the species in the above two families. from belonging to distinct families. Fritz Muller. while all the other important organs were closely similar. in order to arrive at the same functional result. in distinct organic beings. and the structures thus acquired would almost necessarily have differed. Hence it might have been expected that in the few species belonging to both families which live on the land. viz. their variability assuredly would not have been exactly the same. or rather. This line of argument seems to have had great weight in leading Fritz Muller to accept the views maintained by me in this volume. but if this be admitted in the one case it is clearly possible in the other. and to breathe the air. open to any one to deny that the eye in either case could have been developed through the natural selection of successive slight variations. and in accordance with the principle that the nature of each variation depends on two factors. are aquatic in their habits. so in the several foregoing cases it appears that natural selection. has produced similar organs. the species agree most closely in all important characters: namely in their sense organs. It is. as in the position of the orifices. Consequently natural selection would have had different materials or variations to work on. which are furnished with hair-claspers. and fundamental differences of structure in the visual organs of two groups might have been anticipated. in the position of the tufts of hair within their complex stomachs.. has argued in the same manner. belonging to distinct sub-families and families. and he found it to differ in each in several important points. Muller was thus led carefully to examine the apparatus in the air-breathing species. even to the microscopical hooks by which they are cleansed. These . the nature of the organism and that of the surrounding conditions. and might even have been expected. for why should this one apparatus. in order to test the conclusions arrived at in this volume. which owe none of their structure in common to inheritance from a common progenitor. as well as most other crustaceans. possessing an air-breathing apparatus and fitted to live out of the water. in accordance with the views advanced by me. and in some accessory details. has followed out with much care a nearly similar line of argument. On the hypothesis of separate acts of creation the whole case remains unintelligible.employed in describing the eyes of the Cephalopoda and Vertebrata. Another distinguished zoologist. circulating systems. working for the good of each being. For these species. the equally important air-breathing apparatus would have been the same. given for the same purpose. it is improbable in the highest degree that their common progenitor should have been adapted for breathing air. In two of these families. and which are nearly related to each other. the late Professor Claparede. As two men have sometimes independently hit on the same invention. be accounted for by inheritance from a common progenitor.

and still more so the four wings of a butterfly. Some authors maintain that organic beings have been formed in many ways for the sake of mere variety. though not in development. From such structures we may advance till we come to such a case of extraordinary adaptation as that lately described by Dr. the two wings of a fly. so as to be wafted by every breeze. and is itself hollowed out into a sort of chamber with two lateral entrances. and of appendages on the under side of the hind part of the body. though very different plan occurs in many plants in which a symmetrical flower secretes a few drops of nectar. we see the same end gained and the same function performed. could never have imagined what purpose all these parts serve. How differently constructed is the feathered wing of a bird and the membrane-covered wing of a bat. Cruger saw crowds of large humble-bees visiting the gigantic flowers of this orchid. but on what a number of patterns is the hinge constructed. and by being furnished with wings and plumes. sometimes forming trap-like contrivances. as well as conspicuously coloured. and sometimes capable of neatly adapted movements through irritability or elasticity. with the stamens and pistils modified in many ways. though hermaphrodites. of the maxillae or lips. The basal part of the labellum stands over the bucket. and is consequently visited by insects. closely similar. into which drops of almost pure water continually fall from two secreting horns which stand above it. being blown by the wind through mere chance on to the stigma. in doing this they frequently pushed each other into the bucket. in beings not at all or only remotely allied. The most ingenious man. and the two wings with the elytra of a beetle. as different in shape as they are elegant in structure. An almost equally simple. almost like toys in a shop. and in the several groups they are formed by the modification of the fore legs. if he had not witnessed what takes place. and these carry the pollen from the anthers to the stigma. so as to adhere to the fur of quadrupeds. I will give one other instance: for this subject of the same end being gained by the most diversified means well deserves attention. the pollen does not spontaneously fall on the stigma. . but such a view of nature is incredible. by being embedded in pulp or flesh. and with those in which. by their capsule being converted into a light balloon-like envelope. and when the bucket is half-full. The nectar may be stored in variously shaped receptacles. it is a common rule throughout nature that the same end should be gained. formed of the most diverse parts. from the long row of neatly interlocking teeth in a Nucula to the simple ligament of a Mussel! Seeds are disseminated by their minuteness. But Dr. by organs in appearance. within this chamber there are curious fleshy ridges. the water overflows by a spout on one side. but to gnaw off the ridges within the chamber above the bucket. and this is the simplest plan which can well be conceived. even sometimes in the case of closely related beings. all for the same purpose and effected in essentially the same manner. With several kinds this is effected by the pollen-grains. but entailing changes in every part of the flower. so as to attract and be devoured by birds. by having hooks and grapnels of many kinds and serrated awns. of the hind legs. by the most diversified means. which are light and incoherent. Bivalve shells are made to open and shut. In the foregoing cases. as they could not have been inherited from a common progenitor. some aid is necessary for their fertilisation. not in order to suck nectar. and rendered nutritious. On the other hand. Cruger in the Coryanthes.organs must have been independently developed. This orchid has part of its labellum or lower lip hollowed out into a great bucket. With plants having separated sexes. From this simple stage we may pass through an inexhaustible number of contrivances.

which already differ from each other in some slight degree. yet. Dr. and is equally curious. namely. We should also bear in mind that every highly developed organism has passed through many changes. and is pushed by its comrades into the bucket and then crawls out by the passage. for whatever purpose it may serve. Cruger saw a "continual procession" of bees thus crawling out of their involuntary bath. and retain the pollen. with a bee which he had killed before it had quite crawled out. or to the same flower a second time. The pollen-mass of the male plant (for the sexes are separate in this orchid) is thus carried to the flower of the female plant. which is viscid enough to break certain elastic threads. thus effecting fertilisation. and adheres by its viscid extremity to the back of the bee. where it is brought into contact with the stigma. but may be again and again further altered. considering how small the proportion of living and known forms is to the extinct and unknown. the variability will not be of the same exact nature. Finally. then." We meet with this admission in the . in order to gnaw the labellum. but were compelled to crawl out through the passage formed by the spout or overflow. Hence. so that a bee. Bees visit these flowers. transmits a sensation or vibration to a certain membrane which is instantly ruptured. in doing this they inevitably touch a long. in the foregoing and in innumerable other instances. as already remarked. The construction of the flower in another closely allied orchid. this sets free a spring by which the pollen-mass is shot forth. though serving the same end. first rubs its back against the viscid stigma and then against the viscid glands of the pollen-masses. How. the structure of each part of each species. Cruger sent me a flower in spirits of wine. sensitive projection. in the right direction. Now at last we see the full use of every part of the flower. like those of the Coryanthes. towards which no transitional grade is known to lead. tapering. and consequently the results obtained through natural selection for the same general purpose will not be the same. as indeed is shown by that old. that new organs appearing as if created for some special purpose rarely or never appear in any being. and the flower is fertilised. as I have called it. It is certainly true. I have been astonished how rarely an organ can be named. The passage is narrow. although in many cases it is most difficult even to conjecture by what transitions organs could have arrived at their present state. of the water-secreting horns of the bucket half-full of water. the Catasetum. with a pollen-mass still fastened to its back. and rub against the properly placed viscid pollen-masses and the viscid stigma. This antenna. thus provided. and are thus carried away. which prevents the bees from flying away. When the bee. or. and forces them to crawl out through the spout. and adheres to it. it may be asked. but somewhat exaggerated. Dr. like an arrow. the antenna. when touched. in forcing its way out. so that each modification will not readily be quite lost. through which the species has passed during its successive adaptations to changed habits and conditions of life. and that each modified structure tends to be inherited.and their wings being thus wetted they could not fly away. is the sum of many inherited changes. is widely different. canon in natural history of "Natura non facit saltum. The pollen-masses are thus glued to the back of the bee which first happens to crawl out through the passage of a lately expanded flower. the pollen-mass necessarily comes first into contact with the viscid stigma. that when two forms vary. can we understand the graduated scale of complexity and the multifarious means for gaining the same end. and is roofed over by the column. flies to another flower. The answer no doubt is.

for the hare. as Milne Edwards has well expressed it. but must advance by the short and sure. though the aid in this latter respect must be slight. "Nature is prodigal in variety. AS AFFECTED BY NATURAL SELECTION. In the first place. as in the case of the dog. or to escape from beasts of prey. might assuredly be acted on by natural selection. for so trifling an object as to drive away flies. or from determining the attacks of insects. with hardly any tail. ORGANS OF LITTLE APPARENT IMPORTANCE. which in their lungs or modified swim-bladders betray their aquatic origin. its general presence and use for many purposes in so many land animals. we can clearly understand why she should not. or as an aid in turning. Seeing how important an organ of locomotion the tail is in most aquatic animals. as a fly-flapper. would be able to range into new pastures and thus gain a great advantage.writings of almost every experienced naturalist. by the survival of the fittest. almost as great. so that they are more subject to disease. should there be so much variety and so little real novelty? Why should all the parts and organs of many independent beings. The tail of the giraffe looks like an artificially constructed fly-flapper. the colour of the skin and hair of quadrupeds. Organs now of trifling importance have probably in some cases been of high importance to an early progenitor. can double still more quickly. as in the case of the most perfect and complex organs. I have sometimes felt great difficulty in understanding the origin or formation of parts of little importance. it might subsequently come to be worked in for all sorts of purposes. though of a very different kind. such as the down on fruit and the colour of its flesh. though slow steps. or not so well enabled in a coming dearth to search for food. It is not that the larger quadrupeds are actually destroyed (except in some rare cases) by flies. and it seems at first incredible that this could have been adapted for its present purpose by successive slight modifications. In a former chapter I have given instances of very trifling characters. As natural selection acts by life and death. yet we should pause before being too positive even in this case. but niggard in innovation. each supposed to have been separately created for its own proper place in nature. on the theory of Creation." Why. for we know that the distribution and existence of cattle and other animals in South America absolutely depend on their power of resisting the attacks of insects: so that individuals which could by any means defend themselves from these small enemies. A well-developed tail having been formed in an aquatic animal. for natural selection acts only by taking advantage of slight successive variations. we are much too ignorant in regard to the whole economy of any one organic being to say what slight modifications would be of importance or not. be so commonly linked together by graduated steps? Why should not Nature take a sudden leap from structure to structure? On the theory of natural selection. may perhaps be thus accounted for. and by the destruction of the less well-fitted individuals. or. which. have been transmitted to existing species in nearly the same state. although now of very slight use. she can never take a great and sudden leap. an organ of prehension. and. each better and better fitted. but they are incessantly harassed and their strength reduced. but any actually injurious deviations in their structure would of course have been checked by natural selection. . after having been slowly perfected at a former period. from being correlated with constitutional differences.

and no doubt they facilitate. and in believing that they have been developed through natural selection. of the tendency to reversion to long-lost characters. as is the liability to be poisoned by certain plants. though of no use to the sex. and are exposed to a certain extent to natural selection. the colour is probably in chief part due to sexual selection. We are profoundly ignorant of the cause of each slight variation or individual difference. we may infer that this structure has arisen from the laws of growth. or may be indispensable for this act. and we did not know that there were many black and pied kinds. more especially in the less civilized countries. by which characters of use to one sex are often gained and then transmitted more or less perfectly to the other sex. but we should be very cautious in drawing any such inference. etc. and then . as there is reason to believe from the distribution of the thorn-bearing species in Africa and South America. A trailing palm in the Malay Archipelago climbs the loftiest trees by the aid of exquisitely constructed hooks clustered around the ends of the branches. and that with the hair the horns are correlated. and we are immediately made conscious of this by reflecting on the differences between the breeds of our domesticated animals in different countries. We must by no means overlook the effects of the definite action of changed conditions of life. The naked skin on the head of a vulture is generally considered as a direct adaptation for wallowing in putridity. of the complex laws of growth. and consequently that it was a character of importance. under new conditions of life and newly acquired habits. may subsequently have been taken advantage of by its modified descendants. and has been taken advantage of in the parturition of the higher animals. or it may possibly be due to the direct action of putrid matter. But structures thus indirectly gained. but as sutures occur in the skulls of young birds and reptiles.In the second place. although at first of no advantage to a species. such as of correlation. as it underwent further modification and became a climber. which seem to depend in a quite subordinate degree on the nature of the conditions. so that even colour would be thus subjected to the action of natural selection. so the spikes on the palm may at first have been developed for this object. is of the highest service to the plant. as it is. Animals kept by savages in different countries often have to struggle for their own subsistence. and had been acquired through natural selection. and subsequently have been improved and taken advantage of by the plant. we may easily err in attributing importance to characters. and which. and finally of sexual selection. The sutures in the skulls of young mammals have been advanced as a beautiful adaptation for aiding parturition. and individuals with slightly different constitutions would succeed best under different climates. Some observers are convinced that a damp climate affects the growth of the hair. I dare say that we should have thought that the green colour was a beautiful adaptation to conceal this tree-frequenting bird from its enemies. comprehension.. and this contrivance. where there has been but little methodical selection. and possibly even the form of the pelvis. when we see that the skin on the head of the clean-feeding male turkey is likewise naked. of the pressure of one part on another. no doubt. Mountain breeds always differ from lowland breeds. and so it may be. and a mountainous country would probably affect the hind limbs from exercising them more. With cattle susceptibility to the attacks of flies is correlated with colour. serve as a defence against browsing quadrupeds. of so-called spontaneous variations. but as we see nearly similar hooks on many trees which are not climbers. which have only to escape from a broken egg. If green woodpeckers alone had existed.

are of special use to these animals. We may safely attribute these structures to inheritance. to delight man or the Creator (but this latter point is beyond the scope of scientific discussion). UTILITARIAN DOCTRINE. But a still more important consideration is that the chief part of the organisation of every living creature is due to inheritance. and may never have been of any use to their progenitors. which nevertheless are generally admitted to have arisen through ordinary generation from one or a few parent-stocks. also. and the various causes of modifications. The effects of lessened exercise. is apparently one chief cause of the great modification which the breeds of swine have undergone. a view already discussed. are of special use to these birds. probably a great effect. as H. and this. It is scarcely possible to decide how much allowance ought to be made for such causes of change. HOW ACQUIRED. No doubt the definite action of changed conditions. if true. and we may further venture to believe that the several bones in the limbs of the monkey. von Nathusius has lately shown in his excellent Treatise. The laborious breathing necessary in high regions tends.by the law of homologous variation. together with abundant food. on the whole organisation is probably still more important. as we have good reason to believe. we ought not to lay too much stress on our ignorance of the precise cause of the slight analogous differences between true species. or was formerly. the front limbs and the head would probably be affected. I fully admit that many structures are now of no direct use to their possessors. They believe that many structures have been created for the sake of beauty. But webbed feet no doubt were as useful to the progenitor of the upland goose and of the frigate-bird. so-called spontaneous variations. probably through the reduction of more numerous bones in the fin of some ancient fish-like progenitor of the whole class. The shape. or of the frigate-bird. and the complex laws of growth. Such doctrines. of some direct or indirect use to its possessor. and again correlation would come into play. would be absolutely fatal to my theory. if we are unable to account for the characteristic differences of our several domestic breeds. and in the flipper of the seal. as they now are to the most aquatic of living birds. of the pelvis might affect by pressure the shape of certain parts of the young in the womb. but a foot with five toes fitted for walking or grasping. HOW FAR TRUE: BEAUTY. were originally developed. independently of any advantage thus gained. have all produced an effect. to increase the size of the chest. But we are far too ignorant to speculate on the relative importance of the several known and unknown causes of variation. So we may believe that the progenitor of the seal did not possess a flipper. . we may conclude that the structure of every living creature either now is. or for the sake of mere variety. in the fore leg of the horse. as the definite action of external conditions. but with these important exceptions. and I have made these remarks only to show that. Thus. The foregoing remarks lead me to say a few words on the protest lately made by some naturalists against the utilitarian doctrine that every detail of structure has been produced for the good of its possessor. we can hardly believe that the webbed feet of the upland goose. lately specified. though each being assuredly is well fitted for its place in nature. but this does not prove that they were formed solely for beauty or variety. in the wing of the bat. and consequently. horse and bat. we cannot believe that the similar bones in the arm of the monkey. many structures have now no very close and direct relation to present habits of life. on the principle of utility.

and others displeasure. Were the beautiful volute and cone shells of the Eocene epoch. and in many others. A similar line of argument holds good with fruits. instead of to the males alone. the other closed. reptiles. When the female is as beautifully coloured as the male. created that man might ages afterwards admire them in his cabinet? Few objects are more beautiful than the minute siliceous cases of the diatomaceae: were these created that they might be examined and admired under the higher powers of the microscope? The beauty in this latter case. destitute of nectar. in order that the fruit may be devoured and the matured seeds disseminated. one kind open and coloured so as to attract insects. if it be coloured of any brilliant tint. But this beauty serves merely as a guide to birds and beasts. which is not rarely the case with birds and butterflies. in the men of different races admiring an entirely different standard of beauty in their women. and in consequence at the same time beautiful. I infer that this is the case from having as yet found no exception to the rule that seeds are always thus disseminated when embedded within a fruit of any kind (that is within a fleshy or pulpy envelope). the reception of a peculiar kind of pleasure from certain colours. irrespective of any real quality in the admired object. I willingly admit that a great number of male animals. Several plants habitually produce two kinds of flowers. But this has been effected through sexual selection. I have come to this conclusion from finding it an invariable rule that when a flower is fertilised by the wind it never has a gaily-coloured corolla. forms and sounds--was first developed in the mind of man and of the lower animals. we may conclude that. Hence. on grasses. which are all fertilised through the agency of the wind. spinach. but they have been rendered conspicuous in contrast with the green leaves. If beautiful objects had been created solely for man's gratification. docks and nettles. the cause apparently lies in the colours acquired through sexual selection having been transmitted to both sexes. How the sense of beauty in its simplest form--that is. is a very obscure subject. On the other hand. and mammals. and that the idea of what is beautiful. our plants would not have been decked with beautiful flowers. some fishes. if insects had not been developed on the face of the earth. and not for the delight of man. We see this. that is. or rendered conspicuous by being white or black. as all our most gorgeous birds. so that they may be easily observed by insects. We may infer from all this that a nearly similar taste for beautiful colours and for musical sounds runs through a large part of the animal kingdom. and never visited by insects. is not innate or unalterable. is apparently wholly due to symmetry of growth. and the gracefully sculptured ammonites of the Secondary period. Flowers rank among the most beautiful productions of nature. have been rendered beautiful for beauty's sake. nut and ash trees. that a ripe strawberry or cherry is as pleasing to the eye as to the palate--that the gaily-coloured fruit of the spindle-wood tree and the scarlet berries of the holly are beautiful objects--will be admitted by everyone. for instance. So it is with the music of birds. not coloured. but would have produced only such poor flowers as we see on our fir.With respect to the belief that organic beings have been created beautiful for the delight of man--a belief which it has been pronounced is subversive of my whole theory--I may first remark that the sense of beauty obviously depends on the nature of the mind. it ought to be shown that before man appeared there was less beauty on the face of the earth than since he came on the stage. oak. The same sort of difficulty is presented if we enquire how it is that certain flavours and odours give pleasure. by the more beautiful males having been continually preferred by the females. and a host of magnificently coloured butterflies. Habit in all these cases appears .

But natural selection can and does often produce structures for the direct injury of other animals. each will be found on the whole advantageous. and profits by the structures of others. but there must be some fundamental cause in the constitution of the nervous system in each species. The correction for the aberration of light is said by Muller not to be perfect even in that most perfect organ. It is a much more probable view that the rattlesnake uses its rattle. And we see that this is the standard of perfection attained under nature. I cannot find even one which seems to me of any weight. the human eye. Natural selection will not produce absolute perfection. it will be modified. Snakes act on the same principle which makes the hen ruffle her feathers and expand her wings when a dog approaches her chickens. for natural selection acts solely by and for the good of each. The endemic productions of New Zealand. namely.to have come to a certain extent into play. But I have not space here to enlarge on the many ways by which animals endeavour to frighten away their enemies. If a fair balance be struck between the good and evil caused by each part. it would annihilate my theory. the being will become extinct. in order to warn the doomed mouse. as we see in the fang of the adder. or if it be not so. Natural selection cannot possibly produce any modification in a species exclusively for the good of another species. after describing in the strongest terms the wonderful powers of the human eye. for instance. I would almost as soon believe that the cat curls the end of its tail when preparing to spring. Natural selection tends only to make each organic being as perfect as. by which its eggs are deposited in the living bodies of other insects. for such could not have been produced through natural selection. one compared with another. adds these remarkable words: "That which we have discovered in the way of inexactness and imperfection in the optical machine and in the image on the retina. Although many statements may be found in works on natural history to this effect. to warn its prey." If our reason leads us to admire with enthusiasm a multitude of inimitable . It is admitted that the rattlesnake has a poison-fang for its own defence and for the destruction of its prey. Natural selection will never produce in a being any structure more injurious than beneficial to that being. in order to alarm the many birds and beasts which are known to attack even the most venomous species. as myriads have become extinct. the cobra expands its frill and the puff-adder swells while hissing so loudly and harshly. with this high standard under nature. or slightly more perfect than the other inhabitants of the same country with which it comes into competition. as Paley has remarked. whose judgment no one will dispute. is as nothing in comparison with the incongruities which we have just come across in the domain of the sensations. One might say that nature has taken delight in accumulating contradictions in order to remove all foundation from the theory of a pre-existing harmony between the external and internal worlds. are perfect. but some authors suppose that at the same time it is furnished with a rattle for its own injury. nor do we always meet. as far as we can judge. No organ will be formed. under changing conditions of life. Helmholtz. if any part comes to be injurious. for the purpose of causing pain or for doing an injury to its possessor. and in the ovipositor of the ichneumon. but they are now rapidly yielding before the advancing legions of plants and animals introduced from Europe. though throughout nature one species incessantly takes advantage of. After the lapse of time. If it could be proved that any part of the structure of any one species had been formed for the exclusive good of another species.

If we admire the several ingenious contrivances by which orchids and many other plants are fertilised through insect agency. and when the conditions of life did not insensibly graduate away from one part to another. from existing in greater numbers. which are utterly useless to the community for any other purpose. we can perhaps understand how it is that the use of the sting should so often cause the insect's own death: for if on the whole the power of stinging be useful to the social community. such as to produce galls. the intermediate variety will usually exist in lesser numbers than the two forms which it connects. Many of them are serious. can we consider as equally perfect the elaboration of dense clouds of pollen by our fir-trees. and that it has since been modified but not perfected for its present purpose. for instance. or to perish herself in the combat. cannot be withdrawn. We have seen in this chapter how cautious we should be in concluding that the most different habits of life could not graduate into each other. as a boring and serrated instrument.contrivances in nature. though we may easily err on both sides. and partly because the very process of natural selection implies the continual supplanting and extinction of preceding and intermediate gradations. that a bat. When two varieties are formed in two districts of a continuous area. owing to the backward serratures. but I think that in the discussion light has been thrown on several facts. and maternal love or maternal hatred. since intensified. and thus inevitably causes the death of the insect by tearing out its viscera? If we look at the sting of the bee. Closely allied species. and will thus generally succeed in supplanting and exterminating it. but we ought to admire the savage instinctive hatred of the queen-bee. during the course of further modification. so that a few granules may be wafted by chance on to the ovules? SUMMARY: THE LAW OF UNITY OF TYPE AND OF THE CONDITIONS OF EXISTENCE EMBRACED BY THE THEORY OF NATURAL SELECTION. though the latter fortunately is most rare. is all the same to the inexorable principles of natural selection. and are not linked together by a multitude of intermediate gradations. must often have been formed when the area was not continuous. an intermediate variety will often be formed. which. can we admire the production for this single purpose of thousands of drones. as soon as they are born. We have seen that species at any one period are not indefinitely variable. and at any one time acts only on a few forms. like that in so many members of the same great order. We have in this chapter discussed some of the difficulties and objections which may be urged against the theory. consequently the two latter. with the poison originally adapted for some other object. but from reasons assigned. which urges her to destroy the young queens. this same reason tells us. will have a great advantage over the less numerous intermediate variety. her daughters. If we admire the truly wonderful power of scent by which the males of many insects find their females. though it may cause the death of some few members. when used against many kinds of enemies. partly because the process of natural selection is always very slow. and which are ultimately slaughtered by their industrious and sterile sisters? It may be difficult. that some other contrivances are less perfect. fitted for an intermediate zone. as having existed in a remote progenitor. which on the belief of independent acts of creation are utterly obscure. could not have been formed by natural . it will fulfil all the requirements of natural selection. for undoubtedly this is for the good of the community. now living on a continuous area. Can we consider the sting of the bee as perfect.

ground woodpeckers. In many other cases. independently of any good having been thus gained. For instance. though it has become of such small importance that it could not. the one having been perfected whilst aided by the other. Hence we can understand. each good for its possessor. generally the smaller one. diving thrushes. organs. only in accordance with the standard of that particular country. that modifications in its structure could not have been slowly accumulated by means of natural selection. is enough to stagger any one. For in the larger country there will have existed more . we should be extremely cautious in concluding that none can have existed. and excretions highly useful or even indispensable. a swim-bladder has apparently been converted into an air-breathing lung. or it may have diversified habits. and then having been in part or in whole specialised for one function. In the cases in which we know of no intermediate or transitional states. then under changing conditions of life. and two distinct organs having performed at the same time the same function. if we know of a long series of gradations in complexity. The same organ having performed simultaneously very different functions. In each well-stocked country natural selection acts through the competition of the inhabitants and consequently leads to success in the battle for life. essential differences in their structure can almost always be detected. On the other hand. We may. modifications are probably the direct result of the laws of variation or of growth. organs serving for the same purpose and in external appearance closely similar may have been separately and independently formed. though it may well produce parts. been subsequently taken advantage of. have been acquired by means of natural selection. with some very unlike those of its nearest congeners.selection from an animal which at first only glided through the air. bearing in mind that each organic being is trying to live wherever it can live. also. We have seen that in two beings widely remote from each other in the natural scale. how it has arisen that there are upland geese with webbed feet. Hence the inhabitants of one country. Natural selection can produce nothing in one species for the exclusive good or injury of another. must often have largely facilitated transitions. Although the belief that an organ so perfect as the eye could have been formed by natural selection. for the good of species under new conditions of life. often yield to the inhabitants of another and generally the larger country. in its present state. and this again naturally follows from the same great principle. and still further modified. and this naturally follows from the principle of natural selection. or highly injurious to another species. for the metamorphoses of many organs show what wonderful changes in function are at least possible. We have seen that a species under new conditions of life may change its habits. But even such structures have often. as we may feel assured. believe that a part formerly of high importance has frequently been retained (as the tail of an aquatic animal by its terrestrial descendants). there is no logical impossibility in the acquirement of any conceivable degree of perfection through natural selection. but in all cases at the same time useful to the possessor. the common rule throughout nature is infinite diversity of structure for gaining the same end. but when such organs are closely examined. and petrels with the habits of auks. In many cases we are far too ignorant to be enabled to assert that a part or organ is so unimportant for the welfare of a species. yet in the case of any organ.

"Natura non facit saltum. On my theory.individuals. Natural selection will not necessarily lead to absolute perfection. Longevity--Modifications not necessarily simultaneous--Modifications apparently of no direct service--Progressive development--Characters of small functional importance. and the Conditions of Existence. have remained so. the law of the Conditions of Existence is the higher law. that of Unity of Type. but it would be useless to discuss all of them." This canon. but if we include all those of past times. For natural selection acts by either now adapting the varying parts of each being to its organic and inorganic conditions of life. It is generally acknowledged that all organic beings have been formed on two great laws--Unity of Type. I will devote this chapter to the consideration of various miscellaneous objections which have been advanced against my views. as well as the number and kinds of its inhabitants. is not strictly correct. being affected by the direct action of external conditions of life. the most constant--Supposed incompetence of natural selection to account for the incipient stages of useful structures--Causes which interfere with the acquisition through natural selection of useful structures--Gradations of structure with changed functions--Widely different organs in members of the same class. even if they were at any one time perfectly adapted to their conditions of life. that all are not as perfect as they might have been in relation to their conditions. and which is quite independent of their habits of life. as many have been made by writers who have not taken the trouble to understand the subject. and no one will dispute that the physical conditions of each country. Thus a distinguished German naturalist has asserted that the weakest part of my theory is. as some of the previous discussions may thus be made clearer. it must on this theory be strictly true. unless they themselves likewise changed. developed from one and the same source--Reasons for disbelieving in great and abrupt modifications. Hence. whether known or unknown. and subjected in all cases to the several laws of growth and variation. is fully embraced by the principle of natural selection. CHAPTER VII. through the inheritance of former variations and adaptations. unity of type is explained by unity of descent. can absolute perfection be everywhere predicated. On the theory of natural selection we can clearly understand the full meaning of that old canon in natural history. and this is shown to be the case by so many native forms in many quarters of the world having yielded their places to intruding foreigners. if we look to the present inhabitants alone of the world. as far as we can judge by our limited faculties. as it includes. or by having adapted them during past periods of time: the adaptations being aided in many cases by the increased use or disuse of parts. and thus the standard of perfection will have been rendered higher. The expression of conditions of existence. nor. MISCELLANEOUS OBJECTIONS TO THE THEORY OF NATURAL SELECTION. and the competition will have been severer. have undergone many . Nor can organic beings. and more diversified forms. that I consider all organic beings as imperfect: what I have really said is. in fact. so often insisted on by the illustrious Cuvier. By unity of type is meant that fundamental agreement in structure which we see in organic beings of the same class. when their conditions changed.

Ray Lankester has recently discussed this subject. if slight. the conditions of life. are closely similar or even identical with those now living. that longevity is generally related to the standard of each species in the scale of organisation. Moreover. can a variety live side by side with the parent species? If both have become fitted for slightly different habits of life or conditions. been largely determined through natural selection. be acquired by successive variations. as none of the animals and plants of Egypt. their varieties seem to be generally confined to distinct regions. they would . and he asks. inhabiting distinct stations. The fact of little or no modification having been effected since the glacial period. A critic has lately insisted. but this will be effected only under certain favourable circumstances. in the case of animals which wander much about and cross freely. The most striking modifications. that longevity is a great advantage to all species. whereas. Bronn. E. so probably have none in any part of the world. first in one part and then in another. and as they would be transmitted all together. at the close of his German translation of this work. as well as to the amount of expenditure in reproduction and in general activity. such as high land or low land. during the last several thousand years. as far as its extreme complexity allows him to form a judgment. the more permanent varieties are generally found. Bronn also insists that distinct species never differ from each other in single characters. this line of argument proves too much. survive from year to year by means of its seeds or ova? Mr.. in Egypt. on the principle of natural selection.mutations. in which the variability seems to be of a peculiar nature. The many animals which have remained unchanged since the commencement of the glacial period. how it always comes that many parts of the organisation should have been modified at the same time through variation and natural selection? But there is no necessity for supposing that all the parts of any being have been simultaneously modified. asks how. have changed during the last three or four thousand years. albinism. which implies that when variations or individual differences of a beneficial nature happen to arise. as Mr. they might live together. It has been argued that. yet all naturalists admit that such races have been produced through the modification of their original types. or embalmed. owing to advantages gained through natural selection. and yet. but is powerless against the doctrine of natural selection or the survival of the fittest. etc. might. of which we know anything.H. these will be preserved. it is probable. The celebrated palaeontologist. as was formerly remarked. as far as we know. for the ancient domestic races figured on the Egyptian monuments. as far as I can discover. and all mere temporary variations. would have been of some avail against those who believe in an innate and necessary law of development. And these conditions have. dry or moist districts. for these have been exposed to great changes of climate and have migrated over great distances. have remained absolutely uniform. so that he who believes in natural selection "must arrange his genealogical tree" in such a manner that all the descendants have longer lives than their progenitors! Cannot our critics conceive that a biennial plant or one of the lower animals might range into a cold climate and perish there every winter. such as size. G. but in many parts. and if we lay on one side polymorphic species. with some parade of mathematical accuracy. and he concludes. But. Lewes has remarked. would have been an incomparably stronger case. excellently adapted for some purpose.

and of the leaves on the axis. we have to allow for the direct and definite action of changed conditions of life. as cases in which natural selection could not have acted. A much more serious objection has been urged by Bronn. to be extremely cautious in pretending to decide what structures now are. however. Even when selection has been applied by man to some one character alone--of which our cultivated plants offer the best instances--it will invariably be found that although this one part. so will be other parts. an early developed part affecting one subsequently developed. almost all the other parts have been slightly modified. mutual pressure. but even in these cases. to the above objection is afforded by those domestic races which have been modified. but first one part and then another slightly modified and improved. due to some change in the conditions. Bud-variations. This may be attributed partly to the principle of correlated growth. the shape of the seed. of use to each species. such as an increased or diminished flow of nutriment to a part. chiefly through man's power of selection. namely. for some special purpose. and recently by Broca. it is almost certain that all the individuals of the species would be similarly modified. if we bear in mind the power of a minute drop of poison in producing complex galls. Nevertheless. He admits that natural selection has effected much. which we do not in the least understand. we ought. as well as for more strongly marked variations which occasionally arise. but if we could trace each step in the history of their transformation--and the latter steps can be traced--we should not see great and simultaneous changes.appear to us as if they had been simultaneously developed. . for the sake of brevity. this subject has been discussed by Nageli in an admirable essay. or at the greyhound and mastiff. and therefore cannot have been influenced through natural selection. or of a nectarine on a peach-tree. which appear to be quite unimportant for the welfare of the species. under the expression of the laws of growth. There is much force in the above objection. it should always be borne in mind that when one part is modified. Look at the race and dray-horse. whether it be the flower. fruit. and for so-called spontaneous variations. or have formerly been. or leaves. Their whole frames. With respect to plants. that many characters appear to be of no service whatever to their possessors. and if the unknown cause were to act persistently. and so forth--as well as through other causes which lead to the many mysterious cases of correlation. in which the nature of the conditions apparently plays a quite subordinate part. has been greatly changed. There must be some efficient cause for each slight individual difference. and partly to so-called spontaneous variation. and even their mental characteristics. the position of the ovules. but he insists that the families of plants differ chiefly from each other in morphological characters. we ought not to feel too sure that the above variations are not the effect of some local change in the nature of the sap. In the third place. offer good instances of spontaneous variations. through certain dimly seen causes. He consequently believes in an innate tendency towards progressive and more perfect development. The best answer. and a multitude of analogous cases. In the second place. when not of any use for dissemination. Bronn adduces the length of the ears and tails in the different species of hares and mice--the complex folds of enamel in the teeth of many animals. have been modified. in the first place. To these may be added the numerical divisions in the parts of the flower. etc. such as the appearance of a moss-rose on a common rose. These agencies may be all grouped together. He specifies the arrangement of the cells in the tissues.

Schobl. the external ears of the common mouse are supplied in an extraordinary manner with nerves. but now we know this to be the case. and in others the lower ones alone are fertilised. and their arrangement. hence the length of the ears can hardly be quite unimportant. I may mention that. but they are now known to be of the highest importance for the fertilisation of the species through the aid of insects. which a few years ago would have been considered as mere morphological differences without any special function. and the . Several plants belonging to distinct orders habitually produce flowers of two kinds--the one open. the position of the ovules. as they yield with the utmost safety a large stock of seed. and he suggests that this probably depends on the direction in which the pollen-tubes enter the ovarium. yet their use has not been. manifestly of high importance. of the ordinary structure. with the expenditure of wonderfully little pollen. could have been of any service. or of no physiological signification. in structure. As Bronn gives the length of the ears and tail in the several species of mice as instances. so that they no doubt serve as tactile organs. which are so highly developed that no one doubts that they are of importance. With respect to plants. though trifling ones. of differences in structure which can be of no special use. one ovule holds the former and a second ovule the latter position. and its use would be much influence by its length. and have probably been gained through natural selection. and the production of seed. as just stated. The petals in the imperfect flowers almost always consist of mere rudiments. even when one is erect and the other suspended within the same ovarium. With respect to the assumed inutility of various parts and organs. as it now seems probable. These two kinds of flowers sometimes differ wonderfully in structure. No one until lately would have imagined that in dimorphic and trimorphic plants the different lengths of the stamens and pistils.In the earlier editions of this work I underrated. yet may be seen to graduate into each other on the same plant. according to Dr. which before the principle of selection by man was well understood. to which on account of Nageli's essay I shall confine myself in the following remarks. the other closed and imperfect. and the benefits which certainly are derived from this process are thus secured. We shall. it is hardly necessary to observe that even in the higher and best-known animals many structures exist. These positions seem at first purely morphological. The two kinds of flowers often differ much. I can no more believe in this than that the well-adapted form of a race-horse or greyhound. presently see that the tail is a highly useful prehensile organ to some of the species. Hooker informs me that within the same ovarium the upper ovules alone in some cases. The ordinary and open flowers can be intercrossed. or has only recently been. it will be admitted that the flowers of the orchids present a multitude of curious structures. the frequency and importance of modifications due to spontaneous variability. excited so much surprise in the minds of the older naturalists. but Dr. If so. In certain whole groups of plants the ovules stand erect. But it is impossible to attribute to this cause the innumerable structures which are so well adapted to the habits of life of each species. and in others they are suspended. It may be worth while to illustrate some of the foregoing remarks. can thus be explained. would follow the selection of any slight deviations in position which favoured their fertilisation. The closed and imperfect flowers are. however. also. and within the same ovarium of some few plants. ascertained.

while all the other flowers on the plant are tetramerous. It is a more curious fact. de Jussieu. one flower. but being of very small size. opens first. In these several cases. according to A. two-valved and two-seeded siliques. one-seeded nutlets. and it can hardly be doubted that if all the flowers and leaves on the same plant had been subjected to the same external and internal condition. and in the upper part of the spike. All these modifications follow from the relative position and inter-action of the parts. which are considered by botanists to be generally of a highly important nature. or only in a quite subordinate manner. for the plants have never produced with me perfect flowers). In six out of thirty of the closed flowers in an Indian violet (name unknown. In one section of the Malpighiaceae the closed flowers. all would have been modified in the same manner. the sepals are reduced from the normal number of five to three. a sixth stamen standing opposite to a petal being alone developed. or occurring on . colour and other characters. are still further modified.pollen-grains are reduced in diameter. for the five stamens which stand opposite to the sepals are all aborted. lanceolate. and to reduce the amount of pollen. ribbed. with the exception of that of the well-developed ray-florets. during the progress of the reduction of the pollen and the closure of the flowers. but must have followed from the laws of growth. according to Tausch. the style is aborted. In the common rue and some other plants. In many Compositae and Umbelliferae (and in some other plants) the circumferential flowers have their corollas much more developed than those of the centre. are orthospermous. In the Spanish chestnut. that I will give some additional cases of another kind. Professor Braun mentions a Fumariaceous genus. It is so necessary to appreciate the important effects of the laws of growth. In numerous other cases we find modifications of structure. according to Schacht. in the nearly horizontal and in the upright branches. In Ononis columnae five of the alternate stamens are rudimentary. usually the central or terminal one. and in some species of Viola three stamens are in this state. and in Hyoseris the same head yields achenes of three different forms. the angles of divergence of the leaves differ. and the central one coelospermous. due to differences in relative position on the same plant. and this stamen is not present in the ordinary flowers of this species. in which the flowers in the lower part of the spike bear oval. natural selection cannot. and has five sepals and petals. which are of service in making the flowers conspicuous to insects. In Carthamus and some other Compositae the central achenes alone are furnished with a pappus. have come into play. yet hardly any of the above special modifications can have been thus determined. as far as we can judge. and five divisions to the ovarium. affecting only some of the flowers on the same plant. namely of differences in the same part or organ. while the surrounding flowers generally have three calyx-lobes with the other organs pentamerous. and the ovaria are reduced from three to two. as are the flowers and leaves in certain positions. when rendered by the closure of the flowers superfluous. Now although natural selection may well have had the power to prevent some of the flowers from expanding. including the functional inactivity of parts. previously referred to. that the achenes or seeds of the circumference and centre sometimes differ greatly in form. In the British Adoxa the uppermost flower generally has two calyx-lobes with the other organs tetramerous. and this is a character which was considered by De Candolle to be in other species of the highest systematic importance. two retaining their proper function. and in certain fir-trees. In certain Umbelliferae the exterior seeds. and this seems often connected with the abortion of the reproductive organs.

closed flowers. we cannot even attribute them. to any proximate agency. with either one or two ovaries. pentamerous. from the nature of the organism and of the conditions. etc. The manner in which the petals are folded in the bud is in most groups a very constant morphological character. From the fact of the above characters being unimportant for the welfare of the species. it must be one of retrogression rather than of progression. The acquisition of a useless part can hardly be said to raise an organism in the natural scale. but in some species flowers may be found on the same plant. of whatever importance they may generally be to us for our classifications. any slight variations which occurred in them would not have been accumulated and augmented through natural selection. A structure which has been developed through long-continued selection. they may be. independently of natural selection. I may mention that. that the plants have been caught in the act of progressing towards a higher state of development? On the contrary.distinct plants. the aestivation is almost as frequently that of the Rhinanthideae as of the Antirrhinideae. As these variations seem of no special use to the plants. I should infer from the mere fact of the parts in question differing or varying greatly on the same plant. for it will no longer be regulated by this same power of selection. that I need not give examples. and in the case of the imperfect. and in H. s'etend entre le pericarpe et le placenta. if any new principle has to be invoked. and . to which latter tribe the genus belongs." We thus see that with plants many morphological changes may be attributed to the laws of growth and the inter-action of parts. mutabile. and in this case all the individuals of the species would be modified in the same manner. they cannot have been influenced by natural selection. but he subsequently saw them growing on the same bush. but if the unknown cause were to act almost uniformly for a length of time. we may infer that the result would be almost uniform. when it ceases to be of service to a species. But with respect to Nageli's doctrine of an innate tendency towards perfection or progressive development. Hilaire gives the following cases: the genus Zanthoxylon belongs to a division of the Rutaceae with a single ovary. "Voila donc dans un meme individu des loges et un style qui se rattachent tantot a un axe verticale et tantot a un gynobase. and even in the same panicle. Masters has observed instances of both marginal and free central placentation. but as numerical variations are comparatively rare when the parts are few. Hilaire found towards the southern extreme of the range of Gomphia oleaeformis two forms which he did not at first doubt were distinct species. the flowers of Papaver bracteatum offer either two sepals with four petals (which is the common type with poppies). as in the last class of cases. "Une lame PLUS OU MOINS LARGE. In Helianthemum the capsule has been described as unilocular or tri-locular. but Professor Asa Gray states that with some species of Mimulus. flowers indifferently tetramerous. Aug. can it be said in the case of these strongly pronounced variations. as we see with rudimentary organs. St. St. Of their cause we are quite ignorant. above described. or three sepals with six petals. and he then adds. generally becomes variable. such as relative position. But when. according to De Candolle. that such modifications were of extremely small importance to the plants themselves. which grow close together under the same conditions. We are ignorant of the exciting cause of the above specified modifications. Lastly. It is so common to observe on the same plant." In the flowers of Saponaria officinalis Dr. modifications have been induced which are unimportant for the welfare of the species. I will give only a few instances.. and so it must be with many parasitic and degraded animals.

For the best definition which has ever been given of a high standard of organisation. Mivart's objections will be. Although we have no good evidence of the existence in organic beings of an innate tendency towards progressive development. independently of natural selection. as well as through the intercrossing of distinct individuals. The one new point which appears to have struck many readers is. for as these morphological characters do not affect the welfare of the species. as propounded by Mr. who may wish to weigh the evidence on both sides." This subject is intimately connected with that of the gradation of the characters. When thus marshalled. I never before felt so strongly convinced of the general truth of the conclusions here arrived at. and scales to all true reptiles. It cannot have been of much importance to the greater number of mammals. otherwise modified. of course. is the degree to which the parts have been specialised or differentiated. inasmuch as the parts are thus enabled to perform their functions more efficiently. the divisions of the flower or of the ovarium. feathers or scales. in so intricate a subject. When discussing special cases. Mr. All Mr. that characters of slight vital importance to the species. no slight effort of reason and memory is left to the reader. has recently collected all the objections which have ever been advanced by myself and others against the theory of natural selection. Mivart passes over the effects of the increased use and disuse of parts. whatever it may be. and have treated in my "Variation under Domestication" at greater length than. but not through natural selection. but. points which were discussed in the last chapter . yet hair has been transmitted to almost all mammals. often accompanied by a change of function. subject. "That natural selection is incompetent to account for the incipient stages of useful structures. as I have attempted to show in the fourth chapter. Mivart's book. for instance. considered in the present volume. which I have always maintained to be highly important. or reptiles. My judgment may not be trustworthy. Wallace and myself. A distinguished zoologist. which we consider as important--such as the arrangement of the leaves. and has illustrated them with admirable art and force. St. or have been. as I am inclined to believe. through the continued action of natural selection. and consequently is often assumed to be of high vital importance to the species. He likewise often assumes that I attribute nothing to variation. the position of the ovules.. Mr. as we shall hereafter see when we treat of the genetic principle of classification.apparently often have been. as I believe. etc. they make a formidable array. any slight deviations in them could not have been governed or accumulated through this latter agency. feathers to all birds. yet this necessarily follows. A structure. birds. Thus. It is a strange result which we thus arrive at. George Mivart. descendants. are the most important to the systematist. whether they were clothed with hair. which is common to many allied forms. to much partial error. transmitted in nearly the same state to numerous. this is by no means so paradoxical as it may at first appear. and comparing each section with what I have said on the same head. and natural selection tends towards this end. Mivart's plan to give the various facts and considerations opposed to his conclusions. first appeared in many cases as fluctuating variations. any other writer. whereas in the work just referred to I have collected a greater number of well-established cases than can be found in any other work known to me. but after reading with care Mr. namely. is ranked by us as of high systematic importance. and as it forms no part of Mr. which sooner or later became constant through the nature of the organism and of the surrounding conditions. morphological differences. the conversion of a swim-bladder into lungs.

while the individuals less favoured in the same respects will have been the most liable to perish. if not fed by their owners. selecting those which are the most illustrative. to a supply of food. which exactly corresponds with what I have called unconscious selection by man.under two headings. will often have been preserved. These will have intercrossed and left offspring. The giraffe. One is that the increased size of the body would obviously require an increased supply of food." But as the giraffe does actually exist in large numbers in Africa. allowing them freely to intercross. as man does. These slight proportional differences. and this must be a great advantage to it during dearths. as with the race-horse and greyhound. at each stage of increased size. Mivart's objections. combined.. either inheriting the same bodily peculiarities. no doubt. why should we doubt that. in which careful measurements are given. head and tongue. By this process long-continued. or with a tendency to vary again in the same manner. by breeding from the victorious birds. are not of the slightest use or importance to most species. or as with the game-cock. The Niata cattle in South America show us how small a difference in structure may make. left untouched by the other hoofed quadrupeds of the country. reeds. it may be well to explain once again how natural selection will act in all ordinary cases. It can thus obtain food beyond the reach of the other Ungulata or hoofed animals inhabiting the same country. and he considers it as "very problematical whether the disadvantages thence arising would not. But it will have been otherwise with the nascent giraffe. To this conclusion Mr. for they will have roamed over the whole country in search of food. much elongated neck. would have . in times of scarcity. These cattle can browse as well as others on grass. fore legs. but from the projection of the lower jaw they cannot. etc. a great difference in preserving an animal's life. That the individuals of the same species often differ slightly in the relative lengths of all their parts may be seen in many works of natural history. subjected as now to severe dearths. Man has modified some of his animals. by its lofty stature. for those individuals which had some one part or several parts of their bodies rather more elongated than usual. Assuredly the being able to reach. it seems to me almost certain that an ordinary hoofed quadruped might be converted into a giraffe. has its whole frame beautifully adapted for browsing on the higher branches of trees. without necessarily having attended to special points of structure. more than counterbalance the advantages. would generally have survived. as far as size is concerned. during such periods. considering its probable habits of life. Mivart brings forward two objections. and as some of the largest antelopes in the world. Before coming to Mr. We here see that there is no need to separate single pairs. due to the laws of growth and variation. by simply preserving and breeding from the fleetest individuals. and will destroy all the inferior individuals. Nevertheless. in a most important manner with the inherited effects of the increased use of parts. I will here consider in some detail several of the cases advanced by Mr. So under nature with the nascent giraffe. when he methodically improves a breed: natural selection will preserve and thus separate all the superior individuals. the individuals which were the highest browsers and were able during dearths to reach even an inch or two above the others. during the often recurrent droughts. so that at these times the Niatas perish. to which food the common cattle and horses are then driven. as want of space prevents me from considering all. Mivart. intermediate gradations could formerly have existed there. taller than an ox. abound there. browse on the twigs of trees.

This animal also uses its long neck as a means of offence or defence. why has not any member of the group acquired a long proboscis? With respect to South Africa. and against this animal. for which hoofed animals are singularly ill-constructed) implies greatly increased bulk of body. we see the lower branches trimmed or planed to an exact level by the browsing of the horses or cattle. see in a general manner that various causes might have interfered with the development of a long neck or proboscis. guanaco and macrauchenia? Or. Mr. and it is almost equally certain that this one kind alone could have its neck elongated for this purpose. in a lesser degree. Nor must we overlook the fact. though it is so luxuriant. nor why the later tertiary periods should have been much more favourable for their existence than the present time. for instance South America. and that some species are much more variable than others.been of some advantage to the nascent giraffe. it does not follow that natural selection would be able to act on them and . as Sir S. Chauncey Wright has remarked. and if high browsing be so great an advantage. and what advantage would it be. if natural selection be so potent. various animals belonging to this same order have not acquired either an elongated neck or a proboscis. for instance. in other quarters of the world. the camel. by violently swinging its head armed with stump-like horns. its tall neck--and the taller the better--would. great and small. again. serve as a watch-tower. In order that an animal should acquire some structure specially and largely developed. which was formerly inhabited by numerous herds of the giraffe. It is from this cause. it is almost indispensable that several other parts should be modified and coadapted. through natural selection and the effects of increased use. besides the giraffe. why has not any other hoofed quadruped acquired a long neck and lofty stature. if kept there. the answer is not difficult. and can best be given by an illustration. Why. but by the union of all. that increased bulk would act as a protection against almost all beasts of prey excepting the lion. In every meadow in England. and we know that some areas support singularly few large quadrupeds. Baker remarks. We are ignorant with respect to the conditions which determine the numbers and range of each species. but it is as unreasonable to expect a distinct answer to such a question as why some event in the history of mankind did not occur in one country while it did in another. Whatever the causes may have been. Although every part of the body varies slightly. however. that no animal is more difficult to stalk than the giraffe. With the different species of our domesticated animals we know that the parts vary in a different manner and degree. and not with the other ungulate animals. it does not follow that the necessary parts should always vary in the right direction and to the right degree. we can see that certain districts and times would have been much more favourable than others for the development of so large a quadruped as the giraffe. to sheep. cannot be distinctly answered. To reach the foliage at a considerable height (without climbing. as Mr. while South Africa abounds with them to an unparalleled degree. The preservation of each species can rarely be determined by any one advantage. Mivart then asks (and this is his second objection). We can. and. and we cannot even conjecture what changes of structure would be favourable to its increase in some new country. Even if the fitting variations did arise. in which trees grow. In South Africa the competition for browsing on the higher branches of the acacias and other trees must be between giraffe and giraffe. Why this should be so we do not know. to acquire slightly longer necks? In every district some one kind of animal will almost certainly be able to browse higher than the others.

might. In each case various causes. natural selection is a slow process. but when the power of true flight had once been acquired. or for avoiding falls. and assigns certain reasons in answer. is a highly complex affair. Objections of the same nature as the foregoing have been advanced by many writers.produce a structure which apparently would be beneficial to the species. might at last be converted into an animal so thoroughly aquatic as to brave the open ocean. it would never be reconverted back. in modifying any particular structure for obtaining food. yet as some of these bats are peculiar species. Gradations of structure. Lastly. then in streams or lakes. in many quarters of the world. Bats. by the aid of their hind legs alone. Lyell asks. with each step beneficial.--as seems often to be the case. besides the general ones just indicated. for the bats ground-insects would serve as food. etc. or will be greatly retarded. seeing that no one can solve the . as formerly shown. have probably interfered with the acquisition through natural selection of structures. they must have long inhabited their present homes. or completely lost. as such development would be advantageous to all? Why have not apes acquired the intellectual powers of man? Various causes could be assigned. A strictly terrestrial animal. But seals would not find on oceanic islands the conditions favourable to their gradual reconversion into a terrestrial form. at least for the above purposes. through disuse. probably acquired their wings by at first gliding through the air from tree to tree. and the same favourable conditions must long endure in order that any marked effect should thus be produced. but as they are conjectural. will be favoured only under certain peculiar conditions. A definite answer to the latter question ought not to be expected. One writer asks. by occasionally hunting for food in shallow water. Oceanic islands are inhabited by bats and seals. which first colonise and abound on most oceanic islands. with each stage beneficial to a changing species. for the former there would be no prey. like many birds. which it is thought would be beneficial to certain species. have had their wings greatly reduced in size. why have not seals and bats given birth on such islands to forms fitted to live on the land? But seals would necessarily be first converted into terrestrial carnivorous animals of considerable size. and that there is nothing strange in a transition not having occurred in any particular case. Except by assigning such general and vague reasons. but these would already be largely preyed on by the reptiles or birds. For instance. Bats. like the so-called flying squirrels. then natural selection will be able to do little. Lastly. for the sake of escaping from their enemies. so as to compete with birds or other ground animals. but by no terrestrial mammals. These conjectural remarks have been made merely to show that a transition of structure. but in this case it would be necessary that they should first have acquired the power of running quickly on the ground. and their relative probability cannot be weighed. and for such a change a bat seems singularly ill-fitted. into the less efficient power of gliding through the air. more than one writer has asked why have some animals had their mental powers more highly developed than others. indeed. if the number of individuals existing in a country is determined chiefly through destruction by beasts of prey--by external or internal parasites. and bats into terrestrial insectivorous animals. why has not the ostrich acquired the power of flight? But a moment's reflection will show what an enormous supply of food would be necessary to give to this bird of the desert force to move its huge body through the air. we cannot explain why. it would be useless to give them. Therefore Sir C. hoofed quadrupeds have not acquired much elongated necks or other means for browsing on the higher branches of trees.

Insects often resemble for the sake of protection various objects. The caterpillars which project motionless like dead twigs from the bushes on which they feed." But in all the foregoing cases the insects in their original state no doubt presented some rude and accidental resemblance to an object commonly found in the stations frequented by them. for in every group the characters which differ in the several species are the most apt to vary. and in these having become more or less green-coloured. which resembles "a stick grown over by a creeping moss or jungermannia. spines. why. offer an excellent instance of a resemblance of this kind. excrement of birds. or other object. but as the case stands there is none. that a native Dyak maintained that the foliaceous excrescences were really moss. for natural selection to seize upon and perpetuate." as in the case given by Mr. Mr. Mivart's objection. there is nothing improbable in this insect having varied in the irregularities on its surface.simpler problem. As some rude resemblance is necessary for the first start. and this chiefly in colour. of two races of savages. are the most . are rare and exceptional. and this apparently implies increased brain power. "As. The cases of the imitation of such objects as the excrement of birds. and is not confined to colour. Wallace. The resemblance is often wonderfully close. we can understand how it is that the larger and higher animals do not (with the exception. and that it varied slightly in many ways. dead twigs. bits of lichen. flowers. and every grade in resemblance which aided an insect to escape notice or detection. then all the variations which rendered the insect at all more like any such object. Mivart remarks. Nor can I see any force in Mr. they would be eliminated. one has risen higher in the scale of civilisation than the other. or. but to this latter point I shall hereafter recur. Mivart's difficulty with respect to "the last touches of perfection in the mimicry. Darwin's theory. On this head. there is a constant tendency to indefinite variation. according to Mr. and the more perfect the resemblance so much the better for the insect. and even to the manner in which the insects hold themselves. and living insects. would be preserved." So close was this resemblance. they must tend to neutralize each other. Assuming that an insect originally happened to resemble in some degree a dead twig or a decayed leaf. but only the surface which commonly surrounds them. and thus favoured its escape. and at first to form such unstable modifications that it is difficult. such as green or decayed leaves. There would indeed be force in Mr. while the generic characters. of one fish) resemble for the sake of protection special objects. bamboo. of a walking-stick insect (Ceroxylus laceratus). if not impossible. as far as I know. Considering the nature of the differences between the species in the group which includes the above Ceroxylus. and as the minute incipient variations will be in ALL DIRECTIONS. considering the almost infinite number of surrounding objects and the diversity in form and colour of the hosts of insects which exist. independently of natural selection. while other variations would be neglected and ultimately lost. but extends to form. Nor is this at all improbable. if they rendered the insect at all less like the imitated object. or those common to all the species. Insects are preyed on by birds and other enemies whose sight is probably sharper than ours. if we were to attempt to account for the above resemblances. Mivart's other objections. to see how such indefinite oscillations of infinitesimal beginnings can ever build up a sufficiently appreciable resemblance to a leaf. through mere fluctuating variability. We will return to Mr. would tend towards its preservation.

and placed transversely to the longer axis of the mouth. or even fifteen feet in length. subsist by sifting the mud and water. which clothe the whole gigantic palate. two-thirds of the length of the baleen in this species of whale. and the baleen. I wish only to show that this is not incredible. why should not the early progenitors of the whales with baleen have possessed a mouth constructed something like the lamellated beak of a duck? Ducks. on each side of the upper jaw. four feet. and in being thus furnished it differs conspicuously from the lower jaw of a whale. elastic lamellae. in another eighteen inches. Within the main row there are some subsidiary rows. With respect to the baleen. each of service to its possessor. being about one-third of an inch in length. as they project inward. instead of straight downward. Those standing towards the middle are the longest. the upper mandible alone is furnished with lamellae. of about 300 plates or laminae. in another three.constant. is about one-eighteenth of the length of the head of a moderately large Balaenoptera rostrata. in which species the baleen is only nine inches long. They arise from the palate. In the genus Prion. like whales. Mr. and serve to strain or sift the water. and the family has sometimes been called Criblatores. but finer. one of its greatest peculiarities. Mivart remarks that if it "had once attained such a size and development as to be at all useful. obliquely bevelled so as to be pointed. I hope that I may not be misconstrued into saying that the progenitors of whales did actually possess mouths lamellated like the beak of a duck. the middle lamina being in one species. so that the beak of this . The extremities and inner margins of all the plates are frayed into stiff bristles. But towards the extremity of the beak they differ much. which is destitute of baleen. and in the Balaenoptera rostrata only about nine inches in length. or sifters. and they project fourteen one-hundredths of an inch beneath the edge. The upper mandible is furnished on each side (in the specimen examined by me) with a row or comb formed of 188 thin. that is. and are attached by flexible membrane to the sides of the mandible. The beak of a shoveller-duck (Spatula clypeata) is a more beautiful and complex structure than the mouth of a whale. it may be asked. The middle and longest lamina in the Greenland whale is ten. which stand close together transversely to the longer axis of the mouth. the extremities of these lower lamellae are frayed into fine bristly points. The Greenland whale is one of the most wonderful animals in the world. but in the different species of Cetaceans there are gradations in length. though incomparably less bulky. The lower mandible of the shoveller-duck is furnished with lamellae of equal length with these above. and that the immense plates of baleen in the Greenland whale might have been developed from such lamellae by finely graduated steps. The baleen consists of a row. a member of the distinct family of the Petrels. On the other hand. according to Scoresby. In these several respects they resemble the plates of baleen in the mouth of a whale. so that they thus curiously resemble the plates of baleen. and thus to secure the minute prey on which these great animals subsist. twelve. the lamellae would be six inches in length. At their bases there is a short subsidiary row of obliquely transverse lamellae. But how to obtain the beginning of such useful development?" In answer. which are well developed and project beneath the margin. or whalebone. so that if we were to make the head of the shoveller as long as that of the Balaenoptera. then its preservation and augmentation within serviceable limits would be promoted by natural selection alone. The quality of the whalebone also differs in the different species. The entire head of the shoveller.

The Hyperoodon bidens is destitute of true teeth in an efficient condition.bird resembles in this respect the mouth of a whale. In this latter bird the lamellae of the upper mandible are much coarser than in the common duck. is grass. through the beak of the Merganetta armata." Its chief food. yet this bird. and which. as far as fitness for sifting is concerned. The edges of the lower mandible are crossed by numerous fine ridges. Salvin). but its palate is roughened. constantly uses it for this purpose. Bartlett. Turning to another group of the same family. but uses its beak exclusively for tearing or cutting herbage. In this latter species the lamellae are much coarser than in the shoveller. recurved teeth. for seizing or tearing solid food. and terminating upward in teeth-like knobs. which project very little. lastly. The edges of the lower mandible are serrated with teeth much more prominent. almost confluent. they are only about fifty in number on each side. and in some respects through that of the Aix sponsa. like those of the Merganser (a member of the same family). coarser and sharper than in the duck. From the highly developed structure of the shoveller's beak we may proceed (as I have learned from information and specimens sent to me by Mr. about twenty-seven in number on each side. hard points of horn. serving for the widely different purpose of securing live fish. The palate is also covered with hard rounded knobs. except the hooked tip. as if for crushing food. They are square-topped. The common goose does not sift the water. with a beak constructed like that of a common goose and adapted solely for grazing. nor so distinct from each other. as I hear from Mr. for which purpose it is so well fitted that it can crop grass closer than almost any other animal. without any great break. but rather more regularly placed. it will hardly be denied that the points might have been converted through variation and natural selection into lamellae as well-developed as those of the Egyptian . Returning to the whales. but the lamellae are not so numerous. therefore. The beak of a goose. in which the lamellae are less developed than in the common goose. with small unequal. There are other species. Although the beak is thus very inferior as a sifter to that of a shoveller. as I may add. nothing improbable in supposing that some early Cetacean form was provided with similar points of horn on the palate. into one like the shoveller. in which the lamellae are considerably less developed than in the common duck. E. as I hear from Mr. as I am informed by Mr. to the beak of the common duck. like the knobs on the beak of the goose. If so. There are other species of geese. which it crops like the common goose. Salvin. however. Bartlett. We thus see that a member of the duck family. as every one knows. provided with a beak almost exclusively adapted for sifting the water. according to Lacepede. "uses its bill like a duck by throwing the water out at the corners. but I do not know whether they use their beaks for sifting the water. for this bird could hardly use any part of its beak. might also be converted by small changes into one provided with prominent. aided it in seizing or tearing its food. nor do they project so much inward. yet this goose. and are edged with translucent. and are firmly attached to the sides of the mandible. There is. and do not project at all beneath the margin. In the Egyptian goose (Chenalopex) the beak closely resembles that of the common duck. hardish tissue. might be converted by small changes into a species like the Egyptian goose--this into one like the common duck--and. or even a member with a beak having less well-developed lamellae.

far from clear. such as soles. and they do this so vigorously that the eye is pressed hard against the upper part of the orbit. however. cannot long retain a vertical position. We should bear in mind that each species of duck is subjected to a severe struggle for existence. The Pleuronectidae. The different members. they stand opposite to each other. The forehead between the eyes consequently becomes. Soon the eye proper to the lower side begins to glide slowly round the head to the upper side. in which I quite agree with him. however. and the whole body is then symmetrical. it could not be used by the fish while lying in its habitual position on one side. then into lamellae like those of the domestic duck. While thus at rest they often twist. as could be . which does not in any considerable degree alter the shape in which it leaves the ovum. soon growing tired. and to their being destitute of a swim-bladder. and facility for feeding on the ground. "a long series of forms exhibiting a gradual transition from Hippoglossus pinguis. Mivart has taken up this case. it is of a white colour. gradations. less developed in many ways than the upper side. in which case they would have served exclusively as a sifting apparatus. which are entirely thrown to one side. that such an incipient transformation must rather have been injurious. with the functions of the parts slowly changing during the progress of development. The lower eye would. From this stage. or Flat-fish. indeed. while very young and still symmetrical." But he might have found an answer to this objection in the excellent observations published in 1867 by Malm. The lower. He then adds: "If the transit was gradual. are remarkable for their asymmetrical bodies. The chief advantages thus gained seem to be protection from their enemies.goose. the lower eye upward. But the eyes offer the most remarkable peculiarity. is manifest from several species. That the Pleuronectidae are admirably adapted by their flattened and asymmetrical structure for their habits of life. of the family present. have been liable to be abraded by the sandy bottom. until they became as well constructed as those of the shoveller. or resting-surface. lead us onward to the enormous plates of baleen in the Greenland whale. They rest on one side--in the greater number of species on the left. and that the structure of every part of its frame must be well adapted to its conditions of life. and so onward. as Malm observed. the small size of their lateral fins. for they are both placed on the upper side of the head. then how such transit of one eye a minute fraction of the journey towards the other side of the head could benefit the individual is. with the lateral fins often of smaller size. and remarks that a sudden spontaneous transformation in the position of the eyes is hardly conceivable. Hence. as Schiodte remarks. flounders. also. to see above them. The Pleuronectidae. in which the lamellae would be two-thirds of the length of the plates of baleen in the Balaenoptera rostrata. in which case they would have been used both for seizing objects and for sifting the water. which may be observed in still-existing Cetaceans. with their eyes standing on opposite sides of the head. as was formerly thought to be the case. being extremely common. with both sides equally coloured. and occasionally reversed adult specimens occur. It is obvious that unless the lower eye did thus travel round. During early youth. but in some on the right side.. owing to the excessive depth of their bodies. It seems. resembles at first sight the ventral surface of an ordinary fish. etc. Nor is there the least reason to doubt that each step in this scale might have been as serviceable to certain ancient Cetaceans. they fall to the bottom on one side. to the soles. as are the gradations in the beaks of the different existing members of the duck-family. but does not pass right through the skull. even." Mr.

which is under-most. But it cannot be supposed that the peculiar speckled appearance of the upper side of the sole. and swims diagonally through the water. and no permanent effect is thus produced. and their skulls are thus rendered rather crooked. in opposition to some other naturalists. of changing their colour in accordance with the surrounding surface. We may also attribute to the inherited effects of use the fact of the mouth in several kinds of flat-fish being bent towards the lower surface. whether it be the right or left. on the other hand. that the skull yields and is altered in shape. of feeding with ease on the ground. including the lateral fins. to twenty-five to thirty in the lower halves. as "there is so much less room for their action than with the larger fins above. Malm states that the newly-hatched young of perches. of which I have given a figure. however. and he has observed that they often then strain their lower eyes so as to look upward. no doubt beneficial to the individual and to the species. we could understand how it is that certain species. while resting on one side at the bottom. salmon. On one occasion Malm saw a young fish raise and depress the lower eye through an angular distance of about seventy degrees. Disuse. and on the position of the eyes. Mivart considers would be injurious. With the Pleuronectidae. the eyeless side of the head. if the skin or muscles be permanently contracted through disease or some accident. with the jaw bones stronger and more effective on this. are soon able to hold themselves in a vertical position. its weight drags forward all the bones of the skull on the same side. by remarking that "the author gives a very simple explanation of the abnormal condition of the Pleuronectoids. and a permanent effect is thus produced on the form of the head. have the habit of occasionally resting on one side at the bottom. even after early youth. the older they grow the more habitually they rest on one side. which is not a member of the Pleuronectidae. that the adult Trachypterus arcticus. and in this fish. of endeavouring to look upward with both eyes. are due to . so that it readily yields to muscular action. as recently shown by Pouchet. the two sides of the head are said to be somewhat dissimilar. Judging from analogy. and if this be so. so like the sandy bed of the sea. concludes his abstract of Malm's paper. or the power in some species. It is also known with the higher animals. and other species on the right side. Malm adds. we may reasonably suppose that the absence of colour in flat-fish on the side. temporarily contracted in breadth. owing to the increasing flatness of their bodies. With long-eared rabbits. From the colourless state of the ventral surface of most fishes and of many other animals." We thus see that the first stages of the transit of the eye from one side of the head to the other. or the presence of bony tubercles on the upper side of the turbot. These fishes. that the Pleuronectidae are not quite symmetrical even in the embryo. the tendency to distortion would no doubt be increased through the principle of inheritance. which Mr. Gunther. if one ear flops forward and downward. while young. may be attributed to the habit. than on the other. Traquair supposes. rests on its left side at the bottom. will account for the less developed condition of the whole inferior half of the body. though Yarrel thinks that the reduced size of these fins is advantageous to the fish. Schiodte believes. as Dr. in confirmation of the above view. Our great authority on Fishes.plainly seen. We should remember that the skull at this early age is cartilaginous and flexible. for the sake. on the other hand." Perhaps the lesser number of teeth in the proportion of four to seven in the upper halves of the two jaws of the plaice. and several other symmetrical fishes. is due to the exclusion of light. may likewise be accounted for by disuse. Dr. habitually fall over and rest on the left side.

and thus aided themselves in climbing. however. The mammary glands are common to the whole class of mammals. possible that the long tail of this monkey may be of more service to it as a balancing organ in making its prodigious leaps. and we can know nothing positively about their manner of development. so as to partake of the nature of milk." But there is no necessity for any such belief. It is admitted by most evolutionists that mammals are descended from a marsupial form. remarks on this structure: "It is impossible to believe that in any number of ages the first slight incipient tendency to grasp could preserve the lives of the individuals possessing it. almost before they deserved to be thus designated. Lockwood. Now. they must. than as a prehensile organ. as is the case with some members of the same order. and this almost implies that some benefit great or small is thus derived. and at the same time they hooked their little tails round that of their mother. The extremity of the tail in some American monkeys has been converted into a wonderfully perfect prehensile organ. and how much to natural selection. and if so. what chance was there of the perpetuation of such a variation?" But the case is not here put fairly. who has seen a mouse thus suspend itself. Mivart in every detail. that the inherited effects of the increased use of parts. therefore. For all spontaneous variations in the right direction will thus be preserved. If the harvest mouse had been more strictly arboreal. would in all probability suffice for the work. or favour their chance of having and of rearing offspring. Mr. In the case of the fish (Hippocampus) the eggs are hatched. and serves as a fifth hand. and many other peculiarities. Professor Henslow kept in confinement some harvest mice (Mus messorius) which do not possess a structurally prehensive tail. is it not at least possible that the young might have been similarly nourished? And in this case. Mr.the action of the light. as I have before insisted. and an American naturalist. it would be difficult to say. believes from what he has seen of the development of the young. that they are nourished by a secretion from the cutaneous glands of the sack. Why Cercopithecus. Mivart asks: "Is it conceivable that the young of any animal was ever saved from destruction by accidentally sucking a drop of scarcely nutritious fluid from an accidentally hypertrophied cutaneous gland of its mother? And even if one was so. How much to attribute in each particular case to the effects of use. as will those individuals which inherit in the highest degree the effects of the increased and beneficial use of any part. the mammary glands will have been at first developed within the marsupial sack. will be strengthened by natural selection. in some degree or manner the most nutritious. to their habits of life. Here natural selection has probably come into play. Habit. have been developed at an extremely remote period. Brehm saw the young of an African monkey (Cercopithecus) clinging to the under surface of their mother by their hands. A reviewer. has not become thus provided. would in the long run have reared a larger number . and perhaps of their disuse. and are indispensable for their existence. who agrees with Mr. the individuals which secreted a fluid. but he frequently observed that they curled their tails round the branches of a bush placed in the cage. It is. with the early progenitors of mammals. within a sack of this nature. and the young are reared for a time. it would perhaps have had its tail rendered structurally prehensile. Gunther. it seems impossible to decide. considering its habits while young. I may give another instance of a structure which apparently owes its origin exclusively to use or habit. I have received an analogous account from Dr. as well as in adapting the general shape of the body of these fishes. We should keep in mind.

the effects of use. Mivart. and afterwards transmitted to the offspring at an earlier age. and one of these apparently is defence. nearly indefinite . neatly fitting together and placed on the summit of a flexible stem. called pedicellariae. The Echinodermata (star-fishes. could hardly have been used with full force as long as the larynx entered the nasal passage. and so safely attains the gullet behind it. while the milk passes harmlessly on each side of this elongated larynx. only to cling to the nipple of its mother. and Alexander Agassiz has seen an Echinus or sea-urchin rapidly passing particles of excrement from forceps to forceps down certain lines of its body. and Professor Flower has suggested to me that this structure would have greatly interfered with an animal swallowing solid food. These forceps can seize firmly hold of any object. unless the young at the same time were able to partake of the secretion. yet no minute. We will now turn for a short space to the lower divisions of the animal kingdom. and they would then have formed a breast. In such cases the most probable solution seems to be. The development of the mammary glands would have been of no service. whether in part through compensation of growth. that the glands over a certain space of the sack should have become more highly developed than the remainder. they subserve other functions. With respect to these organs. sea-urchins. of a tridactyle forceps--that is. that the habit was at first acquired by practice at a more advanced age. There is no greater difficulty in understanding how young mammals have instinctively learned to suck the breast. in order that its shell should not be fouled. when well developed. nor could the latter have been efficient without the snapping jaws. Through what agency the glands over a certain space became more highly specialised than the others. the young one must infallibly be choked by the intrusion of the milk into the wind-pipe. which consist. as we see in the Ornithorhyncus. would have been improved or rendered more effective. or how a few hours after leaving the shell they have learned to pick up grains of food.) are furnished with remarkable organs. etc. "this at least perfectly innocent and harmless structure?" It may be suggested in answer that the voice." Mr.of well-nourished offspring. asks: "What would be the utility of the FIRST RUDIMENTARY BEGINNINGS of such structures. than in understanding how unhatched chickens have learned to break the egg-shell by tapping against it with their specially adapted beaks. at the base of the mammalian series. but at first without a nipple. and is thus enabled to give free entrance to the air for the lungs. who has the power of injecting milk into the mouth of her helpless. moved by muscles. and thus the cutaneous glands. It accords with the widely extended principle of specialisation. Mivart remarks: "Did no special provision exist. But the young kangaroo is said not to suck. than would the individuals which secreted a poorer fluid. Mivart then asks how did natural selection remove in the adult kangaroo (and in most other mammals. which are the homologues of the mammary glands. But there IS a special provision. On this head Mr. as on so many previous occasions. I will not pretend to decide. and could not have been affected through natural selection. But there is no doubt that besides removing dirt of all kinds. which is certainly of high importance to many animals. "not even the SUDDEN development of the snapping action would have been beneficial without the freely movable stalk. of one formed of three serrated arms. Mr. and how could such insipient buddings have ever preserved the life of a single Echinus?" He adds. on the assumption that they are descended from a marsupial form). or of natural selection. half-formed offspring. The larynx is so elongated that it rises up into the posterior end of the nasal passage.

These differ much in structure in the different species. Perrier as bearing two kinds of pedicellariae. In certain genera of star-fishes these organs. This may be inferred from their manner of development in the individual. but capable of a snapping action. or zoophytes. Thus every gradation. to whose great kindness I am indebted for much information on the subject. to deny this seems to do no less than to affirm a startling paradox. is unmistakable. instead of being fixed or borne on an immovable support. seated on a neck and capable of movement. I wish I had space here to give a fuller abstract of Mr. there can be no reason to doubt that those furnished with serrated and movable branches likewise serve for the same purpose. one resembling those of Echinus. Muller. Certain compound animals. and if so. another group of the Echinodermata. may likewise be found between the pedicellariae of the star-fishes and the hooks of the Ophiurians. articulated to near their bases. with three equi-distant. though short. and such may be seen on the same spine together with the three lower branches. from an ordinary fixed spine to a fixed pedicellariae. in fact. Agassiz's interesting observations on the development of the pedicellariae. and again. in which one of the three arms of the forceps is reduced to a support for the other two. the shell is described by M. namely the Polyzoa. Agassiz. In certain genera of star-fishes. as well as from a long and perfect series of gradations in different species and genera. and in this case they probably subserve some additional function besides defence. serrated. three other movable branches. "the very combinations needed to show that the pedicellariae are only modified branching spines" may be found. that both in star-fishes and sea-urchins the pedicellariae must undoubtedly be looked at as modified spines. and this is intelligible if they serve. and such cases are always interesting as affording the means of apparently sudden transitions. as they have been termed. movable branches. Mr. The gradation extends even to the manner in which ordinary spines and the pedicellariae. are articulated to the shell. and they would thus serve still more effectively as soon as by meeting together they acted as a prehensile or snapping apparatus. to perfect tridactyle pedicellariae. and is thus rendered movable. In the sea-urchins the steps can be followed by which a fixed spine becomes articulated to the shell. In this case the identity in nature between the arms of the pedicellariae and the movable branches of a spine. It is generally admitted that the ordinary spines serve as a protection. All possible gradations. informs me that there are other star-fishes. and again between the pedicellariae of sea-urchins and the anchors of the Holothuriae." Paradoxical as this may appear to Mr. immovably fixed at the base. In their most perfect condition they curiously resemble the head and beak of a vulture in miniature. as a means of defence. and the other those of Spatangus. certainly exist on some star-fishes. as he adds. Thus we have fixed spines. through the abortion of one of the two states of an organ.variations could simultaneously evolve these complex co-ordinations of structure. at least in part. from simple granules to ordinary spines. and higher up. Mivart. also belonging to the same great class. other genera in which the third arm is completely lost. are provided with curious organs called avicularia. as is likewise the . Now when the latter arise from the summit of a spine they form. Mr. are placed on the summit of a flexible and muscular. with their supporting calcareous rods. With respect to the steps by which these curious organs have been evolved. on the same spine. tridactyle forcepses. Agassiz infers from his own researches and those of Mr. a rude tridactyle pedicellariae. In Echinoneus. would be of service. stem.

Busk. Besides the avicularia." having been developed through natural selection in widely distinct divisions of the animal kingdom. acting like long oars. "to sweep slowly and carefully over the surface of the polyzoary. they swept a branch rapidly across the object-glass of my microscope. But. Mivart might have adduced with equal appropriateness this resemblance as a special difficulty. it is believed. It is therefore impossible to conjecture by what serviceable gradations the one could have been converted into the other. Mivart adduces this case. through an angle of about 90 degrees. like the vibracula. Some species are provided with avicularia and vibracula. the movable lip or lid of the cell corresponding with the lower and movable mandible of the avicularium. removing what might be noxious to the delicate inhabitants of the cells when their tentacula are protruded. In one species observed by me. Smitt and Dr. and all of them on the same polyzoary often moved simultaneously. and their movement caused the whole polyzoary to tremble. and may be seen. it may be worth while to show that with the former a long series of serviceable gradations still exists. We next find one corner of the broad penultimate segment slightly prominent. but it by no means follows from this that such gradations have not existed. with its shape. Mr. which he considers as "essentially similar. Busk remarks. are afterwards swept by the currents within reach of the tentacula of the zooids. By an increase in the size of this projection. In the first and simplest stage. which. In one species examined by me the vibracula were slightly curved and serrated along the outer margin.lower jaw or mandible. as well as that of the terminal segment. The avicularia are believed by Mr. or even their resemblance to the head and beak of a bird. The latter resembles somewhat more closely the chelae or pincers of Crustaceans. . Busk. all the avicularia on the same branch often moved simultaneously backwards and forwards. but they also catch and kill small living animals. some with avicularia alone and a few with vibracula alone. As the chelae of Crustaceans resemble in some degree the avicularia of Polyzoa. and they made violent efforts to free themselves. These generally consist of long bristles. Dr. however. as Mr. until we have at last an instrument as efficient as the chelae of a lobster. namely the avicularia of the Polyzoa and the pedicellariae of the Echinodermata. the polyzoa possess curious organs called vibracula. but the limb still serves as an organ of locomotion. does not know of any gradations now existing between a zooid and an avicularium. both serving as pincers. so that. the pincers are rendered more and more perfect. slightly modified and improved. When a branch was placed on its face. When the jaws are touched with a needle they seize it so firmly that the branch can thus be shaken. as far as structure is concerned. Nitsche--naturalists who have carefully studied this group--to be homologous with the zooids and their cells which compose the zoophyte. probably serve for defence. capable of movement and easily excited. and against these the terminal segment shuts down. with the lower jaw widely open. and is thus enabled to catch hold of an object. and Mr. I can see no similarity between tridactyle pedicellariae and avicularia. And all these gradations can be actually traced. chiefly on account of the supposed difficulty of organs. sometimes furnished with irregular teeth. the vibracula became entangled. They are supposed to serve as a defence. the terminal segment of a limb shuts down either on the square summit of the broad penultimate segment. in the course of five seconds." The avicularia. Mr. or against one whole side.

but it seems more probable that they have passed through this stage.It is not easy to imagine two objects more widely different in appearance than a bristle or vibraculum. even when of considerable length and highly developed. This view of the development of the vibracula. the movable mandible is so much produced and is so like a bristle that the presence of the upper or fixed beak alone serves to determine its avicularian nature. The pollinia are by this means transported by insects from one flower to the stigma of another. Mivart only alludes to two cases. the little mass of viscid matter attached to the end of the caudicle. infinitesimal beginnings of structures which are of utility only when they are considerably developed. for supposing that all the species provided with avicularia had become extinct." As I have fully treated this subject in another work. as during the early stages of the transformation. in certain orchids similar viscid matter is secreted. we can understand how it is that these organs graduate in some cases. That this is the origin of the caudicle. but in much larger quantities by one alone of the three stigmas. with the avicularia of several species of Lepralia. there is no difficulty in believing that all the gradations. and an avicularium like the head of a bird. In the vegetable kingdom Mr. the other parts of the cell. namely the structure of the flowers of orchids. each of plain service to the plant. With respect to the former. but as these are not confined to orchids. namely. is interesting. I will here give only a few details on one alone of the most striking peculiarities of the flowers of orchids. Thus. With respect to the second chief peculiarity. Busk. In some orchids there is no caudicle to the pollen-masses. when highly developed. yet I may mention that at the base of the orchidaceous series. if trustworthy. affixed to an elastic foot-stalk or caudicle. no one with the most vivid imagination would ever have thought that the vibracula had originally existed as part of an organ. as I am informed by Mr. In other orchids the threads cohere at one end of the pollen-masses. and as the movable lip of the cell serves as a protection to the zooid. perhaps in consequence of the copious secretion. consists of a mass of pollen-grains. resembling a bird's head. we have good evidence in the aborted pollen-grains which can sometimes be detected embedded within the central and solid parts. and the movements of climbing plants. could hardly have disappeared at once. with the included zooid. by which the lip became converted first into the lower mandible of an avicularium. and the grains are merely tied together by fine threads. namely. they need not here be considered. A pollinium. though this support in some species is quite absent. he says: "The explanation of their ORIGIN is deemed thoroughly unsatisfactory--utterly insufficient to explain the incipient. namely a zooid with its cell. a long series of gradations can be specified. or an irregular box or hood. without having passed through the avicularian stage. and then into an elongated bristle. Hence. The vibracula may have been directly developed from the lips of the cells. we can see how the threads were probably first developed. and this forms the first or nascent trace of a caudicle. which seems to represent the fixed beak. and this to a little mass of extremely viscid matter. in Cypripedium. is rendered . their pollinia. In most flowers belonging to other orders the stigma secretes a little viscid matter. likewise served as a protection in different ways and under different circumstances. yet they are almost certainly homologous and have been developed from the same common source. It is interesting to see two such widely different organs developed from a common origin. Now. In many cases the vibracula have a grooved support at the base. and this stigma. into each other.

and. there are endless gradations--to species in which the pollen-mass terminates in a very short. single species and single genera possess the power of revolving. if the foot-stalks had possessed in any slight degree the requisite sensitiveness to a touch. The gradations from leaf-climbers to tendril bearers are wonderfully close. free caudicle--to others in which the caudicle becomes firmly attached to the viscid matter. secondly. and thus at the same time drags away some of the pollen-grains. on their continually bending to all points of the compass. as it is hopeless to attempt answering. firstly. with the sterile stigma itself much modified. are excited to bend round and clasp the touching object. For instance. by which means the foot-stalks of the leaves or flowers. In these two latter classes the stems have generally. I was led to predict that some slight tendency to a movement of this kind would be found to be far from uncommon with plants which did . which the tendrils likewise possess. it may naturally be asked how did plants acquire this power in an incipient degree. Hence. admit that all the many gradations in function and structure between simple twiners and tendril-bearers are in each case beneficial in a high degree to the species. I think. afterwards to be improved and increased through natural selection. In this. to a highly complex pollinium. and to those provided with tendrils. As soon as the lower part of a stem strikes against any object and is stopped. We will now turn to climbing plants. with the stigma differing but little from that of the ordinary flowers. namely sensitiveness to a touch. He who will carefully examine the flowers of orchids for himself will not deny the existence of the above series of gradations--from a mass of pollen-grains merely tied together by threads. but as we do not know the full history of any one group of beings. lost the power of twining. These can be arranged in a long series. As in many widely separated families of plants.sterile. He who will read my memoir on these plants will. From this simple condition. or these modified and converted into tendrils. in the same order. it is clearly a great advantage to a twining plant to become a leaf-climber. and forms the basis of our series. and thus necessarily twines round and up the support. and in almost every other case. which differs but little from that of a multitude of common flowers. The power of twining depends. admirably adapted for transportal by insects. such questions. but not always. When an insect visits a flower of this kind. and certain plants may be differently placed in either class. and have thus become twiners. it is as useless to ask. As twining is the simplest means of ascending a support. But in ascending the series from simple twiners to leaf-climbers. In this latter case we have a pollinium in its most highly developed and perfect condition. it rubs off some of the viscid matter. they must have independently acquired it. The revolving movement ceases after the early growth of each shoot. the enquiry may be pushed further backwards. the upper part still goes on bending and revolving. to those which I have called leaf-climbers. and it is probable that every twiner which possessed leaves with long foot-stalks would have been developed into a leaf-climber. an important quality is added. nor will he deny that all the gradations in the several species are admirably adapted in relation to the general structure of each flower for its fertilisation by different insects. on the stems while young being extremely flexible (but this is a character common to many plants which are not climbers). from those which simply twine round a support. and are made to move round and round. though they retain the power of revolving. and cannot have inherited it from a common progenitor. By this movement the stems are inclined to all sides. one after the other in succession. and it may be asked how did the stigma of an ordinary flower become viscid.

As a vast number of species. this movement. Soon afterwards Fritz Muller discovered that the young stems of an Alisma and of a Linum--plants which do not climb and are widely separated in the natural system--revolved plainly. in the young and growing organs of plants. only during the early stages of growth that the foot-stalks and tendrils are sensitive. So with plants it appears that. in others it was extremely slight. which are of manifest importance to them. due to a touch or shake. When I made this prediction. it is this tendency which has been taken advantage of and increased through natural selection. being the incidental result of . they are not of the least use in the way of climbing. until they had become converted into well-developed twining species. Morren found in several species of Oxalis that the leaves and their foot-stalks moved. of the young flower-peduncles of a Maurandia which revolved slightly and irregularly. like the stems of twining plants. but without making any use of this habit. I repeated these observations on some other species of Oxalis with the same result. nearly the same remarks are applicable as in the case of the revolving movements of twining plants. which were at first of no use to them. which is the point that concerns us. But plants possess. can be of any functional importance to them. in obedience to various stimuli. from reasons which I have assigned in my memoir. I knew of only one imperfect case. towards and more rarely from the light--in opposition to. however. It is a more important fact that according to the high authority of Hofmeister. and with climbing plants it is. as well as that due to a touch or shake. and that this had afforded the basis for natural selection to work on and improve. Hence there is no great difficulty in admitting that in the case of leaf-climbers and tendril-bearers. that this will have occurred only with plants which had already acquired the power of revolving. for the nerves and muscles have not been rendered specially sensitive to these stimuli. I have already endeavoured to explain how plants became twiners. It is scarcely possible that the above slight movements. probable. for instance. the attraction of gravity. and if under the conditions to which they are exposed it had profited them to ascend to a height. When the nerves and muscles of an animal are excited by galvanism or by the absorption of strychnine. it ought to be found in a nascent condition in many plants which have not become climbers. they are excited in an incidental manner by a touch. These slight movements appear to be of no service to the plants in question. namely.not climb. It is. though irregularly. or by being shaken. the consequent movements may be called an incidental result. especially after exposure to a hot sun. This is the case: I observed that the young flower-peduncles of the above Maurandia curved themselves a little towards the side which was touched. by the increase of a tendency to slight and irregular revolving movements. namely. the young shoots and leaves of all plants move after being shaken. and more rarely in the direction of. but was best seen in the young leaves. Nevertheless we can see that if the stems of these plants had been flexible. then the habit of slightly and irregularly revolving might have been increased and utilised through natural selection. and had thus become twiners. and he states that he has reason to suspect that this occurs with some other plants. are endowed with this kind of sensitiveness. and of tendrils. in some of them the movement was distinct. from having the power of movement in obedience to certain stimuli. belonging to widely distinct groups. or when the plant was shaken. powers of movement. With respect to the sensitiveness of the foot-stalks of the leaves and flowers. when they were gently and repeatedly touched. anyhow. as we know.

as I hope. which had the longest necks. the continued preservation of the individuals of some extinct high-reaching ruminant. and that these glands were improved in function through natural selection. With the many insects which imitate various objects. in which case they would have formed a mamma. A good opportunity has thus been afforded for enlarging a little on gradations of structure. With such structures as the above lamellae of horn or whalebone. legs. In the family of the ducks. together with inheritance. until the points were converted. will have aided in an important manner in their co-ordination. In certain species of whales there is a tendency to the formation of irregular little points of horn on the palate. which served as a defence. Whether. often associated with strange functions--an important subject. together with inheritance. There is no more difficulty in understanding how the branched spines of some ancient Echinoderm. as in the mouth of the Greenland whale. became developed through natural selection into tridactyle pedicellariae. and I have shown. but we know that certain periodical movements. perhaps more than enough. to prove that natural selection is incompetent to account for the incipient stages of useful structures. may be attributed almost wholly to continued use. the lamellae are first used as teeth. I have now considered enough. which was at first used solely for locomotion.the power of moving. selected with care by a skilful naturalist. With respect to the mammae of the higher animals. are governed by habit. the transportal of the lower eye of a flat-fish to the upper side of the head. for instance the so-called sleep of plants. the most probable conjecture is that primordially the cutaneous glands over the whole surface of a marsupial sack secreted a nutritious fluid. since perfected through the occasional preservation of slight variations which made the resemblance at all closer. and at last almost exclusively for this latter purpose. like those on the beak of a goose--then into short lamellae. gained for other and beneficial purposes. as perfect as those of the shoveller-duck--and finally into the gigantic plates of baleen. etc. than in understanding the development of the pincers of crustaceans. that there is no great difficulty on this head.. habit or use can have done little or nothing. then partly as teeth and partly as a sifting apparatus. and it seems to be quite within the scope of natural selection to preserve all favourable variations. With the giraffe. there is no improbability in the belief that an accidental resemblance to some common object was in each case the foundation for the work of natural selection. but the prolonged use of all the parts. and the continued destruction of those which could not browse so high. On the other hand. which was not treated at sufficient length in the former editions of this work. as far as we can judge. through slight. I will not pretend to decide. of the cases. and concentrated into a confined area. serviceable modifications in the ultimate and penultimate segments of a limb. would have sufficed for the production of this remarkable quadruped. and the formation of a prehensile tail. first into lamellated knobs or teeth. In the avicularia and vibracula of the Polyzoa we have organs widely different in appearance developed from the same source. towards their development. and this will have been carried on as long as the insect continued to vary. during the gradual development of climbing plants. natural selection has been aided by the inherited effects of use. like those of the domestic ducks--and then into lamellae. . I will now briefly recapitulate the foregoing cases. and as long as a more and more perfect resemblance led to its escape from sharp-sighted enemies. and could browse a little above the average height.

and the steps can likewise be followed by which viscid matter. why has not this or that structure been gained by certain species. would have been beneficial to a species. He thinks it difficult to believe that the wing of a bird "was . but in some few cases a retrogression." about which it is not pretended that anything is known. and the requisite conditions may seldom have concurred. and it may often have happened that the requisite parts did not vary in the right manner or to the right degree. he supposes that the differences between the extinct three-toed Hipparion and the horse arose suddenly. They seem to me to partake little of the character of demonstration. can be traced cohering into caudicles. and which. aided by the other agencies often specified. if natural selection be so potent. such as that secreted by the stigmas of ordinary flowers. which we think. as it seems to me. His chief arguments have now been considered. Mr. and some naturalists agree with him." At the present day almost all naturalists admit evolution under some form. With respect to climbing plants. to invoke any internal force beyond the tendency to ordinary variability. would equally well give rise by graduated steps to natural races or species. but in some few cases special reasons. can be assigned. but he considers it as "demonstrably insufficient" to account for the phenomena which I explain by its agency. The belief that any given structure. often erroneously. Mr. considering our ignorance of the past history of each species. I am bound to add. is opposed to what we can understand of its manner of action. would have been gained under all circumstances through natural selection. by man has given rise to many well-adapted domestic races. The final result will generally have been. and the others will hereafter be considered. Mivart is further inclined to believe. the threads which originally served to tie together the pollen-grains. that new species manifest themselves "with suddenness and by modifications appearing at once. as the struggle for life did not depend on such structures. but there is no need. and of the conditions which at the present day determine its numbers and range. through the aid of natural selection." For instance. That species have a capacity for change will be admitted by all evolutionists. in organisation. often of a peculiar nature. In this case. It has often been asked. which we imagine would have been gained through natural selection from appearing to us advantageous to the species. Mivart believes that species change through "an internal force or tendency. and to have little weight in comparison with those in favour of the power of natural selection. Mivart does not deny that natural selection has effected something. which stood in no relation to certain structures. that some of the facts and arguments here used by me. many co-ordinated modifications are almost indispensable. as already explained. and still subserving nearly but not quite the same purpose. In many cases complex and long-enduring conditions. With the pollinia of orchids. an advance. Thus to adapt a species to new habits of life. Many species must have been prevented from increasing in numbers through destructive agencies. Mr. are necessary for the development of a structure. I need not repeat what has been so lately said. they could not have been acquired through natural selection. have been advanced for the same purpose in an able article lately published in the "Medico-Chirurgical Review. In most cases only general reasons. which through the aid of selection. became attached to the free ends of the caudicles--all these gradations being of manifest benefit to the plants in question. to which it would apparently have been advantageous? But it is unreasonable to expect a precise answer to such questions.and with the vibracula we can understand how the successive gradations might have been of service.

etc. Hence. That many species have been evolved in an extremely gradual manner. which implies great breaks or discontinuity in the series. Everyone who believes in slow and gradual evolution. to be lost by accidental causes of destruction and by subsequent intercrossing. Nor should it be forgotten. are as follows. So it is if we look to past times. doubtful species. etc. It is indeed manifest that multitudes of species are related in the closest manner to other species that still exist. and the characters which thus reappear were. The species and even the genera of many large natural families are so closely allied together that it is difficult to distinguish not a few of them. it is probable. there can hardly be a doubt. My reasons for doubting whether natural species have changed as abruptly as have occasionally domestic races. or have lately existed. I am compelled to allude to subjects hereafter to be discussed. or if we compare the fossil species embedded in the sub-stages of the same geological formation. in proceeding from north to south. Mivart. A still greater number must be called monstrosities. This conclusion. they would be liable. if found in a state of nature. Look at the many outlying islands round a continent. abrupt and strongly marked variations occur in our domesticated productions. as are known occasionally to arise under domestication. and compare the species which have just passed away with those still living within the same areas. such as six-fingered men. Mivart.developed in any other way than by a comparatively sudden modification of a marked and important kind. This difficulty. On every continent. will of course admit that specific changes may have been as abrupt and as great as any single variation which we meet with under nature.." and apparently he would extend the same view to the wings of bats and pterodactyles. Of these latter variations several may be attributed to reversion. But as species are more variable when domesticated or cultivated than under their natural conditions. as formerly explained. and for entirely disbelieving that they have changed in the wonderful manner indicated by Mr. If such occurred under nature. which we have reason to believe were formerly connected. and it will hardly be maintained that such species have been developed in an abrupt or sudden manner. as in the case of unconscious selection by man. appears to me improbable in the highest degree. in many cases at first gained in a gradual manner. Niata cattle. and see how many of their inhabitants can be raised only to the rank of doubtful species. porcupine men. According to our experience. it is almost necessary to believe. closely related to their parental types. and as they are widely different in character from natural species. singly and at rather long intervals of time. in opposition to all analogy. But in making these and the following remarks. and so it is known to be under domestication. in order that a new species should suddenly appear in the manner supposed by Mr. or even under domestication. is avoided on the theory of gradual evolution. and of the destruction of a large number which varied in an opposite manner. . Excluding such cases of abrupt variations. unless abrupt variations of this kind are specially preserved and separated by the care of man.. from lowland to upland. as we likewise do on certain distinct continents. that several wonderfully changed individuals appeared simultaneously within the same district. we meet with a host of closely related or representative species. the few which remain would at best constitute. which varied more or less in any favourable direction. they throw very little light on our subject. it is not probable that such great and abrupt variations have often occurred under nature. Ancon sheep. through the preservation of a large number of individuals.

embryology enters a strong protest. and yet should not bear even a trace in its embryonic condition of any sudden modification. for instance. it is incredible that an animal should have undergone such momentous and abrupt transformations as those above indicated.when we look to the special parts of allied species. On this same principle we can understand how it is that specific characters are more variable than generic characters. or the sudden conversion of a Hipparion into a horse. But against the belief in such abrupt changes. however. one furnished with wings. If the record is as fragmentary as many geologists strenuously assert. hardly any light is thrown by the belief in abrupt modifications on the deficiency of connecting links in our geological formations. On this view of the meaning of embryological resemblances. and serves as a record of the past condition of the species. and how the parts which are developed in an extraordinary degree or manner are more variable than other parts of the same species. and that they become differentiated by insensibly fine steps. yet it may be maintained that some have been developed in a different and abrupt manner. which have been advanced in favour of this view. But the value of this evidence depends entirely on the perfection of the geological record. every detail in its structure being developed by insensibly fine steps. Although very many species have almost certainly been produced by steps not greater than those separating fine varieties. For instance. Embryological resemblances of all kinds can be accounted for. by the progenitors of our existing species having varied after early youth. in opposition to all analogy. The vague and in some respects false analogies. the fact that the species included in the larger genera are more closely related to each other. and having transmitted their newly-acquired characters to their offspring. or the falling of a facetted spheroid from one facet to another. as they have been shown to be by Mr. ought not to be made without strong evidence being assigned. however. such as the sudden crystallisation of inorganic substances. like varieties round species. It is notorious that the wings of birds and bats. . Such an admission. will be almost compelled to assume. hardly deserve consideration. and the legs of horses or other quadrupeds. Chauncey Wright. connecting together widely different structures. Hence it is that existing species during the early stages of their development so often resemble ancient and extinct forms belonging to the same class. namely. Mivart. that numerous and wonderfully fine gradations can be traced. at a corresponding age. there is nothing strange in new forms appearing as if suddenly developed. are undistinguishable at an early embryonic period. and indeed on any view. One class of facts. The former are also grouped in little clusters. in relation to periods remote in the history of the world. The embryo is thus left almost unaffected. as was shown in our second chapter. all pointing in the same direction. instead of to distinct species. such as the sudden development of the wings of birds or bats. Many large groups of facts are intelligible only on the principle that species have been evolved by very small steps. and they present other analogies with varieties. He who believes that some ancient form was transformed suddenly through an internal force or tendency into. could be added. as we shall hereafter see. Unless we admit transformations as prodigious as those advocated by Mr. the sudden appearance of new and distinct forms of life in our geological formations supports at first sight the belief in abrupt development. Many analogous facts. and present a greater number of varieties than do the species in the smaller genera.

more especially by a very young one. have been suddenly produced. he is generally forced to go back to recover the habitual train of thought: so P. Huber found it was with a caterpillar. and when performed by many individuals in the same way. He will further be compelled to believe that many structures beautifully adapted to all the other parts of the same creature and to the surrounding conditions. when performed by an animal. INSTINCT. and to leave those of science. Many instincts are so wonderful that their development will probably appear to the reader a difficulty sufficient to overthrow my whole theory. molothrus. they often remain constant throughout life. Habits easily become associated with other habits. ostrich. their origin--Natural instincts of the cuckoo. he will not be able to assign a shadow of an explanation. as Pierre Huber expresses it. It would be easy to show that several distinct mental actions are commonly embraced by this term. without experience. that I have nothing to do with the origin of the mental powers. Frederick Cuvier and several of the older metaphysicians have compared instinct with habit. any more than I have with that of life itself. but different in their origin--Instincts graduated--Aphides and ants--Instincts variable--Domestic instincts. but not necessarily of its origin. to enter into the realms of miracle. A little dose of judgment or reason. To admit all this is. as it seems to me. when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds' nests. or in repeating anything by rote. An action. This comparison gives. so in instincts. an accurate notion of the frame of mind under which an instinctive action is performed. It cannot be denied that such abrupt and great changes of structure are widely different from those which most species apparently have undergone. often comes into play. I may here premise. but every one understands what is meant. and of such complex and wonderful co-adaptations. indeed not rarely in direct opposition to our conscious will! yet they may be modified by the will or reason. I will not attempt any definition of instinct. He will be forced to admit that these great and sudden transformations have left no trace of their action on the embryo.that many individuals varied simultaneously. We are concerned only with the diversities of instinct and of the other mental faculties in animals of the same class. which makes a very complicated . without their knowing for what purpose it is performed. Several other points of resemblance between instincts and habits could be pointed out. When once acquired. with certain periods of time and states of the body. As in repeating a well-known song. But I could show that none of these characters are universal. I think. Instincts comparable with habits. one action follows another by a sort of rhythm. and parasitic bees--Slave-making ants--Hive-bee. is usually said to be instinctive. which we ourselves require experience to enable us to perform. its cell-making instinct--Changes of instinct and structure not necessarily simultaneous--Difficulties of the theory of the Natural Selection of instincts--Neuter or sterile insects--Summary. How unconsciously many habitual actions are performed. CHAPTER VIII. even with animals low in the scale of nature. if a person be interrupted in a song.

to the third stage. we ought to find in nature. where it had left off. be might truly be said to have done so instinctively. it was much embarrassed. a caterpillar were taken out of a hammock made up. and thus tried to complete the already finished work. And such instances of diversity of instinct in the same species can be . Changes of instinct may sometimes be facilitated by the same species having different instincts at different periods of life. and if it can be shown that instincts do vary ever so little.--that is of variations produced by the same unknown causes which produce slight deviations of bodily structure. and put it into a hammock completed up only to the third stage. But I believe that the effects of habit are in many cases of subordinate importance to the effects of the natural selection of what may be called spontaneous variations of instincts. It is thus. in which case either the one or the other instinct might be preserved by natural selection. or when placed under different circumstances. and this we certainly can do. making allowance for the instincts of animals having been but little observed. I have been surprised to find. under its present conditions of life. that all the most complex and wonderful instincts have originated. and sixth stages of construction.hammock. It will be universally admitted that instincts are as important as corporeal structures for the welfare of each species. variations. and then transmitted by inheritance to succeeding generations. and are increased by. so I do not doubt it has been with instincts. can be discovered. slight. for instance. seemed forced to start from the third stage. far from deriving any benefit from this. But it would be a serious error to suppose that the greater number of instincts have been acquired by habit in one generation. No complex instinct can possibly be produced through natural selection. so that much of its work was already done for it. except in Europe and North America. use or habit. As modifications of corporeal structure arise from. If we suppose any habitual action to become inherited--and it can be shown that this does sometimes happen--then the resemblance between what originally was a habit and an instinct becomes so close as not to be distinguished. and. as in the case of corporeal structures. the sixth stage of construction. and were put into one finished up to the sixth stage. or we ought at least to be able to show that gradations of some kind are possible. it is at least possible that slight modifications of instinct might be profitable to a species. fifth. and for no instinct being known among extinct species. for if he took a caterpillar which had completed its hammock up to. however. those of the hive-bee and of many ants. say. in order to complete its hammock. namely. could not possibly have been acquired by habit. yet profitable. except by the slow and gradual accumulation of numerous. If. then I can see no difficulty in natural selection preserving and continually accumulating variations of instinct to any extent that was profitable. or at different seasons of the year. Hence. and are diminished or lost by disuse. leading to the most complex instincts. had played a tune with no practice at all. It can be clearly shown that the most wonderful instincts with which we are acquainted. Under changed conditions of life. etc.. as I believe. instead of playing the pianoforte at three years old with wonderfully little practice. the caterpillar simply re-performed the fourth. how very generally gradations. If Mozart. not the actual transitional gradations by which each complex instinct has been acquired--for these could be found only in the lineal ancestors of each species--but we ought to find in the collateral lines of descent some evidence of such gradations.

Although there is no evidence that any animal performs an action for the exclusive good of another species. but often from causes wholly unknown to us. As some degree of variation in instincts under a state of nature. but not one excreted. and prevented their attendance during several hours. as well as I could. as I have elsewhere shown. and in its total loss. has it not granted to the bee "the ability to use some other material when wax was deficient?" But what other natural material could bees use? They will work. It has lately been shown that bees. and each. as I have seen. the migratory instinct. and not the result of experience. therefore probably they do not excrete solely for the good of the ants. The fear of man is slowly acquired. though it is strengthened by experience. I allowed an ant to visit them. namely. and by the sight of fear of the same enemy in other animals. immediately lifted up its abdomen and excreted a limpid drop of sweet juice. After this interval. But as the excretion is extremely viscid. showing that the action was instinctive. are indispensable for the action of natural selection. it has been asked. with which he had covered decorticated trees. as far as we can judge. but not one excreted. both in extent and direction. from the observations of Huber. I removed all the ants from a group of about a dozen aphides on a dock-plant. and we see an instance of this. by its eager way of running about to be well aware what a rich flock it had discovered. is that of aphides voluntarily yielding. as the ants do with their antennae. which was eagerly devoured by the ant.shown to occur in nature. Why. instead of searching for pollen. their sweet excretion to ants: that they do so voluntarily. Audubon has given several remarkable cases of differences in the nests of the same species in the northern and southern United States. which vary partly in dependence on the situations chosen. . but want of space prevents me. but as details on this and other such points are not indispensable. by the various animals which inhabit desert islands. Fear of any particular enemy is certainly an instinctive quality. as may be seen in nestling birds. been produced for the exclusive good of others. as each takes advantage of the weaker bodily structure of other species. with wax hardened with vermilion or softened with lard. the instinct of each species is good for itself. Afterwards. even in England. One of the strongest instances of an animal apparently performing an action for the sole good of another. as in the case of corporeal structure. that the aphides show no dislike to the ants: if the latter be not present they are at last compelled to eject their excretion. Even the quite young aphides behaved in this manner. oatmeal. I felt sure that the aphides would want to excrete. So it is with the nests of birds. So again certain instincts cannot be considered as absolutely perfect. as was first observed by Huber. will gladly use a very different substance. in the greater wildness of all our large birds in comparison with our small birds. with which I am acquainted. It is certain. yet each tries to take advantage of the instincts of others. and on the nature and temperature of the country inhabited. instead of laboriously collecting propolis. I can only assert that instincts certainly do vary--for instance. it then began to play with its antennae on the abdomen first of one aphis and then of another. and conformably to my theory. but has never. it is no doubt a convenience to the aphides to have it removed. as soon as it felt the antennae. if instinct be variable. the following facts show. and it immediately seemed. they may be here passed over. I then tickled and stroked them with a hair in the same manner. I watched them for some time through a lens. and the inheritance of such variations. Again. as many instances as possible ought to be given. used a cement of wax and turpentine. Andrew Knight observed that his bees.

always brought home game birds. and how curiously they become mingled. It is notorious how much domestic animals vary in their mental qualities. stand motionless like a statue. and a cross with a greyhound has given to a whole family of shepherd-dogs a tendency to hunt hares. We may safely attribute the greater wildness of our large birds to this cause. that I do not speak without good evidence. is tame in Norway. A number of curious and authentic instances could be given of various shades of disposition and taste. for in uninhabited islands large birds are not more fearful than small. a herd of deer. which in a like manner become curiously blended together. St. associated with certain frames of mind or periods of time. performed with eager delight by each breed. as they may be called. born in a state of nature. a flock of sheep. instead of at. retrieving is certainly in some degree inherited by retrievers. by shepherd-dogs. could be shown by many facts. That the mental qualities of animals of the same kind. which. One cat. The possibility. or even probability. habits. and likewise of the oddest tricks. as soon as it scented its prey. but they have been acted on by far less rigorous selection. than the white butterfly knows why she lays her eggs on the leaf of the cabbage--I cannot see that these actions differ essentially from true instincts. I can only repeat my assurance. How strongly these domestic instincts. of inherited variations of instinct in a state of nature will be strengthened by briefly considering a few cases under domestication. and these tendencies are known to be inherited. INHERITED CHANGES OF HABIT OR INSTINCT IN DOMESTICATED ANIMALS. performed without experience by the young. we should assuredly call these actions instinctive. when thus tested by crossing. instead of at. and have been transmitted for an incomparably shorter period. and another hunted on marshy ground and almost nightly caught woodcocks or snipes. Several cases could also be adduced of occasional and strange habits in wild animals. John. resemble natural instincts. and then slowly crawl forward with a peculiar gait. under less fixed conditions of life. as is the hooded crow in Egypt. We shall thus be enabled to see the part which habit and the selection of so-called spontaneous variations have played in modifying the mental qualities of our domestic animals. and another mice. another hares or rabbits. and dispositions are inherited. I cannot see that these actions. without the facts in detail. to new instincts. But I am well aware that these general statements. one naturally takes to catching rats. can produce but a feeble effect on the reader's mind. and the magpie.for the large birds have been most persecuted by man. and without the end being known--for the young pointer can no more know that he points to aid his master. But let us look to the familiar case of the breeds of dogs: it cannot be doubted that young pointers (I have myself seen striking instances) will sometimes point and even back other dogs the very first time that they are taken out. These domestic instincts. and another kind of wolf rushing round. and a tendency to run round. are certainly far less fixed than natural instincts. and driving them to a distant point. so wary in England. according to Mr. With cats. if advantageous to the species. If we were to behold one kind of wolf. and for a long period . for instance. and in nearly the same manner by each individual. Domestic instincts. through natural selection. vary much. when young and without any training. is well shown when different breeds of dogs are crossed. Thus it is known that a cross with a bull-dog has affected for many generations the courage and obstinacy of greyhounds. might have given rise.

All wolves. the Gallus bankiva. so that habit and some degree of selection have probably concurred in civilising by inheritance our dogs. as I once saw." that is. It is scarcely possible to doubt that the love of man has become instinctive in the dog. where the savages do not keep these domestic animals. are at first excessively wild. as I have witnessed. and this is known occasionally to happen. by not coming in a straight line to his master. but this is not true. for if the hen gives the danger chuckle they will run (more especially young turkeys) from under her and conceal themselves in the surrounding grass or thickets. sheep and pigs. How rarely. and unconscious selection is still in progress. On the other hand. without intending to improve the breed. as we see in wild ground-birds. jackals and species of the cat genus. they are destroyed. sheep. only the exaggerated pause of an animal preparing to spring on its prey. the tumbler-pigeon to tumble--an action which. even when quite young. but fear only of dogs and cats.exhibit traces of the instincts of either parent: for example. When the first tendency to point was once displayed. So it is with young pheasants reared in England under a hen. so that we must attribute at least the greater part of the inherited change from extreme wildness to extreme tameness. for the mother-hen has almost lost by disuse the power of . dogs which stand and hunt best. Familiarity alone prevents our seeing how largely and how permanently the minds of our domestic animals have been modified. when reared in India under a hen. had not some one dog naturally shown a tendency in this line. scarcely any animal is tamer than the young of the tame rabbit. to habit and long-continued close confinement. methodical selection and the inherited effects of compulsory training in each successive generation would soon complete the work. hardly any animal is more difficult to tame than the young of the wild rabbit. and that the long-continued selection of the best individuals in successive generations made tumblers what they now are. do our civilised dogs. and this tendency has been found incurable in dogs which have been brought home as puppies from countries such as Tierra del Fuego and Australia. as many have thought. whose great-grandfather was a wolf. is performed by young birds. and near Glasgow there are house-tumblers. or probably could have taught. but I can hardly suppose that domestic rabbits have often been selected for tameness alone. when kept tame. on the other hand. Brent. Domestic instincts are sometimes spoken of as actions which have become inherited solely from long-continued and compulsory habit. foxes. in a pure terrier: the act of pointing is probably. and if not cured. We may believe that some one pigeon showed a slight tendency to this strange habit. which cannot fly eighteen inches high without going head over heels. and are then beaten. their mother to fly away. It may be doubted whether any one would have thought of training a dog to point. as each man tries to procure. It is not that chickens have lost all fear. as I hear from Mr. Natural instincts are lost under domestication: a remarkable instance of this is seen in those breeds of fowls which very rarely or never become "broody. never wish to sit on their eggs. But this instinct retained by our chickens has become useless under domestication. and pigs! No doubt they occasionally do make an attack. for I am informed by Captain Hutton that the young chickens of the parent stock. Le Roy describes a dog. No one would ever have thought of teaching. habit alone in some cases has sufficed. On the other hand. and this dog showed a trace of its wild parentage only in one way. young chickens have lost wholly by habit. and this is evidently done for the instinctive purpose of allowing. are most eager to attack poultry. that have never seen a pigeon tumble. require to be taught not to attack poultry. when called. that fear of the dog and cat which no doubt was originally instinctive in them.

but in most cases habit and selection have probably concurred. that the cuckoo occasionally lays her eggs on the bare ground. the instinct which leads the cuckoo to lay her eggs in other birds' nests. that he once found in Illinois a young cuckoo. not daily. those first laid would have to be left for some time unincubated or there would be eggs and young birds of different ages in the same nest. all at the same time. partly by habit and partly by man selecting and accumulating. we may conclude that under domestication instincts have been acquired and natural instincts have been lost. It has been both asserted and denied that the American cuckoo occasionally lays her eggs in other birds' nests. encumbered as she could hardly fail to be by having eggs and young of different ages at the same time. which at first appeared from what we must in our ignorance call an accident. SPECIAL INSTINCTS. in other cases compulsory habit has done nothing. This rare event is probably a case of reversion to the long-lost. and thus be more successful in rearing their young. also recently been ascertained on sufficient evidence. or if the young were made more vigorous by advantage being taken of the mistaken instinct of another species than when reared by their own mother. I will select only three. We shall. and as both were nearly full feathered. It is supposed by some naturalists that the more immediate cause of the instinct of the cuckoo is that she lays her eggs. and in their turn would be apt to lay their eggs in other birds' nests. Merrill. Hence. By a continued process of this nature. by Adolf Muller.flight. of Iowa. peculiar mental habits and actions. It has. and that she occasionally laid an egg in another bird's nest. during successive generations. aboriginal instinct of . more especially as she migrates at a very early period. if she were to make her own nest and sit on her own eggs. then the old birds or the fostered young would gain an advantage. namely. and all has been the result of selection. so that. If the old bird profited by this occasional habit through being enabled to emigrate earlier or through any other cause. I could also give several instances of various birds which have been known occasionally to lay their eggs in other birds' nests. and the cell-making power of the hive-bee: these two latter instincts have generally and justly been ranked by naturalists as the most wonderful of all known instincts. Now let us suppose that the ancient progenitor of our European cuckoo had the habits of the American cuckoo. But the American cuckoo is in this predicament. INSTINCTS OF THE CUCKOO. but I have lately heard from Dr. but at intervals of two or three days. there could be no mistake in their identification. together with a young jay in the nest of a blue jay (Garrulus cristatus). perhaps. I believe that the strange instinct of our cuckoo has been generated. and the first hatched young would probably have to be fed by the male alone. In some cases compulsory habit alone has sufficed to produce inherited mental changes. And analogy would lead us to believe that the young thus reared would be apt to follow by inheritance the occasional and aberrant habit of their mother. sits on them and feeds her young. for she makes her own nest and has eggs and young successively hatched. best understand how instincts in a state of nature have become modified by selection by considering a few cases. the slave-making instinct of certain ants. If this were the case the process of laying and hatching might be inconveniently long. pursued both methodically and unconsciously.

In the case of the European cuckoo. speculation on an instinct known to us only in a single species. the . One of these was replaced in the nest by the observer. as is probably the case. But he has now received a trustworthy account of a young cuckoo which was actually seen. lays only one egg in a nest. to an extraordinary degree in colour. not exceeding those of the skylark--a bird about one-fourth as large as the cuckoo. with rare exceptions. The eggs of the Australian bronze cuckoo vary. but not rarely departs from it. the strength and a properly shaped back for ejecting its foster-brothers. Thirdly. and as the latter at this age is in a most helpless condition. that the common cuckoo. for we have hitherto had no facts to guide us. Gould was formerly inclined to believe that the act of ejection was performed by the foster-parents themselves. if it were of great importance for the young cuckoo. The chief points to be referred to are three: first.nidification. This has been boldly called a beneficent arrangement. which lay their eggs in other birds' nests. that the eggs are remarkably small. natural selection might have secured and fixed any advantageous variation. now. while still blind and not able even to hold up its own head. But in all cases. or. The European species apparently manifests some tendency towards a similar instinct. then there is no difficulty in believing that a race or species might have been formed which would have laid smaller and smaller eggs. Now. during successive generations. the offspring of the foster-parents are commonly ejected from the nest within three days after the cuckoo is hatched. so that in this respect. Secondly. it would assuredly have been added to those which it is assumed must all have been acquired together. and that its foster-brothers may perish before they had acquired much feeling! Turning now to the Australian species: though these birds generally lay only one egg in a nest. That the small size of the egg is a real case of adaptation we may infer from the fact of the mon-parasitic American cuckoo laying full-sized eggs. when they lay their eggs in an open nest. in the act of ejecting its foster-brothers. if it had been of an advantage to this species to have laid eggs even smaller than those now laid. owing to Mr. for these would have been more safely hatched and reared. In the bronze cuckoo the eggs vary greatly in size. that the young cuckoo. to receive as much food as possible soon after birth. Ramsay remarks that two of the Australian cuckoos. in order that the young cuckoo may get sufficient food. Mr. according to Mr. from eight to ten lines in length. we have learned something about three Australian species. is useless. which then perish from cold and hunger. has the instinct. It has been objected that I have not noticed other related instincts and adaptations of structure in the cuckoo. as well as in size. to have been hatched within a shorter period (for it is asserted that there is a relation between the size of eggs and the period of their incubation). Mr. manifest a decided preference for nests containing eggs similar in colour to their own. which are spoken of as necessarily co-ordinated. Until recently the instincts of the European and of the non-parasitic American cuckoo alone were known. and was again thrown out. Ramsay. as is shown by her laying her dull and pale-coloured eggs in the nest of the hedge-warbler with bright greenish-blue eggs. I can see no special difficulty in its having gradually acquired. as is more probable. With respect to the means by which this strange and odious instinct was acquired. so as to have deceived certain foster-parents. Ramsay's observations. it is not rare to find two and even three eggs in the same nest. Had our cuckoo invariably displayed the above instinct. soon after birth. so that the large and voracious young bird receives ample food.

Hudson is a strong disbeliever in evolution. the M. complete a nest for themselves. This instinct may probably be accounted for by the fact of the hens laying a large number of eggs. a widely distinct genus of American birds. and sometimes to pair. as in the case of the Molothrus bonariensis. are much more highly developed than those of the last. whether of their own species or of their foster parents. The first step towards the acquisition of the proper instinct might have been mere unintentional restlessness on the part of the young bird. which are thus wasted. sometimes to live promiscuously together in flocks. The parasitic habits of another species of Molothrus. have parasitic habits like those of the cuckoo. The instinct. a transitory sharp tooth for cutting through the tough egg-shell. the habit having been afterwards improved. as Owen has remarked. Mr. as on the leaves of a large thistle. as has already been remarked. but as small consequences of one general law. and the variations tend to be inherited at a corresponding or earlier age--propositions which cannot be disputed--then the instincts and structure of the young could be slowly modified as surely as those of the adult. the M. so that the young bird is securely reared. "Must we consider these habits. and these are hatched by the males. They have. occasionally lay their eggs in the nests of other birds. however. The sexes of Molothrus badius are stated by an excellent observer. Hudson. which they find in the appropriated nests. They never. but he appears to have been so much struck by the imperfect instincts of the Molothrus bonariensis that he quotes my words. and both cases must stand or fall together with the whole theory of natural selection. . Mr. but it is remarkable that several together sometimes commence to build an irregular untidy nest of their own. placed in singular ill-adapted situations. the extraordinary habit of pecking holes in the eggs. and structure necessary for the work of ejection. for it never lays more than one egg in a foster-nest. and transmitted to an earlier age. for he has seen the young of this species following old birds of a distinct kind and clamouring to be fed by them. transition?" Various birds. as with the cuckoo. Hudson says it is probable that they are occasionally parasitic. however. They either lay their eggs in the nest thus appropriated. but. has acquired instincts as perfect as those of the cuckoo. They either build a nest of their own or seize on one belonging to some other bird. This habit is not very uncommon with the Gallinaceae. of the American ostrich. bonariensis. moreover. A third species. as far as Mr. or oddly enough build one for themselves on the top of it. but Mr. namely. as far as it is known. They usually sit on their own eggs and rear their own young. when somewhat advanced in age and strength. or than in young snakes acquiring in their upper jaws. allied to our starlings. has not as yet been perfected. that few or none can possibly be hatched. I can see no more difficulty in this than in the unhatched young of other birds acquiring the instinct to break through their own shells. In this family several hen birds unite and lay first a few eggs in one nest and then in another. and throws some light on the singular instinct of the ostrich. not as especially endowed or created instincts. and asks. for those cuckoos which had such habits and structure best developed would be the most securely reared.blind desire. For if each part is liable to individual variations at all ages. but are still far from perfect. pecoris of North America. This bird. Hudson has ascertained. Some species of Molothrus. occasionally throwing out the nestlings of the stranger. They drop also many eggs on the bare ground. and the species present an interesting gradation in the perfection of their instincts. invariably lays its eggs in the nests of strangers. the strength. at intervals of two or three days. They often lay so many eggs--from fifteen to twenty--in the same foster-nest.

The males and fertile females do no work of any kind. I will give the observations which I made in some little detail. and she instantly set to work. so that the contrast in their appearance is great. They are incapable of making their own nests. Some species of Sphegidae (wasp-like insects) are likewise parasitic. and M. they could not even feed themselves. and have never been observed in the nests of F. as with that of the Molothrus or cuckoo. as any one may well be excused for doubting the existence of so extraordinary an instinct as that of making slaves. to whom I am much indebted for information on this and other subjects. the slaves work . sanguinea. Another species. but with plenty of the food which they like best. and with their larvae and pupae to stimulate them to work. and like their masters are much agitated and defend the nest: when the nest is much disturbed. and put all to rights. was likewise first discovered by P. when this insect finds a burrow already made and stored by another sphex. Hence. and many perished of hunger. This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber. So utterly helpless are the masters. I tried to approach the subject in a sceptical frame of mind. and becomes for the occasion parasitic. for these bees have not only had their instincts but their structure modified in accordance with their parasitic habits. Although fully trusting to the statements of Huber and Mr. or of feeding their own larvae. they did nothing. that when Huber shut up thirty of them without a slave. be not thus exterminated. if of advantage to the species. Formica sanguinea. do no other work. and its habits have been attended to by Mr. and actually carry their masters in their jaws. I opened fourteen nests of F. it would have been hopeless to speculate how so wonderful an instinct could have been perfected. the slaves occasionally come out. fusca). though most energetic and courageous in capturing slaves. Smith. Huber to be a slave-making ant. Smith. F. and found a few slaves in all. fed and saved the survivors. Many bees are parasitic. The slaves are black and not above half the size of their red masters. fusca) are found only in their own proper communities. Fabre has lately shown good reason for believing that. This case is more remarkable than that of the cuckoo. I can see no difficulty in natural selection making an occasional habit permanent. so that in one day's hunting I picked up no less than twenty lost and wasted eggs. it is the slaves which determine the migration. What can be more extraordinary than these well-ascertained facts? If we had not known of any other slave-making ant. and the larvae and pupae are exposed. In this case. for they do not possess the pollen-collecting apparatus which would have been indispensable if they had stored up food for their own young. and the workers or sterile females. although the Tachytes nigra generally makes its own burrow and stores it with paralysed prey for its own larvae. and regularly lay their eggs in the nests of other kinds of bees. SLAVE-MAKING INSTINCT. sanguinea. of the British Museum. made some cells and tended the larvae. Huber then introduced a single slave (F. it takes advantage of the prize.for a surprising number of eggs lie strewed over the plains. and they have to migrate. Males and fertile females of the slave-species (F. a better observer even than his celebrated father. the species would certainly become extinct in a single year. and if the insect whose nest and stored food are feloniously appropriated. yet that. without their aid. This species is found in the southern parts of England. When the old nest is found inconvenient. This ant is absolutely dependent on its slaves. When the nest is slightly disturbed.

As. who perhaps fancied that. Now I was curious to ascertain whether F. it is very courageous. fusca. According to Huber. probably depends merely on the slaves being captured in greater numbers in Switzerland than in England. During the months of June and July. This species is sometimes. their principal office is to search for aphides. I watched for many hours several nests in Surrey and Sussex. during these months. sometimes as many as three of these ants clinging to the legs of the slave-making F. and never saw a slave either leave or enter a nest. and I have seen it ferociously attack other ants. on three successive years. June and August. Hence. and it was a most interesting spectacle to behold the masters carefully carrying their slaves in their jaws instead of being carried by them. may be constantly seen bringing in materials for the nest. Smith informs me that he has watched the nests at various hours during May. and put them down on a bare spot near the place of combat. which they ascended together. but they were prevented from getting any pupae to rear as slaves. and when I had accidentally disturbed both nests. rufescens. they had been victorious in their late combat. I came across a community with an unusually large stock of slaves. and it was evident that they did at once distinguish them. The latter ruthlessly killed their small opponents and carried their dead bodies as food to their nest. in the month of July. Smith. it is clear that the slaves feel quite at home. Although so small a species. made into slaves. but in about a quarter of an hour. shortly after all the little yellow ants . I then dug up a small parcel of the pupae of F. on the other hand. probably in search of aphides or cocci. which they habitually make into slaves. One day I fortunately witnessed a migration of F. flava under a stone beneath a nest of the slave-making F. and marching along the same road to a tall Scotch-fir tree. they approached and were vigorously repulsed by an independent community of the slave species (F. Hence. as has been described by Mr. for we have seen that they eagerly and instantly seized the pupae of F. and they alone open and close the doors in the morning and evening. of F. fusca). the little ants attacked their big neighbours with surprising courage. either leave or enter the nest. F. sanguinea could distinguish the pupae of F. or even the earth from the nest. though present in large numbers in August. as Huber expressly states. from those of the little and furious F. which they rarely capture. The masters.energetically together with their masters in carrying them away to a place of safety. This difference in the usual habits of the masters and slaves in the two countries. sanguinea from one nest to another. sanguinea. the slaves in Switzerland habitually work with their masters in making the nest. whereas they were much terrified when they came across the pupae. the slaves are very few in number. after all. twenty-five yards distant. he considers them as strictly household slaves. who had ample opportunities for observation. they were eagerly seized and carried off by the tyrants. At the same time I laid on the same place a small parcel of the pupae of another species. and food of all kinds. as in the case of F. flava. In one instance I found to my surprise an independent community of F. though rarely. Another day my attention was struck by about a score of the slave-makers haunting the same spot. however. flava. with a few of these little yellow ants still clinging to the fragments of their nest. and has never seen the slaves. twenty-nine yards distant. both in Surrey and Hampshire. sanguinea. I thought that they might behave differently when more numerous. and I observed a few slaves mingled with their masters leaving the nest. flava. and quickly ran away. and evidently not in search of food. and. fusca. fusca from another nest. but Mr. During the year 1860.

their slaves and larvae. Let it be observed what a contrast the instinctive habits of F. We hear from mathematicians that bees have practically solved a recondite problem. carrying the dead bodies of F. to a very thick clump of heath. In England the masters alone usually leave the nest to collect building materials and food for themselves. as we have seen. sanguinea originated I will not pretend to conjecture. and when they migrate. I traced a long file of ants burdened with booty. But as ants which are not slave-makers. must have been close at hand. and one was perched motionless with its own pupa in its mouth on the top of a spray of heath. carrying a pupa. and the masters alone go on slave-making expeditions. One evening I visited another community of F. so beautifully adapted to its end. and the foreign ants thus unintentionally reared would then follow their proper instincts. sanguinea present with those of the continental F. but I was not able to find the desolated nest in the thick heath. and thus both collect food for the community. but will merely give an outline of the conclusions at which I have arrived. in regard to the wonderful instinct of making slaves. it is possible that such pupae originally stored as food might become developed. If their presence proved useful to the species which had seized them--if it were more advantageous to this species. rufescens. Formica sanguinea. the masters carry the slaves. which. fusca were rushing about in the greatest agitation. When the instinct was once acquired. CELL-MAKING INSTINCT OF THE HIVE-BEE. on the other hand. if scattered near their nests. does not determine its own migrations. and have made their cells of the proper shape to hold the . and do what work they could. is less aided by its slaves than the same species in Switzerland. sanguinea. though they did not need confirmation by me. The masters determine when and where a new nest shall be formed. By what steps the instinct of F.had crawled away. The nest. as I have seen. Such are the facts. whence I saw the last individual of F. originally for food. but chiefly the slaves. sanguinea. So that the masters in this country receive much less service from their slaves than they do in Switzerland. might by natural selection be strengthened and rendered permanent for the very different purpose of raising slaves. however. The latter does not build its own nest. and found a number of these ants returning home and entering their nests. I will not here enter on minute details on this subject. their aphides. carry off pupae of other species. if carried out to a much less extent even than in our British F. In Switzerland the slaves and masters work together. does not collect food for itself or its young. Both in Switzerland and England the slaves seem to have the exclusive care of the larvae. and cannot even feed itself: it is absolutely dependent on its numerous slaves. sanguinea emerge. possesses much fewer slaves. to capture workers than to procreate them--the habit of collecting pupae. making and bringing materials for the nest: both. and in the early part of the summer extremely few. an image of despair over its ravaged home. tend and milk as it may be called. will. He must be a dull man who can examine the exquisite structure of a comb. for two or three individuals of F. fusca (showing that it was not a migration) and numerous pupae. they took heart and carried off the pupae. without enthusiastic admiration. natural selection might increase and modify the instinct--always supposing each modification to be of use to the species--until an ant was formed as abjectly dependent on its slaves as is the Formica rufescens. for about forty yards back.

The Melipona itself is intermediate in structure between the hive and humble bee. When one cell rests on three other cells. placed in a double layer: each cell. and had made them of equal sizes and had arranged them symmetrically in a double layer. Accordingly I wrote to Professor Miller. sometimes adding to them short tubes of wax. and see whether Nature does not reveal to us her method of work. with the basal edges of its six sides bevelled so as to join an inverted pyramid. in addition. it seems at first quite inconceivable how they can make all the necessary angles and planes. with fitting tools and measures. the bees building perfectly flat walls of wax between the spheres which thus tend to intersect. Waterhouse. but are of the same thickness as the outer spherical portions. it occurred to me that if the Melipona had made its spheres at some given distance from each other. and yet each flat portion forms a part of two cells. In the series between the extreme perfection of the cells of the hive-bee and the simplicity of those of the humble-bee. It is obvious that the Melipona saves wax. is manifestly a gross imitation of the three-sided pyramidal base of the cell of the hive-bee. Let us look to the great principle of gradation. and. each cell consists of an outer spherical portion. enter into the composition of the bases of three adjoining cells on the opposite side. and the following view may. Granting whatever instincts you please. and this pyramid. from the spheres being nearly of the same size. in which the young are hatched. and what is more important. three. and likewise making separate and very irregular rounded cells of wax. Hence. but this is never permitted. and this geometer has kindly read over the following statement. These rhombs have certain angles. but more nearly related to the latter: it forms a nearly regular waxen comb of cylindrical cells. that these cells are always made at that degree of nearness to each other that they would have intersected or broken into each other if the spheres had been completed. or even perceive when they are correctly made. It has been remarked that a skilful workman. I was led to investigate this subject by Mr. with the least possible consumption of precious wax in their construction. be considered only as a modification of his theory. carefully described and figured by Pierre Huber. Reflecting on this case. who has shown that the form of the cell stands in close relation to the presence of adjoining cells. of three rhombs. so here. perhaps. by this manner of building. some large cells of wax for holding honey. of Cambridge. and the three which form the pyramidal base of a single cell on one side of the comb. I think. is very frequently and necessarily the case. At one end of a short series we have humble-bees. At the other end of the series we have the cells of the hive-bee. the resulting structure would have been as perfect as the comb of the hive-bee.greatest possible amount of honey. the three flat surfaces are united into a pyramid. though this is effected by a crowd of bees working in a dark hive. which. and tells me that it is strictly correct:-- . and of two. or more flat surfaces. But the important point to notice is. These latter cells are nearly spherical and of nearly equal sizes. drawn up from his information. is an hexagonal prism. would find it very difficult to make cells of wax of the true form. according as the cell adjoins two. and are aggregated into an irregular mass. the three plane surfaces in any one cell necessarily enter into the construction of three adjoining cells. which use their old cocoons to hold honey. as is well known. labour. as Huber has remarked. for the flat walls between the adjoining cells are not double. But the difficulty is not nearly so great as at first appears: all this beautiful work can be shown. we have the cells of the Mexican Melipona domestica. to follow from a few simple instincts. As in the cells of the hive-bee. three or more other cells.

rectangular strip of wax: the bees instantly began to excavate minute circular pits in it. the bees ceased to excavate. who has made numerous careful measurements. a thin and narrow. realised. Tegetmeier. as she already does her cylindrical cells. apparently by turning round on a fixed point. if we could slightly modify the instincts already possessed by the Melipona. and put between them a long. seeing that she already does so to a certain extent. and this is the greatest difficulty. so that each hexagonal prism was built upon the scalloped edge of a smooth basin. I separated two combs. wherever several bees had begun to excavate these basins near together. Hence we may safely conclude that. that the accuracy of the workmanship of the bee has been greatly exaggerated. that . then. if planes of intersection between the several spheres in both layers be formed. with the centre of each sphere at the distance of radius x sqrt(2) or radius x 1. the rims of the basins intersected or broke into each other. As soon as this occurred. that she always describes her spheres so as to intersect to a certain extent. in the same way as before. knife-edged ridge. But I hear from Professor Wyman. instead of on the straight edges of a three-sided pyramid as in the case of ordinary cells. rectangular piece of wax. through natural selection. from the centres of the six surrounding spheres in the same layer. and seeing what perfectly cylindrical burrows many insects make in wood. We must suppose the Melipona to arrange her cells in level layers. there will result a double layer of hexagonal prisms united together by pyramidal bases formed of three rhombs. that whatever the typical form of the cell may be. her inimitable architectural powers. if ever. so much so. and we must further suppose. they had begun their work at such a distance from each other that by the time the basins had acquired the above stated width (i. this bee would make a structure as wonderfully perfect as that of the hive-bee. and in themselves not very wonderful. We must suppose the Melipona to have the power of forming her cells truly spherical. and began to build up flat walls of wax on the lines of intersection between the basins. thick. instead of a thick. The bees instantly began on both sides to excavate little basins near to each other. and at the same distance from the centres of the adjoining spheres in the other and parallel layer.If a number of equal spheres be described with their centres placed in two parallel layers. but the ridge of wax was so thin. they made them wider and wider until they were converted into shallow basins. and of equal sizes.e. It was most interesting to observe that. and then she unites the points of intersection by perfectly flat surfaces. and as they deepened these little pits. it is rarely. and the rhombs and the sides of the hexagonal prisms will have every angle identically the same with the best measurements which have been made of the cells of the hive-bee. I then put into the hive. about the width of an ordinary cell). and were in depth about one sixth of the diameter of the sphere of which they formed a part. appearing to the eye perfectly true or parts of a sphere. coloured with vermilion. that she can somehow judge accurately at what distance to stand from her fellow-labourers when several are making their spheres. But this theory can be tested by experiment. By such modifications of instincts which in themselves are not very wonderful--hardly more wonderful than those which guide a bird to make its nest--I believe that the hive-bee has acquired. and of about the diameter of a cell. and this would not be very surprising.41421 (or at some lesser distance). Following the example of Mr. but she is already so far enabled to judge of distance.

I put the comb back into the hive. In ordinary combs it has appeared to me that the bees do not always succeed in working at exactly the same rate from the opposite sides. and allowed the bees to go on working for a short time. and these flat bases. The bees must have worked at very nearly the same rate in circularly gnawing away and deepening the basins on both sides of the ridge of vermilion wax. and they gnaw this away from the opposite sides. by standing at the proper distance from each other. but never allowing the spheres to break into each other. In one well-marked instance. and by endeavouring to make equal spherical hollows. and thus flatten it. I could show that they are conformable with my theory. do make a rough. where I suppose that the bees had excavated too quickly. as far as the eye could judge. if they had been excavated to the same depth as in the former experiment. for I have noticed half-completed rhombs at the base of a just-commenced cell. as soon as they had been a little deepened. but the work. until the hexagonal walls are commenced. and if I had space. in other parts. exactly along the planes of imaginary intersection between the basins on the opposite side of the ridge of wax. by stopping work at the planes of intersection. I do not see that there is any difficulty in the bees. as far as I have seen. Now bees. Considering how flexible thin wax is. by excavating at the same rate. along the plane of intersection between . and they stopped their excavations in due time. if the bees were to build for themselves a thin wall of wax. is not. as the case may be. only small portions. or the two plates. would have broken into each other from the opposite sides. were situated. From the experiment of the ridge of vermilion wax we can see that. and convex on the opposed side where the bees had worked less quickly. did not suffer this to happen. In some parts. from the extreme thinness of the little plate. had not been neatly performed. but I will not here enter on details.the bottoms of the basins. strictly correct. formed by thin little plates of the vermilion wax left ungnawed. and then stopping their work. circumferential wall or rim all round the comb. and again examined the cell. the first commencement having always been a little hood of wax. and had become PERFECTLY FLAT: it was absolutely impossible. always working circularly as they deepen each cell. They do not make the whole three-sided pyramidal base of any one cell at the same time. so that the basins. large portions of a rhombic plate were thus left between the opposed basins. which were slightly concave on one side. Some of these statements differ from those made by the justly celebrated elder Huber. and they never complete the upper edges of the rhombic plates. that the very first cell is excavated out of a little parallel-sided wall of wax. but it would be a great error to suppose that the bees cannot build up a rough wall of wax in the proper position--that is. in order to have thus succeeded in leaving flat plates between the basins. and I suspect that the bees in such cases stand in the opposed cells and push and bend the ductile and warm wax (which as I have tried is easily done) into its proper intermediate plane. and I found that the rhombic plate had been completed. that they could have effected this by gnawing away the convex side. but I am convinced of their accuracy. from the unnatural state of things. whilst at work on the two sides of a strip of wax. came to have flat bases. We see how important a part excavation plays in the construction of the cells. The bees. as may be clearly seen by examining the edge of a growing comb. they could make their cells of the proper shape. perceiving when they have gnawed the wax away to the proper thinness. Huber's statement. but only that one rhombic plate which stands on the extreme growing margin. however.

by striking imaginary spheres. We shall understand how they work. in its strictly proper place. by being largely gnawed away on both sides. I have several specimens showing clearly that they can do this. It seems at first to add to the difficulty of understanding how the cells are made. From all the cells. When bees have a place on which they can stand in their proper positions for working--for instance. from twenty-one measurements. by covering the edges of the hexagonal walls of a single cell. strength is continually given to the comb. The work of construction seems to be a sort of balance struck between many bees. how often the bees would pull down and rebuild in different ways the same cell. so that. having a mean thickness. as far as I have seen. and then building up. being thus crowned by a strong coping of wax. the bees can cluster and crawl over the comb without injuring the delicate hexagonal walls. and I invariably found that the colour was most delicately diffused by the bees--as delicately as a painter could have done it with his brush--by atoms of the coloured wax having been taken from the spot on which it had been placed. the masons always piling up the cut-away cement. so that the comb has to be built over one face of the slip--in this case the bees can lay the foundations of one wall of a new hexagon. which will ultimately be left. 1/352 of an inch in thickness. flexures may sometimes be observed. and worked into the growing edges of the cells all round. These walls. all instinctively standing at the same relative distance from each other. The manner in which the bees build is curious. I was able practically to show this fact. on a slip of wood. of 1/229 of an inch. We shall thus have a thin wall steadily growing upward but always crowned by a gigantic coping. or leaving ungnawed. very thin wall is left in the middle. It was really curious to note in cases of difficulty. vary greatly in thickness. they never gnaw away and finish off the angles of a cell till a large part both of that cell and of the adjoining cells has been built. till a smooth. sometimes recurring to a shape which they had at first rejected. all trying to sweep equal spheres. on an average of twelve measurements made near the border of the comb. one bee after working a short time at one cell going to another. whereas the basal rhomboidal plates are thicker. projecting beyond the other completed cells. the planes of intersection between these spheres. being. as Professor Miller has kindly ascertained for me. with an extremely thin layer of melted vermilion wax. corresponding in position to the planes of the rhombic basal plates of future cells. Even in the rude circumferential rim or wall of wax round a growing comb. but. It suffices that the bees should be enabled to stand at their proper relative distances from each other and from the walls of the last completed cells. By the above singular manner of building. This capacity in bees of laying down under certain circumstances a rough wall in its proper place . as Huber has stated. or the extreme margin of the circumferential rim of a growing comb. by supposing masons first to pile up a broad ridge of cement. both those just commenced and those completed. they always make the first rough wall from ten to twenty times thicker than the excessively thin finished wall of the cell. and then to begin cutting it away equally on both sides near the ground. nearly in the proportion of three to two. But the rough wall of wax has in every case to be finished off. as when two pieces of comb met at an angle. a score of individuals work even at the commencement of the first cell. with the utmost ultimate economy of wax. and adding fresh cement on the summit of the ridge.two adjoining spheres. they can build up a wall intermediate between two adjoining spheres. that a multitude of bees all work together. placed directly under the middle of a comb growing downwards. and then.

from the same cause. as it probably often has determined. can be explained by natural selection having taken . Moreover. and building up intermediate planes. and the Melipona would make a comb as perfect as that of the hive-bee. could have profited the progenitors of the hive-bee? I think the answer is not difficult: cells constructed like those of the bee or the wasp gain in strength. natural selection could not lead. and the time consumed in collecting the honey. or on quite distinct causes. is absolutely perfect in economising labour and wax. Again. nearer together. always standing at the proper relative distance from the parts of the cells just begun. it would continually be more and more advantageous to our humble-bees. Of course the success of the species may be dependent on the number of its enemies. and consequently required a store of honey: there can in this case be no doubt that it would be an advantage to our imaginary humble-bee if a slight modification of her instincts led her to make her waxen cells near together. and let us further suppose that the community lived through the winter. as I believe. Hence the saving of wax by largely saving honey. Hence.between two just-commenced cells. Thus. and in the materials of which they are constructed. many bees have to remain idle for many days during the process of secretion. and aggregated into a mass. as far as we can see. for in this case a large part of the bounding surface of each cell would serve to bound the adjoining cells. whether a bee allied to our humble-bees could exist in large numbers in any country. it may reasonably be asked. it would be advantageous to the Melipona. if she were to make her cells closer together. must be an important element of success any family of bees. Beyond this stage of perfection in architecture. as it bears on a fact. With respect to the formation of wax. like the cells of the Melipona. or parasites. the spherical surfaces would wholly disappear and be replaced by plane surfaces. Nor does there seem to me any great difficulty in a single insect (as in the case of a queen-wasp) making hexagonal cells. namely. that the cells on the extreme margin of wasp-combs are sometimes strictly hexagonal. as we have seen. for the comb of the hive-bee. A large store of honey is indispensable to support a large stock of bees during the winter. and more regular in every way than at present. for a wall in common even to two adjoining cells would save some little labour and wax. is important. if they were to make their cells more and more regular. and save much in labour and space. if she were to work alternately on the inside and outside of two or three cells commenced at the same time. the most wonderful of all known instincts. but I have not space here to enter on this subject. Tegetmeier that it has been experimentally proved that from twelve to fifteen pounds of dry sugar are consumed by a hive of bees for the secretion of a pound of wax. and the security of the hive is known mainly to depend on a large number of bees being supported. and much labour and wax would be saved. and so be altogether independent of the quantity of honey which the bees can collect. and I am informed by Mr. But let us suppose that this latter circumstance determined. which seems at first subversive of the foregoing theory. so as to intersect a little. that of the hive-bee. how a long and graduated succession of modified architectural instincts. so that a prodigious quantity of fluid nectar must be collected and consumed by the bees in a hive for the secretion of the wax necessary for the construction of their combs. each profitable to the individual under its conditions of life. sweeping spheres or cylinders. for then. it is known that bees are often hard pressed to get sufficient nectar. As natural selection acts only by the accumulation of slight modifications of structure or instinct. all tending towards the present perfect plan of construction.

No doubt many instincts of very difficult explanation could be opposed to the theory of natural selection--cases. than they know what are the several angles of the hexagonal prisms and of the basal rhombic plates. subsequently to. more and more perfectly led the bees to sweep equal spheres at a given distance from each other in a double layer. having succeeded best.advantage of numerous. believed to be moistened with saliva. It has been objected to the foregoing view of the origin of instincts that "the variations of structure and of instinct must have been simultaneous and accurately adjusted to each other. in which no intermediate gradations are known to exist. led the bird to become more and more of a seed-eater? In this case the beak is supposed to be slowly modified by natural selection. To take as an illustration the case of the larger titmouse. but let the feet of the titmouse vary and grow larger from correlation with the beak. in which we cannot see how an instinct could have originated. should at last produce a species with instincts leading it to neglect other materials and to make its nest exclusively of inspissated saliva? And so in other cases. and even with flakes of this substance. Some birds build their nests of mud. slight modifications of simpler instincts. cases of instincts of such trifling importance. and least waste of honey in the secretion of wax. but in accordance with. which secreted more and more saliva. Now what special difficulty would there be in natural selection preserving all the slight individual variations in the shape of the beak. at the same time that habit. and one of the swifts of North America makes its nest (as I have seen) of sticks agglutinated with saliva. however. or compulsion. that individual swarm which thus made the best cells with least labour. the motive power of the process of natural selection having been the construction of cells of due strength and of the proper size and shape for the larvae. this bird often holds the seeds of the yew between its feet on a branch. that they could hardly have been acted on by natural . It must. Is it then very improbable that the natural selection of individual swifts." The force of this objection rests entirely on the assumption that the changes in the instincts and structure are abrupt. as well constructed for this purpose as that of the nuthatch. be admitted that in many instances we cannot conjecture whether it was instinct or structure which first varied. and hammers with its beak till it gets at the kernel. or spontaneous variations of taste. natural selection having. and to build up and excavate the wax along the planes of intersection. by slow degrees. successive. of course. or from any other unknown cause. The bees. OBJECTIONS TO THE THEORY OF NATURAL SELECTION AS APPLIED TO INSTINCTS: NEUTER AND STERILE INSECTS. no more knowing that they swept their spheres at one particular distance from each other. as a modification in the one without an immediate corresponding change in the other would have been fatal. and it is not improbable that such larger feet would lead the bird to climb more and more until it acquired the remarkable climbing instinct and power of the nuthatch. slowly changing habits or taste. cases. To take one more case: few instincts are more remarkable than that which leads the swift of the Eastern Islands to make its nest wholly of inspissated saliva. this being effected with the greatest possible economy of labour and wax. and having transmitted their newly-acquired economical instincts to new swarms. which were better and better adapted to break open the seeds. (Parus major) alluded to in a previous chapter. In this case a gradual change of structure is supposed to lead to changed instinctive habits. which in their turn will have had the best chance of succeeding in the struggle for existence. until a beak was formed.

and it had been profitable to the community that a number should have been annually born capable of work. the wonderful difference in this respect between the workers and the perfect females would have been better exemplified by the hive-bee. by individuals having been born with slight profitable modifications. and so onwards. and in instinct. But with the working ant we have an insect differing greatly from its parents. I should have unhesitatingly assumed that all its characters had been slowly acquired through natural selection. We have even slight differences in the horns of different breeds of cattle in relation to an artificially imperfect state of the male sex. . as in the nuptial plumage of many birds. relatively to the length of the horns in both the bulls and cows of these same breeds. as in the shape of the thorax. but incapable of procreation.selection. and actually fatal to the whole theory. We have differences correlated not only with one sex. Hence. yet absolutely sterile. so that it could never have transmitted successively acquired modifications of structure or instinct to its progeny. Breeders of cattle wish the flesh and fat to be well marbled together. but will confine myself to one special difficulty. and consequently must believe that they were independently acquired through natural selection. namely. for it can be shown that some insects and other articulate animals in a state of nature occasionally become sterile. but not much greater than that of any other striking modification of structure. though appearing insuperable. disappears. An animal thus characterized has been slaughtered. This difficulty. which were inherited by the offspring. and in the hooked jaws of the male salmon. and yet. cases of instincts almost identically the same in animals so remote in the scale of nature that we cannot account for their similarity by inheritance from a common progenitor. I can see no especial difficulty in this having been effected through natural selection. I allude to the neuters or sterile females in insect communities: for these neuters often differ widely in instinct and in structure from both the males and fertile females. But I must pass over this preliminary difficulty. as I believe. the difficulty lies in understanding how such correlated modifications of structure could have been slowly accumulated by natural selection. and if such insects had been social. I can see no great difficulty in any character becoming correlated with the sterile condition of certain members of insect communities. that of working or sterile ants. when it is remembered that selection may be applied to the family. let it be remembered that we have innumerable instances. for oxen of certain breeds have longer horns than the oxen of other breeds. I will not here enter on these several cases. which at first appeared to me insuperable. of all sorts of differences of inherited structure which are correlated with certain ages and with either sex. If a working ant or other neuter insect had been an ordinary animal. but I will here take only a single case. but with that short period when the reproductive system is active. as well as to the individual. It may well be asked how it is possible to reconcile this case with the theory of natural selection? First. and may thus gain the desired end. As far as instinct alone is concerned. How the workers have been rendered sterile is a difficulty. both in our domestic productions and in those in a state of nature. The subject well deserves to be discussed at great length. they cannot propagate their kind. and in being destitute of wings and sometimes of eyes. from being sterile. is lessened. or. The great difficulty lies in the working ants differing widely from both the males and the fertile females in structure. and that these again varied and again were selected.

that the successive. Such faith may be placed in the power of selection that a breed of cattle. presenting gradations of structure. not only from the fertile females and males. and the double sterile plants with the neuters of the same community. have been rendered different from the fertile males and females through natural selection. be formed by carefully watching which individual bulls and cows. for the sake of gaining a serviceable end. as I believe.but the breeder has gone with confidence to the same stock and has succeeded. correlated with the sterile condition of certain members of the community. and they have an enormously developed abdomen which secretes a sort of honey. but in some few alone. According to this view we ought occasionally to find in the same nest neuter-insects. have proved advantageous. the workers of one caste alone carry a wonderful sort of shield on their heads. and that the extreme forms can be linked together by individuals taken out of the same nest: I have myself compared perfect gradations of this kind. In the simpler case of neuter insects all of one caste. considering how few neuter-insects out of Europe have been carefully examined. namely. until that prodigious amount of difference between the fertile and sterile females of the same species has been produced which we see in many social insects. and not to the individual. all the neuters ultimately came to be thus characterized. But we have not as yet touched on the acme of the difficulty. Smith has shown that the neuters of several British ants differ surprisingly from each other in size and sometimes in colour. being as distinct from each other as are any two species of the same genus. slight. This process must have been repeated many times. but from each other. moreover. sometimes to an almost incredible degree. do not generally graduate into each other. always yielding oxen with extraordinarily long horns. we may conclude from the analogy of ordinary variations. with jaws and instincts extraordinarily different: in Cryptocerus. by which alone the variety can be propagated. we may conclude that slight modifications of structure or of instinct. F. profitable modifications did not first arise in all the neuters in the same nest. some varieties of the double annual stock. in Eciton. and that by the survival of the communities with females which produced most neuters having the advantageous modification. It will indeed be thought that I have an overweening confidence in the principle of natural selection. always produce a large proportion of seedlings bearing double and quite sterile flowers. could. the use of which is quite unknown: in the Mexican Myrmecocystus. As with the varieties of the stock. or the domestic cattle as they may be called. supplying the place of that excreted by the aphides. even not rarely. may be compared with the fertile male and female ants. the fact that the neuters of several ants differ. there are working and soldier neuters. they are fed by the workers of another caste. Hence. and are thus divided into two or even three castes. when matched. it is probable. selection has been applied to the family. which our European ants guard and imprison. or rather as any two genera of the same family. produced oxen with the longest horns. Thus. when I do not admit that such wonderful and well-established facts at once annihilate the theory. Mr. Here is a better and real illustration: According to M. consequently the fertile males and females have flourished. The castes. and this we do find. These latter. and yet no one ox would ever have propagated its kind. but they likewise yield some single and fertile plants. which. It sometimes . from having been long and carefully selected to the right degree. the workers of one caste never leave the nest. and transmitted to their fertile offspring a tendency to produce sterile members with the same modifications. Verlot. but are perfectly well defined. so with social insects.

of the equally complex case. For the workers of Myrmica have not even rudiments of ocelli. of certain Brazilian crustaceans likewise appearing under two widely distinct male forms. in this species. I may digress by adding. Bates. until all the workers were in this condition. and by Fritz Muller. I speak confidently on this latter point. through the survival of the parents which generated them. and jaws nearly five times as big. or all of small size with widely different jaws. of the jaws which I dissected from the workers of the several sizes. the larger workers have simple eyes (ocelli). while those of an intermediate size are scanty in numbers. yet they graduate insensibly into each other. Smith has observed. Mr." has described analogous cases. F. all of large size with one form of jaw. could form a species which should regularly produce neuters. as in the case of the driver ant. . that if the smaller workers had been the most useful to the community. though the male and female ants of this genus have well-developed ocelli. and in the form and number of the teeth. I believe that natural selection. which produced more and more of the smaller workers. of certain Malayan butterflies regularly appearing under two or even three distinct female forms. Wallace. and many sixteen feet high. With these facts before me. one set of workers of one size and structure. Lubbock made drawings for me. I can affirm that the eyes are far more rudimentary in the smaller workers than can be accounted for merely by their proportionately lesser size. a graduated series having first been formed. So that here we have two bodies of sterile workers in the same nest. differing not only in size. and. by acting on the fertile ants or parents. we should then have had a species of ant with neuters in nearly the same condition as those of Myrmica. I may give one other case: so confidently did I expect occasionally to find gradations of important structures between the different castes of neuters in the same species. But the important fact for us is that. of whom many were five feet four inches high. F. But this subject need not here be discussed. The jaws. that I gladly availed myself of Mr. or lastly. until none with an intermediate structure were produced. An analogous explanation has been given by Mr. with the camera lucida. of the working ants of the several sizes differed wonderfully in shape. Having carefully dissected several specimens of these workers. which. The reader will perhaps best appreciate the amount of difference in these workers by my giving. can be plainly distinguished. and then the extreme forms having been produced in greater and greater numbers. with some few of intermediate size. as Mr. and this is the greatest difficulty. that the workers of intermediate size have their ocelli in an exactly intermediate condition. yet connected by some few members in an intermediate condition. though I dare not assert so positively.happens that the larger or the smaller sized workers are the most numerous. moreover. but a strictly accurate illustration: the difference was the same as if we were to see a set of workmen building a house. as Sir J. but we must in addition suppose that the larger workmen had heads four instead of three times as big as those of the smaller men. or that both large and small are numerous. not the actual measurements. but in their organs of vision. Smith's offer of numerous specimens from the same nest of the driver ant (Anomma) of West Africa. as does the widely-different structure of their jaws. in his interesting "Naturalist on the Amazons. whereas the smaller workers have their ocelli rudimentary. and those males and females had been continually selected. and simultaneously another set of workers of a different size and structure. Formica flava has larger and smaller workers. though small. though the workers can be grouped into castes of different sizes. and I fully believe.

with all my faith in natural selection. which are in any way profitable. as with plants. has originated. on the principle of inheritance. the wonderful fact of two distinctly defined castes of sterile workers existing in the same nest. we can understand. therefore. But I must confess. discussed this case. while man works by acquired knowledge and manufactured instruments. is very interesting. how it is that the thrush of tropical South America lines its nest with mud. both widely different from each other and from their parents. annihilate it. as advanced by Lamarck. species. I have endeavoured in this chapter briefly to show that the mental qualities of our domestic animals vary. Therefore. No one will dispute that instincts are of the highest importance to each animal. also. to the best of my judgment. but distinct. though animals take advantage of the instincts of others. In many cases habit or use and disuse have probably come into play. of "Natura non facit saltum. For instance. on the same principle that the division of labour is useful to civilised man. had not the case of these neuter insects led me to this conclusion. with only a small hole left in the plaster through which the males feed them and their young when hatched. confined to the workers of sterile females. could not possibly affect the males and fertile females. in order to show the power of natural selection. On the other hand. and is plainly explicable on the foregoing views. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. yet often retaining nearly the same instincts.I have now explained how. work by inherited instincts and by inherited organs or tools." to roost . which alone leave descendants. that no instinct can be shown to have been produced for the good of other animals. Ants. I believe. I am surprised that no one has advanced this demonstrative case of neuter insects. I have. against the well-known doctrine of inherited habit." is applicable to instincts as well as to corporeal structure. the fact that instincts are not always absolutely perfect and are liable to mistakes. build "cock-nests. any amount of modification may be effected by the accumulation of numerous. in the same peculiar manner as does our British thrush. when inhabiting distant parts of the world and living under considerably different conditions of life. that the canon in natural history. there is no real difficulty. how it is that the male wrens (Troglodytes) of North America. but none of the cases of difficulty. For peculiar habits. as it proves that with animals. This theory is also strengthened by some few other facts in regard to instincts. and likewise because this is by far the most serious special difficulty which my theory has encountered. as by that common case of closely allied. We can see how useful their production may have been to a social community of ants. however long they might be followed. I should never have anticipated that this principle could have been efficient in so high a degree. The case. slight. under changing conditions of life. spontaneous variations. SUMMARY. how it is that the Hornbills of Africa and India have the same extraordinary instinct of plastering up and imprisoning the females in a hole in a tree. however. at some little but wholly insufficient length. but is otherwise inexplicable--all tend to corroborate the theory of natural selection. I do not pretend that the facts given in this chapter strengthen in any great degree my theory. in natural selection accumulating to any extent slight modifications of instinct which are in any way useful. that. and that the variations are inherited. without exercise or habit having been brought into play.

with reference to my theory. The view commonly entertained by naturalists is that species. as I shall show. The fertility of varieties. cannot have been acquired. the larvae of ichneumonidae feeding within the live bodies of caterpillars. as may be clearly seen in the state of the male element in both plants and animals. It is an incidental result of differences in the reproductive systems of the parent-species. as far as the microscope reveals. it may not be a logical deduction. is. In the first case the two sexual elements which go to form the embryo are perfect. owing to the sterility in both cases being looked on as a special endowment. but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its foster-brothers. but as small consequences of one general law leading to the advancement of all organic beings--namely. though the formative organs themselves are perfect in structure. affected by close interbreeding. have their reproductive organs functionally impotent. have been specially endowed with sterility. namely. like the males of our Kitty-wrens. Finally. yet when intercrossed they produce either few or no offspring. not universal. two classes of facts. more especially as the sterility of species when first crossed. let the strongest live and the weakest die. to a large extent fundamentally different.--a habit wholly unlike that of any other known bird. multiply. Distinction between the sterility of first crosses and of hybrids--Sterility various in degree. beyond the province of our reasoning powers. ants making slaves. which is common to the two cases. has to be considered. The subject is in many ways important for us.in. not accumulated by natural selection--Causes of the sterility of first crosses and of hybrids--Parallelism between the effects of changed conditions of life and of crossing--Dimorphism and trimorphism--Fertility of varieties when crossed and of their mongrel offspring not universal--Hybrids and mongrels compared independently of their fertility--Summary. in the second case they are either not at all developed. when crossed. This view certainly seems at first highly probable. or are imperfectly developed. that is of the forms known or believed to be descended from common parents. in order to prevent their confusion. and likewise the fertility of their mongrel offspring. CHAPTER IX. on the other hand. and the sterility of the hybrids produced from them. the sterility of species when first crossed. but incidental on other differences. The distinction probably has been slurred over. not as specially endowed or created instincts. HYBRIDISM. by the preservation of successive profitable degrees of sterility. In treating this subject. removed by domestication--Laws governing the sterility of hybrids--Sterility not a special endowment. have generally been confounded. for species living together could hardly have been kept distinct had they been capable of freely crossing. This distinction is important. Pure species have of course their organs of reproduction in a perfect condition. and that of their hybrid offspring. when the cause of the sterility. when intercrossed. Hybrids. vary. of equal .

Gartner is obliged carefully to count the seeds. It is also most instructive to compare--but I have not space here to enter on details--the evidence advanced by our best botanists on the question whether certain doubtful forms should be ranked as species or varieties. but then he cuts the knot. in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired. It is certain. he unhesitatingly ranks them as varieties. must be secluded in order to prevent pollen being brought to it by insects from other plants. But causes of serious error here intervene: a plant. It can thus be shown that neither sterility nor fertility affords any certain distinction between species and varieties. which the best botanists rank as varieties.importance with the sterility of species. Kolreuter and Gartner. yet he asserts positively that their fertility never increases. DEGREES OF STERILITY. In regard to the sterility of hybrids in successive generations. makes the rule equally universal. that the sterility of various species when crossed is so different in degree and graduates away so insensibly. First. That these processes are often injurious to the fertility of a plant cannot be doubted. for Gartner gives in his table about a score of cases of plants which he castrated. and (excluding all cases such as the Leguminosae. what is often more important. but generally decreases greatly and suddenly. as Gartner repeatedly crossed some forms. and found them absolutely sterile. with the average number produced by both pure parent-species in a state of nature. namely Kolreuter and Gartner. and is doubtful in the same degree as is the evidence derived from other constitutional and structural differences. Gartner. and the maximum produced by their hybrid offspring. and. It is impossible to study the several memoirs and works of those two conscientious and admirable observers. Nearly all the plants experimented on by Gartner were potted. He always compares the maximum number of seeds produced by two species when first crossed. The evidence from this source graduates away. and were kept in a chamber in his house. for six or seven. for the sterility of species when crossed and of their hybrid offspring. such as the common red and blue pimpernels (Anagallis arvensis and coerulea). I think no better evidence of this can be required than that the two most experienced observers who have ever lived. though Gartner was enabled to rear some hybrids. without being deeply impressed with the high generality of some degree of sterility. and in one case for ten generations. for in ten cases in which he found two forms. in order to show that there is any degree of sterility. for it seems to make a broad and clear distinction between varieties and species. when intercrossed. also. Moreover. quite fertile together. With respect . considered by most authors as distinct species. carefully guarding them from a cross with either pure parent. as he believed. who almost devoted their lives to this subject. we may doubt whether many species are really so sterile. must be castrated. on the other hand. and. or by the same observer from experiments made during different years. and artificially fertilised with their own pollen. to be hybridised. arrived at diametrically opposite conclusions in regard to some of the very same forms. But in these and in many other cases. on the one hand. that for all practical purposes it is most difficult to say where perfect fertility ends and sterility begins. with the evidence from fertility adduced by different hybridisers. Kolreuter makes the rule universal. that the fertility of pure species is so easily affected by various circumstances. and he disputes the entire fertility of Kolreuter's ten cases.

namely. notwithstanding the frequent ill effects from manipulation. though this pollen can be proved to be perfectly sound by fertilising other plants or species. by too close interbreeding. already lessened by their hybrid origin. all the individuals are in this peculiar condition. this is often transmitted in an augmented degree to the offspring. and goes on increasing. the visits of insects must be carefully prevented during the flowering season: hence hybrids. . either from the same plant or from another plant of the same hybrid nature. may. their fertility. certain abnormal individuals. and this would have insured in each generation a cross with pollen from a distinct flower. which I never saw to occur in a case of its natural fecundation. so careful an observer as Gartner would have castrated his hybrids. and this would probably be injurious to their fertility. sometimes decidedly increases. The difference in their results may. in contrast with those spontaneously self-fertilised. showing on the one hand that an occasional cross with a distinct individual or variety increases the vigour and fertility of the offspring. In the genus Hippeastrum. that if even the less fertile hybrids be artificially fertilised with hybrid pollen of the same kind. namely. it may first be noticed that when any deviation in structure or constitution is common to both parents. and in other species all the individuals. and as the parent-species. though probably often on the same plant. and Rev. Of his many important statements I will here give only a single one as an example. revolutum produced a plant. W. Moreover. in the process of artificial fertilisation. I think. I have made so many experiments and collected so many facts. namely. But I believe that their fertility has been diminished in nearly all these cases by an independent cause. generally grow in the same garden." So that here we have perfect. in Corydalis as shown by Professor Hildebrand. Herbert.to this decrease. so that a cross between two flowers. So that with some species. that I cannot doubt the correctness of this conclusion. Scott and Fritz Muller. be accounted for by too close interbreeding having been avoided. or other allied hybrids. be in part accounted for by Herbert's great horticultural skill. whenever complicated experiments are in progress. And thus. He experimented on some of the very same species as did Gartner. the Hon. in a first cross between two distinct species. or even more than commonly perfect fertility. Now. namely. the strange fact of an increase of fertility in the successive generations of ARTIFICIALLY FERTILISED hybrids. and both sexual elements in hybrid plants are already affected in some degree. that "every ovule in a pod of Crinum capense fertilised by C. He is as emphatic in his conclusion that some hybrids are perfectly fertile--as fertile as the pure parent-species--as are Kolreuter and Gartner that some degree of sterility between distinct species is a universal law of nature. namely. but not by pollen from the same plant. as from the anthers of the flower itself which is to be fertilised. in various orchids as shown by Mr. Verbascum and Passiflora. Now let us turn to the results arrived at by a third most experienced hybridiser. as I believe. that individual plants of certain species of Lobelia. Hybrids are seldom raised by experimentalists in great numbers. and on the other hand that very close interbreeding lessens their vigour and fertility. will generally be fertilised during each generation by pollen from the same flower. pollen is as often taken by chance (as I know from my own experience) from the anthers of another flower. if left to themselves. This case of the Crinum leads me to refer to a singular fact. can easily be fertilised by the pollen from a distinct species. would be thus effected. I am strengthened in this conviction by a remarkable statement repeatedly made by Gartner. and by his having hot-houses at his command.

as not one of these breeds freely in confinement. etc. when fairly treated. owing to few animals breeding freely under confinement. Any one may readily convince himself of the efficiency of insect agency by examining the flowers of the more sterile kinds of hybrid Rhododendrons. so as to avoid the ill effects of close interbreeding. with respect to the fertility in successive generations of the more fertile hybrid animals. in opposition to the constantly repeated admonition of every breeder. In regard to animals. though not made with scientific precision. It should. for instance. informs me that he raises stocks for grafting from a hybrid between Rhod. Mr.. for he will find on their stigmas plenty of pollen brought from other flowers. a bulb of Hippeastrum aulicum produced four flowers. much fewer experiments have been carefully tried than with plants. then we may infer that animals more widely distinct in the scale of nature can be crossed more easily than in the case of plants. it is not at all surprising that the inherent sterility in the hybrids should have gone on increasing. The practical experiments of horticulturists.can actually be hybridised much more readily than they can be fertilised by pollen from the same individual plant! To give one instance. be borne in mind that. brothers and sisters have usually been crossed in each successive generation. but the hybrids themselves are. Noble. species most widely dissimilar in general habit. if the genera of animals are as distinct from each other as are the genera of plants. and bore good seed." Had hybrids. Rhododendron. that is. Herbert asserts that a hybrid from Calceolaria integrifolia and plantaginea. or that their hybrids. If our systematic arrangements can be trusted. yet many of these hybrids seed freely. It is notorious in how complicated a manner the species of Pelargonium. On the contrary. deserve some notice. For instance. Although I know of hardly any thoroughly well-authenticated cases of . and such alone are fairly treated. Horticulturists raise large beds of the same hybrid. we have no right to expect that the first crosses between them and the canary. and the injurious influence of close interbreeding is thus prevented. the fact would have been notorious to nurserymen. however. as Gartner believed to be the case. Again. Herbert tried similar experiments during many years. few experiments have been fairly tried: for instance. I hardly know of an instance in which two families of the same hybrid have been raised at the same time from different parents." I have taken some pains to ascertain the degree of fertility of some of the complex crosses of Rhododendrons. whereas the pod impregnated by the pollen of the hybrid made vigorous growth and rapid progress to maturity. Fuchsia. And in this case." Mr. and always with the same result. C. and the fourth was subsequently fertilised by the pollen of a compound hybrid descended from three distinct species: the result was that "the ovaries of the three first flowers soon ceased to grow. for by insect agency the several individuals are allowed to cross freely with each other. "reproduces itself as perfectly as if it had been a natural species from the mountains of Chile. I think. and that this hybrid "seeds as freely as it is possible to imagine. ponticum and catawbiense. Calceolaria. but. and I am assured that many of them are perfectly fertile. These cases serve to show on what slight and mysterious causes the lesser or greater fertility of a species sometimes depends. should be perfectly fertile. the canary-bird has been crossed with nine distinct species of finches. and after a few days perished entirely. always gone on decreasing in fertility in each successive generation. three were fertilised by Herbert with their own pollen. Petunia. have been crossed. which produce no pollen. more sterile. which vegetated freely.

is an extremely general result. yet. which are highly fertile when crossed with one of the parent-species. With our domesticated animals. This was effected by Mr. or that the hybrids subsequently reared under domestication became quite fertile. the various races when crossed together are quite fertile. voice. but as one capable of being removed by domestication. yet in many cases they are descended from two or more wild species. seems by far the most probable. Quatrefages states that the hybrids from two moths (Bombyx cynthia and arrindia) were proved in Paris to be fertile inter se for eight generations. almost certain that our dogs are descended from several wild stocks. in order to prevent their crossing and blending together in utter confusion. with perhaps the exception of certain indigenous domestic dogs of South America. Our chief object will be to see whether or not these laws indicate that species have been specially endowed with this quality. but that it cannot. I have reason to believe that the hybrids from Cervulus vaginalis and Reevesii. whether the several aboriginal species would at first have freely bred together and have produced quite fertile hybrids. where neither pure parent-species exists. not as an indelible characteristic. either give up the belief of the universal sterility of species when crossed. under our present state of knowledge. Blyth on their differences in habits. and from the observations by Rutimeyer on their important osteological differences. and from these two birds he raised no less than eight hybrids (grandchildren of the pure geese) from one nest. constitution.perfectly fertile hybrid animals. It has lately been asserted that two such distinct species as the hare and rabbit. The following conclusions are drawn up chiefly from Gartner's admirable work on the hybridisation . or we must look at this sterility in animals. it may be concluded that some degree of sterility. and from Phasianus colchicus with P. In India. The same remarks may be extended to the two chief races of the pig. Blyth and Captain Hutton. and as they are kept for profit. have often bred in this country with either pure parent. they must certainly be highly or perfectly fertile. cygnoides). are perfectly fertile. indeed. hardly be doubted. be considered as absolutely universal. We must. From this fact we must conclude either that the aboriginal parent-species at first produced perfectly fertile hybrids. The hybrids from the common and Chinese geese (A. Eyton. for instance. namely Mr. all are quite fertile together. So again I have lately acquired decisive evidence that the crossed offspring from the Indian humped and common cattle are inter se perfectly fertile.. It is. but analogy makes me greatly doubt. who raised two hybrids from the same parents. as well as from those by Mr. etc. for I am assured by two eminently capable judges. This latter alternative. Finally. these cross-bred geese must be far more fertile. and in one single instance they have bred inter se. and can. however. when they can be got to breed together. these two forms must be regarded as good and distinct species. considering all the ascertained facts on the intercrossing of plants and animals. produce offspring. species which are so different that they are generally ranked in distinct genera. therefore. but from different hatches. torquatus. that whole flocks of these crossed geese are kept in various parts of the country. LAWS GOVERNING THE STERILITY OF FIRST CROSSES AND OF HYBRIDS. both in first crosses and in hybrids. We will now consider a little more in detail the laws governing the sterility of first crosses and of hybrids. which was first propounded by Pallas. M.

as we have seen. beyond that which the plant's own pollen produces. by the pollen of one of the pure parent-species causing the flower of the hybrid to wither earlier than it otherwise would have done. and. and the early withering of the flower is well known to be a sign of incipient fertilisation. The fertility. for instance in Dianthus. are generally very sterile. From this extreme degree of sterility we have self-fertilised hybrids producing a greater and greater number of seeds up to perfect fertility.of plants. even with the pollen of the pure parents. It is surprising in how many curious ways this gradation can be shown. The hybrids raised from two species which are very difficult to cross. and on the other hand. it depends in part upon the constitution of the individuals which happen to have been chosen for the experiment. So in hybrids themselves. and probably never would produce. and the sterility of the hybrids thus produced--two classes of facts which are generally confounded together--is by no means strict. can be united with unusual facility. Now the fertility of first crosses. and which rarely produce any offspring. This is clearly shown by hybrids never having been raised between species ranked by systematists in distinct families. in which two pure species. yet these hybrids are remarkably sterile. considering how scanty our knowledge is in regard to hybrid animals. than is that of pure species. both of first crosses and of hybrids. I have been surprised to find how generally the same rules apply to both kingdoms. Even within the limits of the same genus. is more easily affected by unfavourable conditions. even to an excess of fertility. and produce numerous hybrid offspring. when at last produced. for it is not always the same in degree when the same two species are crossed under the same circumstances. up to nearly complete or even quite complete fertility. the pollen of different species applied to the stigma of some one species of the same genus. So it is with hybrids. It has been already remarked. and of the hybrids produced from them. From this absolute zero of fertility. and. is largely governed by their systematic affinity. On the other hand. are very fertile. graduates from zero to perfect fertility. I have taken much pains to ascertain how far they apply to animals. a single fertile seed: but in some of these cases a first trace of fertility may be detected. by very closely allied species generally uniting with facility. these two opposite cases occur. or only with extreme difficulty. yields a perfect gradation in the number of seeds produced. there are some which never have produced. There are many cases. as in the genus Verbascum. for their degree of fertility is often found to differ greatly in the several individuals raised from seed out of the same capsule and exposed to the same conditions. both of first crosses and of hybrids. the general resemblance between species in structure and constitution. it exerts no more influence than so much inorganic dust. but only the barest outline of the facts can here be given. or with extreme difficulty. in certain abnormal cases. But the fertility of first crosses is likewise innately variable. and on the other hand . there are species which can be crossed very rarely. But the correspondence between systematic affinity and the facility of crossing is by no means strict. When pollen from a plant of one family is placed on the stigma of a plant of a distinct family. but the hybrids. that the degree of fertility. A multitude of cases could be given of very closely allied species which will not unite. By the term systematic affinity is meant. but the parallelism between the difficulty of making a first cross.

moreover. and such hybrids. an intermediate character between their two parents. He has observed it even between closely related forms (as Matthiola annua and glabra) which many botanists rank only as varieties. There are certain hybrids which. though they rarely differ in external characters. we meet with this same difference. though of course compounded of the very same two species. yet generally differ in fertility in a small. Gartner. exceptional and abnormal individuals sometimes are born. There is often the widest possible difference in the facility of making reciprocal crosses. deciduous and evergreen trees. even when the other hybrids raised from seed from the same capsule have . which is not a particularly distinct species. plants inhabiting different stations and fitted for extremely different climates. and the hybrids thus produced are sufficiently fertile. for they prove that the capacity in any two species to cross is often completely independent of their systematic affinity. jalapa. longiflora. obstinately failed to fertilise. So again among hybrids which are usually intermediate in structure between their parents. longiflora with the pollen of M. Thuret has observed the same fact with certain sea-weeds or Fuci. as Silene. and occasionally in a high degree. Several other singular rules could be given from Gartner: for instance. No one has been able to point out what kind or what amount of difference. excepting in their reproductive systems. can be crossed. The diversity of the result in reciprocal crosses between the same two species was long ago observed by Kolreuter. which closely resemble one of their pure parents. by no less than eight other species of Nicotiana. for instance. can often be crossed with ease. some species have a remarkable power of crossing with other species. to fertilise reciprocally M. In the same family there may be a genus. but Gartner found that N. found that this difference of facility in making reciprocal crosses is extremely common in a lesser degree. as Dianthus. and then a mare by a male-ass: these two species may then be said to have been reciprocally crossed. even in the pollen. though externally so like one of their pure parent-species. Such cases are highly important. I mean the case. other species of the same genus have a remarkable power of impressing their likeness on their hybrid offspring. By a reciprocal cross between two species. is sufficient to prevent two species crossing. the one species having first been used as the father and then as the mother. and in the cotyledons. always closely resemble one of them. in which very many species can most readily be crossed. during eight following years. in which the most persevering efforts have failed to produce between extremely close species a single hybrid. of a female-ass being first crossed by a stallion. but these two powers do not at all necessarily go together. or to be fertilised. It can be shown that plants most widely different in habit and general appearance. To give an instance: Mirabilis jalapa can easily be fertilised by the pollen of M. Many analogous facts could be given. Annual and perennial plants. instead of having. It is also a remarkable fact that hybrids raised from reciprocal crosses. acuminata. and another genus. in the fruit. are with rare exceptions extremely sterile. but Kolreuter tried more than two hundred times. Even within the limits of the same genus. and having strongly marked differences in every part of the flower. and these hybrids are almost always utterly sterile.of very distinct species which unite with the utmost facility. and utterly failed. as is usual. Several other equally striking cases could be given. in any recognisable character. that is of any difference in their structure or constitution. the many species of Nicotiana have been more largely crossed than the species of almost any other genus. for instance.

but not in a reversed direction. in the period of the flow or nature of their sap. This latter statement is clearly proved by the difference in the result of reciprocal crosses between the same two species. Considering the several rules now given. that their fertility. and occasionally the widest possible difference. and then to stop their further propagation by different degrees of sterility. for. These facts show how completely the fertility of a hybrid may be independent of its external resemblance to either pure parent. we see that when forms. when various species are crossed. their fertility graduates from zero to perfect fertility.. on the other hand. there is generally some difference. We can sometimes see the reason why one tree will not take on another from differences in their rate of growth. It will be advisable to explain a little more fully. the differences being of so peculiar and limited a nature. The hybrids. but will admit that it is incidental on differences in the laws of growth of the two plants. and other species cross with extreme difficulty. the male sexual element of the one will often freely act on the female sexual element of the other. that it is by no means always the same in degree in the first cross and in the hybrids produced from this cross. does not always prevent the two . that the facility of making a first cross between any two species is not always governed by their systematic affinity or degree of resemblance to each other. that the fertility of hybrids is not related to the degree in which they resemble in external appearance either parent. it may even be asked. moreover. which govern the fertility of first crosses and of hybrids. in the hardness of their wood. and lastly. Now do these complex and singular rules indicate that species have been endowed with sterility simply to prevent their becoming confounded in nature? I think not. As the capacity of one plant to be grafted or budded on another is unimportant for their welfare in a state of nature. according as the one species or the other is used as the father or the mother. is simply incidental or dependent on unknown differences in their reproductive systems. For why should the sterility be so extremely different in degree. both of first crosses and of hybrids. what I mean by sterility being incidental on other differences. etc. or even to fertility under certain conditions in excess. has the production of hybrids been permitted? To grant to species the special power of producing hybrids. which must be considered as good and distinct species. is innately variable. seems a strange arrangement. are united. one being woody and the other herbaceous. and adaptation to widely different climates. that. The foregoing rules and facts. besides being eminently susceptible to favourable and unfavourable conditions. by an example. Great diversity in the size of two plants. one being evergreen and the other deciduous. appear to me clearly to indicate that the sterility. all of which we must suppose it would be equally important to keep from blending together? Why should the degree of sterility be innately variable in the individuals of the same species? Why should some species cross with facility and yet produce very sterile hybrids. and yet produce fairly fertile hybrids? Why should there often be so great a difference in the result of a reciprocal cross between the same two species? Why. but in a multitude of cases we can assign no reason whatever. in reciprocal crosses between the same two species. produced from reciprocal crosses often differ in fertility. not strictly related to the facility of the first union between their parents.a considerable degree of fertility. and not a specially endowed quality. I presume that no one will suppose that this capacity is a SPECIALLY endowed quality. in the facility of effecting an union.

when thus grafted were rendered barren. so Sagaret believes this to be the case with different individuals of the same two species in being grafted together. yet that there is a rude degree of parallelism in the results of grafting and of crossing distinct species. although there is a clear and great difference between the mere adhesion of grafted stocks and the union of the male and female elements in the act of reproduction. As Gartner found that there was sometimes an innate difference in different INDIVIDUALS of the same two species in crossing. and which could be grafted with no great difficulty on a fourth species. is a different case from the difficulty of uniting two pure species. can usually. so do different varieties of the apricot and peach on certain varieties of the plum. the difficulty is as important for the endurance and stability of specific forms as in the case of grafting it is unimportant for their welfare. which seed much more freely when fertilised with the pollen of a distinct species than when fertilised with pollen from the same plant. On the other hand. for instance. and. for no one has been able to graft together trees belonging to quite distinct families. which seeded freely on their own roots. The facts by no means seem to indicate that the greater or lesser difficulty of either grafting or crossing various species has been a special endowment. than on the apple. the capacity is limited by systematic affinity. Although many distinct genera within the same family have been grafted together. though with difficulty. etc. closely allied species and varieties of the same species. We have seen that the sterility of hybrids which have their reproductive organs in an imperfect condition.. although in the case of crossing. is by no means absolutely governed by systematic affinity. in other cases species of the same genus will not take on each other. to a certain extent. on the gooseberry. We thus see that. so with grafting. At one time it appeared to me probable. that the sterility of first crosses and of hybrids might have been slowly acquired through the natural selection of slightly lessened degrees of . for Thouin found that three species of Robinia. the facility of effecting an union is often very far from equal. But this capacity. on the other hand. yet these two distinct classes of cases run to a large extent parallel. cannot be grafted on the currant. as it has to others. when grafted on other species. which have their reproductive organs perfect. Even different varieties of the pear take with different degrees of facility on the quince. systematic affinity. as might have been expected. Passiflora.grafting together. which is a member of the same genus. whereas the currant will take. These differences in both cases follow. We are reminded by this latter fact of the extraordinary cases of Hippeastrum. certain species of Sorbus. yielded twice as much fruit as when on their own roots. As in reciprocal crosses. Something analogous occurs in grafting. but not invariably. ORIGIN AND CAUSES OF THE STERILITY OF FIRST CROSSES AND OF HYBRIDS. so I believe that the still more complex laws governing the facility of first crosses are incidental on unknown differences in their reproductive systems. be grafted with ease. which is ranked as a distinct genus. And as we must look at the curious and complex laws governing the facility with which trees can be grafted on each other as incidental on unknown differences in their vegetative systems. as in hybridisation. The common gooseberry. The pear can be grafted far more readily on the quince. As in hybridisation. by which term every kind of resemblance and dissimilarity between organic beings is attempted to be expressed. so it sometimes is in grafting.

thus leading to their preservation. that. if it were rendered in some slight degree sterile when crossed with its parent form or with some other variety. it may be remarked that species inhabiting distinct regions are often sterile when crossed. Both Gartner and Kolreuter have proved that in genera including numerous species. now it could clearly have been of no advantage to such separated species to have been rendered mutually sterile. from slightly lessened fertility to absolute sterility. while at the same time the male element of this second form is enabled freely to fertilise the first form. But he who will take the trouble to reflect on the steps by which this first degree of sterility could be increased through natural selection to that high degree which is common with so many species. must have incessantly occurred with many species. After mature reflection. like any other variation. Take the case of any two species which. on the same principle that. now. in rendering species mutually sterile. would not thus itself gain any advantage or indirectly give any advantage to the other individuals of the same variety. when man is selecting at the same time two varieties. if the theory of natural selection be brought to bear. and which thus approached by one small step towards absolute sterility? Yet an advance of this kind. In the first place. But it would be superfluous to discuss this question in detail: for with plants we have conclusive evidence that the sterility of crossed species must be due to some principle. the greatest difficulty will be found to lie in the existence of many graduated steps. for a multitude are mutually quite barren.fertility. if a species was rendered sterile with some one compatriot. will find the subject extraordinarily complex. it seems to me that this could not have been effected through natural selection. when crossed. With sterile neuter insects we have reason to believe that modifications in their structure and fertility have been slowly accumulated by natural selection. to species which never produce a single seed. a series can be formed from species which when crossed yield fewer and fewer seeds. but yet are affected by the pollen of certain other species. if rendered slightly sterile when crossed with some other variety. it is almost as much opposed to the theory of natural selection as to that of special creation. that in reciprocal crosses the male element of one form should have been rendered utterly impotent on a second form. which. from an advantage having been thus indirectly given to the community to which they belonged over other communities of the same species. but an individual animal not belonging to a social community. In considering the probability of natural selection having come into action. quite independent of natural selection. produced few and sterile offspring. what is there which could favour the survival of those individuals which happened to be endowed in a slightly higher degree with mutual infertility. and consequently this could not have been effected through natural selection. spontaneously appeared in certain individuals of one variety when crossed with those of another variety. For it would clearly be advantageous to two varieties or incipient species if they could be kept from blending. it is necessary that he should keep them separate. and which is universal with species which have been differentiated to a generic or family rank. sterility with other species would follow as a necessary contingency. for the germen swells. It may be admitted that it would profit an incipient species. In the second place. for thus fewer bastardised and deteriorated offspring would be produced to commingle their blood with the new species in process of formation. It is here manifestly impossible to select the more sterile . but it may perhaps be argued. for this peculiar state of the reproductive system could hardly have been advantageous to either species.

but I believe. Until becoming acquainted with these facts. but the young chickens had been unable to break through the shell. cannot have been gained through selection. but be incapable of causing an embryo to be developed. it may therefore. I was unwilling to believe in the frequent early death of hybrid embryos. before birth. But a hybrid partakes of only half of the nature and constitution of its mother. who has had great experience in hybridising pheasants and fowls. as we see in the case of the common mule. however. and in the majority of the fertilised eggs. causing the embryo to be imperfectly developed. an embryo may be developed. the male element may reach the female element. the embryos within the eggs of silk moths which had not been fertilised. "without any obvious cause. Hewitt. This latter alternative has not been sufficiently attended to. In the case of first crosses. when the germen alone is effected. as would be the case with a plant having a pistil too long for the pollen-tubes to reach the ovarium. that the early death of the embryo is a very frequent cause of sterility in first crosses. Of the chickens which were born. Mr. There must sometimes be a physical impossibility in the male element reaching the ovule. No explanation can be given of these facts. or within the egg or seed produced by the mother. . be exposed to conditions in some degree unsuitable. But after all. at least it is known that hybrids raised from very distinct species are sometimes weak and dwarfed. so that this acme of sterility. the greater or less difficulty in effecting a union and in obtaining offspring apparently depends on several distinct causes. we may infer that the cause. though the pollen-tubes protrude. more than four-fifths died within the first few days. when once born. or at latest weeks. are differently circumstanced before and after birth: when born and living in a country where their two parents live. of which fact Max Wichura has recently given some striking cases with hybrid willows. whatever it may be. as long as it is nourished within its mother's womb. rather than in the conditions to which it is subsequently exposed. It has also been observed that when the pollen of one species is placed on the stigma of a distantly allied species. are generally healthy and long-lived. more especially as all very young beings are eminently sensitive to injurious or unnatural conditions of life. We will now look a little closer at the probable nature of the differences between species which induce sterility in first crosses and in hybrids. any more than why certain trees cannot be grafted on others.individuals. and perish at an early age. or had become nearly mature. is the same or nearly the same in all cases. the majority of these eggs had been fertilised. and then perish at an early period." so that from the 500 eggs only twelve chickens were reared. they do not penetrate the stigmatic surface. as seems to have been the case with some of Thuret's experiments on Fuci. apparently from mere inability to live. Hybrids. With plants. the cause more probably lies in some imperfection in the original act of impregnation. for hybrids. they are generally placed under suitable conditions of life. Lastly. It may be here worth noticing that in some cases of parthenogenesis. the embryos had either been partially developed and had then perished. and consequently be liable to perish at an early period. which have already ceased to yield seeds. hybridized embryos probably often perish in a like manner. and from the laws governing the various grades of sterility being so uniform throughout the animal and vegetable kingdoms. Salter has recently given the results of an examination of about 500 eggs produced from various crosses between three species of Gallus and their hybrids. pass through their early stages of development and then perish like the embryos produced by a cross between distinct species. Again. from observations communicated to me by Mr.

when animals and plants are removed from their natural conditions. the unequal fertility of hybrids produced from reciprocal crosses. Nor do I pretend that the foregoing remarks go to the root of the matter: no explanation is offered why an organism. or periodical action. the case is somewhat different. All that I have attempted to show is. in some respects allied. or any exotic plant seed freely under culture. they are extremely liable to vary. though in a lesser degree than when sterility ensues. In the one case. in the other . several facts with respect to the sterility of hybrids. When hybrids are able to breed inter se. For it is scarcely possible that two organisations should be compounded into one. and is often accompanied by excess of size or great luxuriance. and certain species in a group will produce unusually fertile hybrids. No one can tell till he tries.In regard to the sterility of hybrids. for their offspring in successive generations are eminently liable to vary. On the other hand. but sometimes the female more than the male. So it is with hybrids. nor can he tell till he tries. whether any two species of a genus will produce more or less sterile hybrids. they are extremely liable to have their reproductive systems seriously affected. in fact. in which the sexual elements are imperfectly developed. does not diminish. the reproductive system. having been blended into one. of too close interbreeding. however. or that of hybrids. Lastly. be owned that we cannot understand. the male element is the most liable to be affected. they transmit to their offspring from generation to generation the same compounded organisation. though often in so slight a degree as to be inappreciable by us. which seems to be partly due to their reproductive systems having been specially affected. the conditions of life have been disturbed. there are many points of similarity. when placed under unnatural conditions. is the great bar to the domestication of animals. in both. including of course the reproductive systems. this being generally the result. is rendered sterile. and when hybrids are produced by the unnatural crossing of two species. for instance. in the other case. that in two cases. the tendency goes to a certain extent with systematic affinity. is affected in a very similar manner. though in some degree variable. or mutual relations of the different parts and organs one to another or to the conditions of life. I have more than once alluded to a large body of facts showing that. the external conditions have remained the same. It must. as every experimentalist has observed. but the organisation has been disturbed by two distinct structures and constitutions. In both cases the sterility occurs in various degrees. In both. without some disturbance occurring in the development. Between the sterility thus superinduced and that of hybrids. and whole groups of species tend to produce sterile hybrids. This. In both cases the sterility is independent of general health. independently of the general state of health. and hence we need not be surprised that their sterility. Thus we see that when organic beings are placed under new and unnatural conditions. or the increased sterility in those hybrids which occasionally and exceptionally resemble closely either pure parent. on the above or any other view. when organic beings are placed during several generations under conditions not natural to them. whether any particular animal will breed under confinement. it is even apt to increase. The above view of the sterility of hybrids being caused by two constitutions being compounded into one has been strongly maintained by Max Wichura. sterility is the common result--in the one case from the conditions of life having been disturbed. one species in a group will sometimes resist great changes of conditions with unimpaired fertility. for whole groups of animals and plants are rendered impotent by the same unnatural conditions. as before explained.

from one soil or climate to another. and on the other hand. It is an old and almost universal belief. These organs are so proportioned in length . the three forms possess altogether six sets of stamens and three kinds of pistils. when subjected. and a multitude of other animals. which would probably have been sterile if aboriginally crossed. as we have seen. The above two parallel series of facts seem to be connected together by some common but unknown bond. produce hybrids which are almost always in some degree sterile. which have been subjected to slightly different conditions. and that close interbreeding continued during several generations between the nearest relations. will be able to explain the primary cause of hybrids being so generally sterile. Hence it seems that. RECIPROCAL DIMORPHISM AND TRIMORPHISM. great benefit is derived from almost any change in their habits of life. which I have elsewhere given. which differ to a certain extent.case from the organisation having been disturbed by two organisations being compounded into one. to a considerable change in their conditions. founded on a considerable body of evidence. very frequently are rendered more or less sterile. I am fully persuaded that this double parallelism is by no means an accident or an illusion. But. the two having differently sized pollen-grains. He who is able to explain why the elephant. that slight changes in the conditions of life are beneficial to all living things. During the convalescence of animals. organic beings long habituated to certain uniform conditions under a state of nature. there is the clearest evidence that a cross between individuals of the same species. one form having a long pistil with short stamens. and when this tendency is slightly disturbed by any change.. gives vigour and fertility to the offspring. With trimorphic plants there are three forms likewise differing in the lengths of their pistils and stamens. are always tending towards an equilibrium. or consists in. and we know that a cross between two forms that have become widely or specifically different. tubers. the other a short pistil with long stamens. that is. almost always leads to decreased size. or which have slightly varied. that slight crosses. slight changes in the conditions of life benefit all organic beings. weakness. are quite fertile together. which is essentially related to the principle of life. crosses between the males and females of the same species. in the size and colour of the pollen-grains. Herbert Spencer. etc. which exist in about equal numbers and which differ in no respect except in their reproductive organs. He will at the same time be able to explain how it is that the races of some of our domesticated animals. which have often been subjected to new and not uniform conditions. if these be kept under the same conditions of life. this principle. which. according to Mr. on the one hand. Several plants belonging to distinct orders present two forms. A similar parallelism holds good with an allied yet very different class of facts. and in some other respects. give vigour and fertility to the offspring. or sterility. are incapable of breeding when kept under only partial confinement in their native country. This subject may be here briefly discussed. both with plants and animals. and will be found to throw some light on hybridism. as throughout nature. the vital forces gain in power. being that life depends on. the incessant action and reaction of various forces. although they are descended from distinct species. We see this acted on by farmers and gardeners in their frequent exchanges of seed. and back again. as under confinement. Again. and as in each of the three forms there are two sets of stamens.

It is well known that if pollen of a distinct species be placed on the stigma of a flower. As the degree of sterility in the latter case depends in an eminent degree on the conditions of life being more or less favourable. and two. and from trimorphic plants all three illegitimate forms. which may be called legitimate. and twenty-four hours afterwards legitimately. This led me carefully to observe during four years many seedlings. and the result has been confirmed by other observers. when both are placed on the same stigma. some being so utterly and incurably sterile that they did not yield during four seasons a single seed or even seed-capsule. If. its action is so strongly prepotent that it generally annihilates the effect of the foreign pollen. or more or less infertile. when illegitimately united. In all these respects. are not fully fertile. just in the same manner as occurs in crossing distinct species. but the latter form did not yield a single seed when fertilised by the longer stamens of the mid-styled form. though applied twenty-four hours subsequently. are more or less infertile. when they are illegitimately fertilised. may be strictly compared with that of hybrids when crossed inter se. I ascertained this by fertilising several flowers. are fully fertile. so it is with the pollen of the several forms of the same species. Now I have shown. had wholly destroyed or prevented the action of the previously applied illegitimate pollen. as they may be called. placed on the same stigma. and its own pollen be afterwards. this shows that the legitimate pollen. and in others which might be added. there is no apparent reason why they should not yield as many seeds as did their parents when legitimately fertilised. on the other hand. When this is done. The infertility which may be observed in various dimorphic and trimorphic plants. up to absolute and utter sterility. It is possible to raise from dimorphic species. for legitimate pollen is strongly prepotent over illegitimate pollen. The chief result is that these illegitimate plants. behave in exactly the same manner as do two distinct species when crossed. But such is not the case. both long-styled and short-styled illegitimate plants. differs much in degree. They are all infertile. first illegitimately. the forms of the same undoubted species. in various degrees. and yielded many seeds. with pollen taken from a peculiarly coloured variety. a hybrid is crossed with either pure parent-species. as in making reciprocal crosses between the same two species. So that with dimorphic species two unions. raised from several illegitimate unions. for instance. that is by pollen taken from stamens not corresponding in height with the pistil. that in order to obtain full fertility with these plants. it is necessary that the stigma of the one form should be fertilised by pollen taken from the stamens of corresponding height in another form. Again. These can then be properly united in a legitimate manner. the sterility is usually much lessened: and so it is when an illegitimate plant is fertilised by a legitimate plant. there is occasionally a great difference in the result. and all the seedlings were similarly coloured. With trimorphic species six unions are legitimate. so the same thing occurs with trimorphic plants. the mid-styled form of Lythrum salicaria was illegitimately fertilised with the greatest ease by pollen from the longer stamens of the short-styled form. when united with each other in a legitimate manner. or fully fertile. and twelve are illegitimate. The sterility of these illegitimate plants. so I have found it with illegitimate unions.to each other that half the stamens in two of the forms stand on a level with the stigma of the third form. In the same manner as the sterility of hybrids does not always run parallel with the difficulty of making the first cross between the two . which may be called illegitimate. even after a considerable interval of time.

or can be united with great difficulty. doubtful whether this is really so. and yet be sterile when united in certain ways.parent-species. first. both in first crosses and in hybrids. while the converse cross can be effected with perfect . With hybrids raised from the same seed-capsule the degree of sterility is innately variable. he would raise plants from his supposed hybridised seed. many hybrids are profuse and persistent flowerers. This will perhaps be made more fully apparent by an illustration. in which the male of one species cannot be united. so that sterility of certain illegitimate plants was unusually great. because they show us. for instance. secondly. in the ordinary unions of the individuals of the same species and with crosses between distinct species. that his two varieties were as good and as distinct species as any in the world. that the sterility of distinct species when crossed and of their hybrid progeny. which results in sterility. while the sterility of the union from which they were derived was by no means great. while ordinary hybrids are produced from an improper union between so-called distinct species. because we may conclude that there is some unknown bond which connects the infertility of illegitimate unions with that of their illegitimate offspring. that two or three forms of the same species may exist and may differ in no respect whatever. and we are led to extend the same view to first crosses and hybrids. infer as probable from the consideration of dimorphic and trimorphic plants. in accordance with the common view. is no safe criterion of specific distinction. Altogether there is the closest identity in character and behaviour between illegitimate plants and hybrids. We may. of two long-styled forms. The facts now given on dimorphic and trimorphic plants are important. depends exclusively on the nature of their sexual elements. miserable dwarfs. because we find. thirdly. relatively to external conditions. and are weak. we may suppose that a botanist found two well-marked varieties (and such occur) of the long-styled form of the trimorphic Lythrum salicaria. and this seems to me of especial importance. and he would find that the seedlings were miserably dwarfed and utterly sterile. For we must remember that it is the union of the sexual elements of individuals of the same form. and that they behaved in all the other above specified respects as if they had been two distinct species. either in structure or in constitution. He might then maintain that he had actually proved. We are also led to this same conclusion by considering reciprocal crosses. exactly similar cases occur with the illegitimate offspring of various dimorphic and trimorphic plants. Lastly. and not on any difference in their structure or general constitution. however. while other and more sterile hybrids produce few flowers. but I will not enlarge on this obscure subject. But to make the case sure. that the physiological test of lessened fertility. and that he determined to try by crossing whether they were specifically distinct. with the female of a second species. It is. and that they behaved in all other respects like ordinary hybrids. but he would be completely mistaken. He would find that they yielded only about one-fifth of the proper number of seed. It is hardly an exaggeration to maintain that illegitimate plants are hybrids. Hence the case appears at first sight exactly the reverse of what occurs. We have also already seen that there is the closest similarity in all respects between first illegitimate unions and first crosses between distinct species. produced within the limits of the same species by the improper union of certain forms. while it is the union of the sexual elements proper to two distinct forms which is fertile. so it is in a marked manner with illegitimate plants. however.

If we turn to varieties. which. NOT UNIVERSAL. which in their natural state probably would have been in some degree sterile when crossed. differing widely from each other in appearance. is that they are descended from aboriginally distinct species. that we have good grounds for admitting the directly opposite doctrine of Pallas. With some exceptions. FERTILITY OF VARIETIES WHEN CROSSED. It may be urged as an overwhelming argument that there must be some essential distinction between species and varieties inasmuch as the latter. and it can hardly be rejected. under domestication. so that the domesticated descendants of species. with domesticated animals. is a remarkable fact. . or supposed to have been produced. looking to varieties produced under nature. more especially when we reflect how many species there are. cross with perfect facility. In the first place. have had so little tendency towards modifying the reproductive system in a manner leading to mutual sterility. Several considerations. are said by Gartner to be quite sterile when crossed. That excellent observer. Now the varying conditions to which domesticated animals and cultivated plants have been subjected. we are still involved in some doubt. are utterly sterile when intercrossed. But the subject is surrounded by difficulties. that in several well-authenticated cases already alluded to. though resembling each other most closely. however. certain plants have been affected in an opposite manner. and why with plants only a few such cases. render the fertility of domestic varieties less remarkable. or of the cabbage.facility. they are at once ranked by most naturalists as species. For instance. for instance. For when it is stated. immediately to be given. and being fertilised by. varieties have not been produced which are mutually sterile. If the Pallasian doctrine of the elimination of sterility through long-continued domestication be admitted. which are considered by most botanists as varieties. produced. and he consequently ranks them as undoubted species. for. that such conditions generally eliminate this tendency. for they have become self-impotent. the explanation which will occur to everyone. have been observed. I fully admit that this is the rule. though in certain cases. however much they may differ from each other in external appearance. With plants. we can understand why. it becomes in the highest degree improbable that similar conditions long-continued should likewise induce this tendency. so that similar differences in the case of varieties would be no sure guide. If we thus argue in a circle. Thus. the fertility of all varieties produced under nature will assuredly have to be granted. so far is cultivation from giving a tendency towards sterility between distinct species. and probably the true one. become perfectly fertile together. as I believe. it may be observed that the amount of external difference between two species is no sure guide to their degree of mutual sterility. while still retaining the capacity of fertilising. presently to be given. Gartner. other species. likewise concluded that species when crossed are sterile owing to differences confined to their reproductive systems. sterility might occasionally be thus caused. and yield perfectly fertile offspring. with species having a peculiar constitution. that certain South American indigenous domestic dogs do not readily unite with European dogs. AND OF THEIR MONGREL OFFSPRING. It is certain that with species the cause lies exclusively in differences in their sexual constitution. those of the pigeon. for instance. Nevertheless the perfect fertility of so many domestic races. the blue and red pimpernel. if two forms hitherto reputed to be varieties be found in any degree sterile together. namely.

which. but only a single head produced any seed. and which can now generally resist with undiminished fertility repeated changes of conditions. The following case is far more remarkable. How far these experiments may be trusted. The evidence is at least as good as that from which we believe in the sterility of a multitude of species. Manipulation in this case could not have been injurious. but the forms experimented on are ranked by Sagaret. and it is important to notice that the hybrid plants thus raised were themselves PERFECTLY fertile. I know not. as the plants have separated sexes. that the yellow and white varieties when crossed produce less seed than the similarly coloured varieties of the same species. Moreover. and although these plants have separated sexes. and a tall variety with red seeds growing near each other in his garden. and this may well make a wide difference in the result. and seems at first incredible. when yellow and white varieties of one species are crossed with yellow and . But we can see that species. they never naturally crossed. I have as yet spoken as if the varieties of the same species were invariably fertile when intercrossed. seeing how profoundly ignorant we are in regard to the normal and abnormal action of the reproductive system. but why this has so generally occurred with natural varieties. who in all other cases consider fertility and sterility as safe criterions of specific distinction. Domesticated productions. But it is impossible to resist the evidence of the existence of a certain amount of sterility in the few following cases. Girou de Buzareingues crossed three varieties of gourd. why domestic varieties have not become mutually infertile when crossed. I believe. has suspected that these varieties of maize are distinct species. We are far from precisely knowing the cause. when taken from their natural conditions and subjected to captivity. and he asserts that their mutual fertilisation is by so much the less easy as their differences are greater. which I will briefly abstract. during several years. so that even Gartner did not venture to consider the two varieties as specifically distinct. as soon as they have been permanently modified in a sufficient degree to take rank as species. and the reproductive functions of organic beings which have always lived under natural conditions would probably in like manner be eminently sensitive to the influence of an unnatural cross. on the other hand. who mainly founds his classification by the test of infertility. The evidence is also derived from hostile witnesses. nor is this surprising. but it is the result of an astonishing number of experiments made during many years on nine species of Verbascum. For we know how commonly wild animals and plants. No one. will have been exposed during long periods of time to more uniform conditions. than have domestic varieties. which would be little liable to have their reproductive powers injuriously affected by the act of crossing with other varieties which had originated in a like manner. as shown by the mere fact of their domestication. as it appears to me. were not originally highly sensitive to changes in their conditions of life. as varieties. he asserts that. and this one head produced only five grains. which like the maize has separated sexes. might be expected to produce varieties.The real difficulty in our present subject is not. Gartner kept. He then fertilised thirteen flowers of the one kind with pollen of the other. by so good an observer and so hostile a witness as Gartner: namely. owing to their struggle for existence with numerous competitors. and Naudin has come to the same conclusion. a dwarf kind of maize with yellow seeds. are rendered sterile.

from man attending only to external characters in his domestic varieties. whose accuracy has been confirmed by every subsequent observer. namely. and although unable to confirm Gartner's results on the crossing of the distinct species. could find very few. than between those which are differently coloured. The general sterility of crossed species may safely be looked at. INDEPENDENTLY OF THEIR FERTILITY. however.white varieties of a DISTINCT species. if proved to be infertile in any degree. not as a special acquirement or endowment. and which he tested by the severest trial. would almost universally be ranked as a species. and he found their mongrel offspring perfectly fertile. But one of these five varieties. The variability. and one variety can sometimes be raised from the seed of another. Scott also has experimented on the species and varieties of Verbascum. Independently of the question of fertility. as it seems to me. HYBRIDS AND MONGRELS COMPARED. he finds that the dissimilarly coloured varieties of the same species yield fewer seeds. but as incidental on changes of an unknown nature in their sexual elements. whose strong wish it was to draw a distinct line between species and varieties. From these facts it can no longer be maintained that varieties when crossed are invariably quite fertile. and crossed with the Nicotiana glutinosa. perhaps. Gartner. and the so-called mongrel offspring of varieties. on the other hand. by reciprocal crosses. always yielded hybrids not so sterile as those which were produced from the four other varieties when crossed with N. Hence the reproductive system of this one variety must have been in some manner and in some degree modified. Gartner further admits that hybrids between very closely allied species are more variable than those from very distinct species. and I have myself seen striking instances of this fact. from these several considerations we may conclude that fertility does not constitute a fundamental distinction between varieties and species when crossed. in the successive generations of mongrels is. in the proportion of eighty-six to 100. When mongrels and the more fertile hybrids are propagated for several generations. more seed is produced by the crosses between the similarly coloured flowers. they agree most closely in many important respects. glutinosa. than the similarly coloured varieties. when crossed with a widely distinct species. From the great difficulty of ascertaining the infertility of varieties in a state of nature. except in the colour of their flowers. has proved the remarkable fact that one particular variety of the common tobacco was more fertile than the other varieties. quite unimportant differences between the so-called hybrid offspring of species. when used either as the father or mother. He experimented on five forms which are commonly reputed to be varieties. Mr. Kolreuter. greater . that in the first generation mongrels are more variable than hybrids. The most important distinction is. and this shows that the difference in the degree of variability graduates away. an extreme amount of variability in the offspring in both cases is notorious. but Gartner admits that hybrids from species which have long been cultivated are often variable in the first generation. for a supposed variety. and. I shall here discuss this subject with extreme brevity. but some few instances of both hybrids and mongrels long retaining a uniform character could be given. And. the offspring of species and of varieties when crossed may be compared in several other respects. Yet these varieties differ in no respect. and from such varieties not having been exposed for very long periods to uniform conditions of life.

For the parents of mongrels are varieties. fails under these circumstances to perform its proper function of producing offspring closely similar in all respects to the parent-form. although most closely allied to each other. These several remarks are apparently applicable to animals. But this conclusion. while Naudin. Now. but more especially owing to prepotency in transmitting likeness running more strongly in one sex than in the other. I think those authors are right who maintain that the ass has a prepotent power over . but hybrids themselves have their reproductive systems seriously affected. Such alone are the unimportant differences which Gartner is able to point out between hybrid and mongrel plants. according to Gartner the same laws. Moreover. and with animals. but this. the degrees and kinds of resemblance in mongrels and in hybrids to their respective parents. are crossed with a third species. by repeated crosses in successive generations with either parent. the hybrids do not differ much. On the other hand. For it bears on the view which I have taken of one of the causes of ordinary variability.than in hybrids. is certainly only a difference in degree. and mostly domestic varieties (very few experiments having been tried on natural varieties). more especially in hybrids produced from nearly related species. is founded on a single experiment. insists in the strongest terms on the almost universal tendency to reversion in hybrids. as far as I can make out. on the other hand. and he experimented chiefly on cultivated plants. the hybrids are widely different from each other. and their descendants are highly variable. This greater variability in mongrels than in hybrids does not seem at all surprising. namely. which would often continue and would augment that arising from the act of crossing. that the reproductive system. follow. When two species are crossed. partly owing to the existence of secondary sexual characters. if it be true. The slight variability of hybrids in the first generation. Both hybrids and mongrels can be reduced to either pure parent form. and this probably explains the singular difference in the results arrived at by different observers. and he experimented on uncultivated species of willows. and they are not variable. in contrast with that in the succeeding generations. one has sometimes a prepotent power of impressing its likeness on the hybrid. and this implies that there has been recent variability. is a curious fact and deserves attention. But to return to our comparison of mongrels and hybrids: Gartner states that mongrels are more liable than hybrids to revert to either parent form. both when one species is crossed with another and when one variety is crossed with another variety. Gartner further states that when any two species. Hybrid plants produced from a reciprocal cross generally resemble each other closely. one variety certainly often has this prepotent power over another variety. Gartner expressly states that the hybrids from long cultivated plants are more subject to reversion than hybrids from species in their natural state. Thus Max Wichura doubts whether hybrids ever revert to their parent forms. and so it is with mongrel plants from a reciprocal cross. For instance. but the subject is here much complicated. and seems directly opposed to the results of several experiments made by Kolreuter. from being eminently sensitive to changed conditions of life. hybrids in the first generation are descended from species (excluding those long cultivated) which have not had their reproductive systems in any way affected. whereas if two very distinct varieties of one species are crossed with another species. So I believe it to be with varieties of plants.

It is generally different.the horse. so that both the mule and the hinny resemble more closely the ass than the horse. but is governed by several curious and complex laws. so that the mule. and is eminently susceptible to action of favourable and unfavourable conditions. and sometimes widely different in reciprocal crosses between the same two species. SUMMARY OF CHAPTER. but closely resemble one of their parents. comes to the conclusion that the laws of resemblance of the child to its parents are the same. deficiency of tail or horns. The sterility is of all degrees. the greater or less facility of one species to unite with another is incidental on unknown differences in their reproductive systems. yet I grant much less frequently than with mongrels. but not universally. are very generally. and of crossed varieties. The degree of sterility does not strictly follow systematic affinity. namely. melanism. that it is only with mongrels that the offspring are not intermediate in character. in their vegetative systems. and which have suddenly appeared--such as albinism. On the whole. the resemblances seem chiefly confined to characters almost monstrous in their nature. . but that the prepotency runs more strongly in the male than in the female ass. so in crossing. I entirely agree with Dr. There is no more reason to think that species have been specially endowed with various degrees of sterility to prevent their crossing and blending in nature. and at varieties as having been produced by secondary laws. In the same manner as in grafting trees. than with hybrids. who. and do not relate to characters which have been slowly acquired through selection. First crosses between forms. which are descended from species slowly and naturally produced. is incidental on differences. or additional fingers and toes. this similarity would be an astonishing fact. in the union of individuals of the same variety. and their hybrids. in all other respects there seems to be a general and close similarity in the offspring of crossed species. It is not always equal in degree in a first cross and in the hybrids produced from this cross. or of different varieties. but this does sometimes occur with hybrids. Independently of the question of fertility and sterility. or of distinct species. than to think that trees have been specially endowed with various and somewhat analogous degrees of difficulty in being grafted together in order to prevent their inarching in our forests. which is an offspring of the male ass and mare. is more like an ass than is the hinny. A tendency to sudden reversions to the perfect character of either parent would. But it harmonises perfectly with the view that there is no essential distinction between species and varieties. after arranging an enormous body of facts with respect to animals. sufficiently distinct to be ranked as species. generally of an unknown nature. Looking to the cases which I have collected of cross-bred animals closely resembling one parent. If we look at species as having been specially created. Prosper Lucas. which is the offspring of the female-ass and stallion. and is often so slight that the most careful experimentalists have arrived at diametrically opposite conclusions in ranking forms by this test. sterile. The sterility is innately variable in individuals of the same species. which are descended from varieties often suddenly produced and semi-monstrous in character. Much stress has been laid by some authors on the supposed fact. whether the two parents differ little or much from each other. also. be much more likely to occur with mongrels. the capacity in one species or variety to take on another.

that. and the sterility of their hybrid offspring. Nor is it surprising that the facility of effecting a first cross. and secondly. or which have varied. perhaps render it probable that some unknown bond in all cases connects the degree of fertility of first unions with that of their offspring. or sufficiently alike to be considered as varieties. The consideration of these facts on dimorphism. and the capacity of being grafted together--though this latter capacity evidently depends on widely different circumstances--should all run. . parallel with the systematic affinity of the forms subjected to experiment. and when we remember that the greater number of varieties have been produced under domestication by the selection of mere external differences. He who will explain these latter cases will be able to explain the sterility of hybrids. invariably fertile. which have been exposed to slightly different conditions of life. that the crossing of forms. In the case of hybrids. But why. for systematic affinity includes resemblances of all kinds. and in their inheritance of characters from both parent-forms. the sterility being closely allied to that which so frequently affects pure species. It is not surprising that the difficulty in crossing any two species. leading to their mutual infertility. when exposed to new and unnatural conditions of life. slight changes in the conditions of life add to the vigour and fertility of all organic beings. even if due to distinct causes: for both depend on the amount of difference between the species which are crossed. and the fertility of the hybrids thus produced. are very generally. should in most cases correspond. The facts given on the sterility of the illegitimate unions of dimorphic and trimorphic plants and of their illegitimate progeny. and that they have not been long exposed to uniform conditions of life. we do not know. although we are as ignorant of the precise cause of the sterility of first crosses and of hybrids as we are why animals and plants removed from their natural conditions become sterile. Nor is this almost universal and perfect fertility surprising. in their variability. In the case of first crosses it seems to depend on several circumstances. Independently of the question of fertility. but it seems to stand in some close relation to species having been exposed for long periods of time to nearly uniform conditions of life. in the case of distinct species. It should also be especially kept in mind. the sexual elements should so generally have become more or less modified. as is so often stated. that long-continued domestication tends to eliminate sterility. in their power of absorbing each other by repeated crosses. it apparently depends on their whole organisation having been disturbed by being compounded from two distinct forms. vigour and fertility of their offspring. clearly leads to the conclusion that the primary cause of the sterility of crossed species is confined to differences in their sexual elements. and their mongrel offspring. firstly. in all other respects there is the closest general resemblance between hybrids and mongrels. when it is remembered how liable we are to argue in a circle with respect to varieties in a state of nature. Finally. This view is strongly supported by a parallelism of another kind: namely. but not. then.The sterility of first crosses and of their hybrid progeny has not been acquired through natural selection. to a certain extent. as well as of the results of reciprocal crosses. and is therefore little likely to induce this same quality. yet the facts given in this chapter do not seem to me opposed to the belief that species aboriginally existed as varieties. favours the size. First crosses between forms known to be varieties. in some instances in chief part on the early death of the embryo.

when looking at any two species. however. that the really governing conditions of life do not graduate away quite insensibly like heat or moisture. it should always be borne in mind what sort of intermediate forms must. of innumerable intermediate links not now occurring everywhere throughout nature depends. One. as I believe. namely. on an extensive and continuous area with graduated physical conditions. is a very obvious difficulty. will generally be beaten out and exterminated during the course of further modification and improvement. combining a tail somewhat expanded with a crop somewhat enlarged. if we possessed all the intermediate varieties which have ever existed. The main cause. be truly enormous. in the extreme imperfection of the geological record. through which new varieties continually take the places of and supplant their parent-forms. ON THE IMPERFECTION OF THE GEOLOGICAL RECORD. as inferred from the rate of denudation and of deposition number--On the lapse of time as estimated by years--On the poorness of our palaeontological collections--On the intermittence of geological formations--On the denudation of granitic areas--On the absence of intermediate varieties in any one formation--On the sudden appearance of groups of species--On their sudden appearance in the lowest known fossiliferous strata--Antiquity of the habitable earth. to show that intermediate varieties. have formerly existed. Most of them have now been discussed. moreover. the characteristic features of these two breeds. but we should have no varieties directly intermediate between the fantail and pouter. On the absence of intermediate varieties at the present day--On the nature of extinct intermediate varieties. therefore. Why then is not every geological formation and every stratum full of such intermediate links? Geology assuredly does not reveal any such finely graduated organic chain. that the life of each species depends in a more important manner on the presence of other already defined organic forms. also. namely. These two breeds. on their number--On the lapse of time. for instance. I have found it difficult. which have formerly existed. that. I endeavoured to show. In the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume. But this is a wholly false view. we should always look for forms intermediate between each species and a common but unknown progenitor. and the progenitor will generally have differed in some respects from all its modified descendants. To give a simple illustration: the fantail and pouter pigeons are both descended from the rock-pigeon. perhaps. on the theory. none. if we had no historical or indirect evidence regarding their origin. But just in proportion as this process of extermination has acted on an enormous scale. than on climate. and this. on the very process of natural selection. I endeavoured. I assigned reasons why such links do not commonly occur at the present day under the circumstances apparently most favourable for their presence.CHAPTER X. to avoid picturing to myself forms DIRECTLY intermediate between them. have become so much modified. The explanation lies. we should have an extremely close series between both and the rock-pigeon. it would . and. so must the number of intermediate varieties. from existing in lesser numbers than the forms which they connect. is the most obvious and serious objection which can be urged against my theory. In the first place. the distinctness of specific forms and their not being blended together by innumerable transitional links.

between child and parent. livia. by differences not greater than we see between the natural and domestic varieties of the same species at the present day. we have no reason to suppose that links directly intermediate between them ever existed. by the theory. By the theory of natural selection all living species have been connected with the parent-species of each genus. oenas. But assuredly. It is just possible. the extent and thickness of our sedimentary formations are the result and the measure of the denudation which the earth's crust has elsewhere undergone. C. for instance. So with natural species. in all such cases. and these parent-species. for in all cases the new and improved forms of life tend to supplant the old and unimproved forms. As Lyell has well remarked. It is hardly possible for me to recall to the reader who is not a practical geologist. But such a case would imply that one form had remained for a very long period unaltered. and in this case DIRECT intermediate links will have existed between them. or even of each stratum. Therefore a man should examine for himself the . and to mark how each author attempts to give an inadequate idea of the duration of each formation. So that the number of intermediate and transitional links. which the future historian will recognise as having produced a revolution in natural science. may at once close this volume. He who can read Sir Charles Lyell's grand work on the Principles of Geology. and learning how deeply the surface of the land has been denuded. unless at the same time we had a nearly perfect chain of the intermediate links. even perhaps more than they differ from each other. and yet does not admit how vast have been the past periods of time. if this theory be true. must have been inconceivably great. all changes having been effected slowly. Not that it suffices to study the Principles of Geology.not have been possible to have determined from a mere comparison of their structure with that of the rock-pigeon. such as C. ON THE LAPSE OF TIME. for instance to the horse and tapir. and how much sediment has been deposited. it may be objected that time cannot have sufficed for so great an amount of organic change. a horse from a tapir. We can best gain some idea of past time by knowing the agencies at work. whilst its descendants had undergone a vast amount of change. now generally extinct. always converging to the common ancestor of each great class. such have lived upon the earth. and so on backwards. will render this a very rare event. even if we closely compared the structure of the parent with that of its modified descendants. between all living and extinct species. but between each and an unknown common parent. and the principle of competition between organism and organism. the facts leading the mind feebly to comprehend the lapse of time. we should be unable to recognise the parent-form of any two or more species. AS INFERRED FROM THE RATE OF DEPOSITION AND EXTENT OF DENUDATION. whether they had descended from this species or from some other allied species. have in their turn been similarly connected with more ancient forms. if we look to forms very distinct. Independently of our not finding fossil remains of such infinitely numerous connecting links. Hence. The common parent will have had in its whole organisation much general resemblance to the tapir and to the horse. but in some points of structure may have differed considerably from both. or to read special treatises by different observers on separate formations. that one of two living forms might have descended from the other.

while our sea-cliffs are everywhere formed by the intersection of various formations. it is good. The tides in most cases reach the cliffs only for a short time twice a day. cannot have been thus formed. by the wind. I remember having been much struck when viewing volcanic islands. have produced great results. especially in arid districts. and the observation is a most striking one. for the gentle slope of the lava-streams. or mud. for each line is composed of one and the same formation. when rapid deepen their channels. On a rainy day. But how often do we see along the bases of retreating cliffs rounded boulders. if we follow for a few miles any line of rocky cliff. which formerly were looked at as ancient sea-coasts. have to be worn away atom by atom. At last the base of the cliff is undermined. we are compelled to admit that the escarpments owe their origin in chief part to the rocks of which they are composed. we find that it is only here and there. the monuments of which we see all around us. and these remaining fixed.great piles of superimposed strata. This being the case. on the one hand. showed at a glance how . more forcibly than the conviction thus gained that subaerial agencies. huge fragments fall down. sand. it is then transported by the streams and rivers. and the waves eat into them only when they are charged with sand or pebbles. due to their formerly liquid state. along a short length or round a promontory. that the cliffs are at the present time suffering. Messrs. When thus impressed with the slow rate at which the land is worn away through subaerial and littoral action. this surface consequently has been gradually lowered. to consider. when formed of moderately hard rocks. the disintegrated matter is carried down even gentle slopes during heavy rain. in the van of many excellent observers--of Jukes. and the waves wearing away the sea-cliffs. and then they are more quickly ground into pebbles. in order to comprehend something about the duration of past time. even in a gently undulating country. and triturate the fragments. and which seem to work so slowly. and on the other hand the thickness of our sedimentary formations. all thickly clothed by marine productions. Geikie. We have. however. Nothing impresses the mind with the vast duration of time. which is undergoing degradation. showing how little they are abraded and how seldom they are rolled about! Moreover. The appearance of the surface and the vegetation show that elsewhere years have elapsed since the waters washed their base. according to our ideas of time. and watch the rivulets bringing down mud. in order to appreciate the past duration of time. recently learned from the observations of Ramsay. we see the effects of subaerial degradation in the muddy rills which flow down every slope. for there is good evidence that pure water effects nothing in wearing away rock. until after being reduced in size they can be rolled about by the waves. having resisted subaerial denudation better than the surrounding surface. which have been worn by the waves and pared all round into perpendicular cliffs of one or two thousand feet in height. Ramsay and Whitaker have shown. It is good to wander along the coast. which. with its dissolved carbonic acid. The whole surface of the land is exposed to the chemical action of the air and of the rainwater. and to a greater extent than might be supposed. the masses of rock which have been removed over many extensive areas. and in colder countries to frost. or the power of the waves. which apparently have so little power. with the lines of harder rock left projecting. that subaerial degradation is a much more important agency than coast-action. that the great lines of escarpment in the Wealden district and those ranging across England. and mark the process of degradation. Croll and others.

that is... which are represented in England by thin beds. and although conglomerates have probably been accumulated at a quicker rate than finer sediments.. would thus be removed from the mean level of the whole area in the course of six million years. Mr. I estimated one mass of conglomerate at ten thousand feet.. Mr. are thousands of feet in thickness on the Continent.000 feet of solid rock." but in estimating them by years. Professor Ramsay has published an account of a downthrow in Anglesea of 2.190 Tertiary strata. This seems an astonishing result. The same story is told still more plainly by faults--those great cracks along which the strata have been upheaved on one side. each of which bears the stamp of time..... or thrown down on the other. the pile of rocks on either side of the crack having been smoothly swept away... but if halved or .. and it makes no great difference whether the upheaval was sudden. relatively to their areas of drainage....000 feet.154 Secondary strata. and the figures are at once pronounced too small. that 1.... and then at the figures representing several million years.... Some of these formations.. they are good to show how slowly the mass must have been heaped together. and some considerations lead to the suspicion that it may be too large. as most geologists now believe.240 --making altogether 72.2. Nevertheless this impression is partly false. in all parts of the world the piles of sedimentary strata are of wonderful thickness.. as it became gradually disintegrated... by calculating the known amount of sediment annually brought down by certain rivers.. and he informs me that he fully believes that there is one in Merionethshire of 12.... rocky beds had once extended into the open ocean.. Moreover.......13.far the hard. was slow and effected by many starts. to the height or depth of thousands of feet. the two produce a totally different effect on the mind. or. The Craven fault. remarks that we do not err "in forming too great a conception of the length of geological periods. for instance.. yet from being formed of worn and rounded pebbles... extends for upward of thirty miles.... The consideration of these various facts impresses the mind almost in the same manner as does the vain endeavour to grapple with the idea of eternity. Professor Ramsay has given me the maximum thickness. Croll shows..... and along this line the vertical displacement of the strata varies from 600 to 3.........57. When geologists look at large and complicated phenomena.000 feet. yet in these cases there is nothing on the surface of the land to show such prodigious movements... in the opinion of most geologists.584 feet. blank periods of enormous length.. between each successive formation we have. So that the lofty pile of sedimentary rocks in Britain gives but an inadequate idea of the time which has elapsed during their accumulation. very nearly thirteen and three-quarters British miles. In regard to subaerial denudation.. In the Cordillera. for since the crust cracked.. and this is the result:-Feet Palaeozoic strata (not including igneous beds).. Croll.. of the successive formations in DIFFERENT parts of Great Britain. from actual measurement in most cases.300 feet. in an interesting paper. On the other hand.... the surface of the land has been so completely planed down that no trace of these vast dislocations is externally visible...

however. according to the standard of years. how long a period it takes to modify a species. due to the occurrence of physical changes of some kind. or through the immigration of new forms. namely. Unfortunately we have no means of determining. that they have formed what well deserves to be called a new sub-breed. if in sand or gravel. should never be forgotten. as the important discoveries made every year in Europe prove. Some of the many . variations or individual differences of the right nature. that new places in the polity of nature do not occur until after long intervals. after an immense interval of time. represented as it is by a measure utterly insignificant in a hall of the above dimensions. and within the same country only a few change at the same time. Throughout an enormously large proportion of the ocean. Now let us turn to our richest museums. by which some of the inhabitants might be better fitted to their new places under the altered circumstance. various breeds have been sensibly changed in the course of two or three centuries. We probably take a quite erroneous view. This tenth of an inch will represent one hundred years. Few men have attended with due care to any one strain for more than half a century. No organism wholly soft can be preserved. probably change much more slowly. which propagate their kind much more slowly than most of the lower animals. and no part with sufficient care. but by this process of unconscious selection. But let it be borne in mind. seem explicable only on the view of the bottom of the sea not rarely lying for ages in an unaltered condition. that is.quartered it is still very surprising. but to the subject of time we must return. will. however. Several eminent breeders. when the beds are upraised. eighty-three feet four inches in length. generally be dissolved by the percolation of rain water charged with carbonic acid. the preservation of the most useful or beautiful animals. This slowness follows from all the inhabitants of the same country being already so well adapted to each other. Few of us. Species. where sediment is not accumulating. Croll gives the following illustration: Take a narrow strip of paper. Moreover. so that a hundred years represents the work of two breeders in succession. Edward Forbes. would not always occur at once. The remark of that admirable palaeontologist. when we assume that sediment is being deposited over nearly the whole bed of the sea. ON THE POORNESS OF PALAEONTOLOGICAL COLLECTIONS. by another and later formation. know what a million really means: Mr. It is not to be supposed that species in a state of nature ever change so quickly as domestic animals under the guidance of methodical selection. the bright blue tint of the water bespeaks its purity. Only a small portion of the surface of the earth has been geologically explored. and stretch it along the wall of a large hall. what a hundred years implies. in relation to the subject of this work. or from a few specimens collected on some one spot. The comparison would be in every way fairer with the effects which follow from unconscious selection. then mark off at one end the tenth of an inch. that very many fossil species are known and named from single and often broken specimens. and what a paltry display we behold! That our collections are imperfect is admitted by every one. at a rate sufficiently quick to embed and preserve fossil remains. during a single lifetime. without the underlying bed having suffered in the interval any wear and tear. with no intention of modifying the breed. The many cases on record of a formation conformably covered. The remains which do become embedded. and the entire strip a million years. have so largely modified some of the higher animals. Shells and bones decay and disappear when left on the bottom of the sea.

far better than pages of detail. without our being able to assign any reason: one of the most striking instances is that of the Flysch formation. until recently not a land-shell was known belonging to either of these vast periods. in every separate territory. will bring home the truth. that is. Nor is their rarity surprising. many great deposits. charged with new and peculiar forms of life. When we see the formations tabulated in written works. requiring a vast length of time for their accumulation. see why the geological formations of each region are almost invariably intermittent. great piles of sediment. but now land-shells have been found in the lias. Lyell and Dr. would never have suspected that during the periods which were blank and barren in his own country. which have been upraised . hardly any idea can be formed of the length of time which has elapsed between the consecutive formations. had elsewhere been accumulated. and although this great mass has been most carefully searched. are entirely destitute of organic remains. Lastly. But the imperfection in the geological record largely results from another and more important cause than any of the foregoing. like E.kinds of animals which live on the beach between high and low water mark seem to be rarely preserved. entirely disbelieve in the change of species. Murchison's great work on Russia. no fossils. I think. what wide gaps there are in that country between the superimposed formations. In regard to mammiferous remains. We can. with the exception of a single Mediterranean species. and that not a cave or true lacustrine bed is known belonging to the age of our secondary or palaeozoic formations. it is difficult to avoid believing that they are closely consecutive. except a few vegetable remains. we may infer that this could nowhere be ascertained. whence the sediment was derived. a glance at the historical table published in Lyell's Manual. which inhabits deep water and this has been found fossil in Sicily. With respect to the terrestrial productions which lived during the Secondary and Palaeozoic periods. have not followed each other in close sequence. But we know. from Sir R. accord with the belief of vast intervals of time having elapsed between each formation. generally implying great changes in the geography of the surrounding lands. it is superfluous to state that our evidence is fragmentary in an extreme degree. occasionally even six thousand feet in thickness. from the several formations being separated from each other by wide intervals of time. Forbes. have been found. and extending for at least 300 miles from Vienna to Switzerland. for instance. several thousand. or when we follow them in nature. The frequent and great changes in the mineralogical composition of consecutive formations. the several species of the Chthamalinae (a sub-family of sessile cirripedes) coat the rocks all over the world in infinite numbers: they are all strictly littoral. whereas not one other species has hitherto been found in any tertiary formation: yet it is known that the genus Chthamalus existed during the Chalk period. when we remember how large a proportion of the bones of tertiary mammals have been discovered either in caves or in lacustrine deposits. Dawson in the carboniferous strata of North America. and in many other parts of the world. namely. The most skilful geologist. Scarcely any fact struck me more when examining many hundred miles of the South American coasts. This doctrine has been emphatically admitted by many geologists and palaeontologists. so it is in North America. how accidental and rare is their preservation. For instance. And if. who. with the exception of one species discovered by Sir C. if his attention had been confined exclusively to these large territories. which consists of shale and sandstone. For instance.

solid. as soon as they are brought up by the slow and gradual rising of the land within the grinding action of the coast-waves. conclude that sediment must be accumulated in extremely thick. All geological facts tell us plainly that each area has undergone numerous slow oscillations of level. in which case the bottom will not be inhabited by so many and such varied forms of life as the more shallow seas. has come to the conclusion that it was accumulated during subsidence. in treating of this or that great formation. or. have thus been formed during subsidence. In the case of an extensive and shallow sea. but which will hardly last to a distant geological age. will have been formed over wide spaces during periods of subsidence. We may. A little reflection will explain why. as well as the subsequent subaerial degradation. Still less can this have happened during the alternate periods of elevation. I may add. which has been bulky enough to resist such degradation as it has as yet suffered. as long as the bed of the sea remained stationary. The explanation.several hundred feet within the recent period. that the only ancient tertiary formation on the west coast of South America. and thus a rich fossiliferous formation. I have watched the progress of geology. when first upraised and during subsequent oscillations of level. formations rich in fossils and sufficiently thick and extensive to resist subsequent degradation. On the other hand. to resist a large amount of denudation. no extensive formations with recent or tertiary remains can anywhere be found. or extensive masses. if it continue slowly to subside. tertiary beds are so poorly developed that no record of several successive and peculiar marine faunas will probably be preserved to a distant age. is that the littoral and sub-littoral deposits are continually worn away. as long as the rate of subsidence and supply of sediment nearly balance each other. though the supply of sediment must for ages have been great. Such thick and extensive accumulations of sediment may be formed in two ways. was deposited during a downward oscillation of level. either in profound depths of the sea. along the rising coast of the western side of South America. the sea will remain shallow and favourable for many and varied forms. and thus gained considerable thickness. but only where the supply of sediment was sufficient to keep the sea shallow and to embed and preserve the remains before they had time to decay. I am convinced that nearly all our ancient formations. to speak more accurately. in order to withstand the incessant action of the waves. from the enormous degradation of the coast rocks and from the muddy streams entering the sea. when upraised. Since publishing my views on this subject in 1845. which is inhabited by a peculiar marine fauna. and the mass when upraised will give an imperfect record of the organisms which existed in the neighbourhood during the period of its accumulation. and have been surprised to note how author after author. I think. and apparently these oscillations have affected wide spaces. may be formed. which are throughout the greater part of their thickness RICH IN FOSSILS. Consequently. no doubt. thick enough. the beds which were then accumulated will generally have been destroyed by being upraised and brought within the limits of the coast-action. These remarks apply chiefly to littoral and sublittoral deposits. than the absence of any recent deposits sufficiently extensive to last for even a short geological period. THICK deposits cannot have been accumulated in the shallow parts. In this latter case. Along the whole west coast. such as that within a large part . Or sediment may be deposited to any thickness and extent over a shallow bottom. which are the most favourable to life.

Italy. for owing to the elevatory movement it would be less than the depth in which it was formed. the deposit formed during the rising movement. and the British Islands. subsided. the granitic area is very large: thus Von Eschwege gives a detailed section of these rocks. One remark is here worth a passing notice. Hopkins also expresses his belief that sedimentary beds of considerable horizontal extent have rarely been completely destroyed. nor would the deposit be much consolidated. and I find that the metamorphic (excluding the "semi-metamorphic") and granite rocks exceed. only one small patch of slightly metamorphosed rock. I have estimated the areas by cutting out and weighing the paper. For it is scarcely possible that such rocks could have been solidified and crystallised while uncovered. which alone could have formed a part of the original capping of the granitic series. I saw. will admit that these latter rocks have been stripped of their covering to an enormous extent. excepting the few who believe that our present metamorphic schists and plutonic rocks once formed the primordial nucleus of the globe. Rogers' beautiful map. part of Germany. that if one part of the area. the former protecting mantle of rock may not have been very thick. collected along the whole coast. diorite. and thus be preserved for a long period. except on the belief that they have subsequently been completely denuded of all overlying strata? That such extensive areas do exist cannot be doubted: the granitic region of Parime is described by Humboldt as being at least nineteen times as large as Switzerland. where the depth varies from thirty or forty to sixty fathoms. been suggested by Mr. stretching from Rio de Janeiro for 260 geographical miles inland in a straight line. nor be capped by overlying formations. were examined by me. however. with not a wreck left behind. granite. and yet not suffer excessively from denudation during its slow upheaval. Hopkins. It has. along the whole northern bank of the Plata.of the Malay Archipelago. During periods of elevation the area of the land and of the adjoining shoal parts of the sea will . France.. and saw nothing but granitic rocks. the whole of the newer Palaeozoic formations. South of the Amazon. might afterwards become protected by fresh accumulations. and I travelled for 150 miles in another direction. if all the sedimentary beds were removed which rest unconformably on them. This region has not been carefully explored. and which could not have formed part of the original mantle under which they were crystallised. how can we account for the naked and extensive areas of such rocks in many parts of the world. after rising and before being denuded. mica-schist. from near Rio de Janeiro to the mouth of the Plata. Hence.D. and they all belonged to this class. so that it would run a good chance of being worn away by atmospheric degradation and by the action of the sea during subsequent oscillations of level. In many regions the metamorphic and granite rocks would be found much more widely extended than they appear to be. Admitting then that gneiss. in the proportion of 19 to 12. Turning to a well-known region. Numerous specimens. besides modern tertiary beds. to the United States and Canada. but from the concurrent testimony of travellers. all conjoined. a distance of 1. though not thick. a widely extended formation might be formed during a period of elevation. But all geologists.5. but if the metamorphic action occurred at profound depths of the ocean. namely. were once necessarily covered up.100 geographical miles. Mr. Inland. Boue colours an area composed of rocks of this nature as equal to that of Spain. etc. as shown in Professor H. it is probable that in some parts of the world whole formations have been completely denuded. but the thickness of the formation could not be great.

for the formation of new varieties and species. time having apparently been required for their migration from the American to the European seas. as it seems to me. it has everywhere been noted. prevent us from coming to any just conclusion on this head. On the other hand. conversely. nor have the several stages of the same formation throughout Europe been correlated with perfect accuracy. when we find a species disappearing before the last layers have been deposited. during subsidence. I am aware that two palaeontologists. it would be rash in the extreme to infer that it had not elsewhere previously existed. have concluded that the average duration of each formation is twice or thrice as long as the average duration of specific forms. When we see a species first appearing in the middle of any formation. namely Bronn and Woodward. as previously explained. So again. but during such periods there will generally be a blank in the geological record. We may safely infer that with marine animals of all kinds there has been a large amount of migration due to climatal and other changes. viewed as a whole. From these several considerations it cannot be doubted that the geological record. but have become extinct in the immediately surrounding sea. and consequently during subsidence. whose opinions are worthy of much deference. In examining the latest deposits. few new varieties or species will be formed. that several species appear somewhat earlier in the palaeozoic beds of North America than in those of Europe. It is an excellent lesson to reflect on the ascertained amount of migration of the inhabitants of Europe during the glacial epoch. the probability is that it only then first immigrated into that area. for instance. Although each formation may mark a very long lapse of years. Although each formation has indisputably required a vast number of years for its deposition. each probably is short compared with the period requisite to change one species into another. it would be equally rash to suppose that it then became extinct.be increased and new stations will often be formed--all circumstances favourable. that some few still existing species are common in the deposit. . and it is during these very periods of subsidence that the deposits which are richest in fossils have been accumulated. but are rare or absent in this particular deposit. Several cases are on record of the same species presenting varieties in the upper and lower parts of the same formation. is extremely imperfect. but I cannot assign due proportional weight to the following considerations. and Hilgendorf has described a most curious case of ten graduated forms of Planorbis multiformis in the successive beds of a fresh-water formation in Switzerland. though there will be much extinction. or. But insuperable difficulties. Thus Trautschold gives a number of instances with Ammonites. that some are now abundant in the neighbouring sea. We forget how small the area of Europe is compared with the rest of the world. It is well known. ON THE ABSENCE OF NUMEROUS INTERMEDIATE VARIETIES IN ANY SINGLE FORMATION. and when we see a species first appearing in any formation. but if we confine our attention to any one formation. it becomes much more difficult to understand why we do not therein find closely graduated varieties between the allied species which lived at its commencement and at its close. several reasons can be given why each should not commonly include a graduated series of links between the species which lived at its commencement and close. in various quarters of the world. the inhabited area and number of inhabitants will decrease (excepting on the shores of a continent when first broken up into an archipelago).

But we have seen that a thick formation. Many instances could be given of beds. representing formations which are elsewhere thousands of feet in thickness. which is necessary that the same marine species may live on the same space. submerged. Yet it may be doubted whether. would be tempted to conclude that the average duration of life of the embedded fossils had been less than that of the glacial period. probable that sediment was deposited during the whole of the glacial period near the mouth of the Mississippi. Dawson found carboniferous beds 1.400 feet thick in Nova Scotia. a formation composed of beds of widely different mineralogical composition. of many long intervals of time and changes of level during the process of deposition. and to keep the depth approximately the same. can accumulate only during a period of subsidence. for it has been observed by more than one palaeontologist that very thick deposits are usually barren of organic remains. the deposit must be a very thick one. the supply of sediment must nearly counterbalance the amount of subsidence. at no less than sixty-eight different levels. and then re-covered by the upper beds of the same formation--facts. In fact. has generally been intermittent in its accumulation. When such beds as were deposited in shallow water near the mouth of the Mississippi during some part of the glacial period shall have been upraised. organic remains will probably first appear and disappear at different levels. Nor will the closest inspection of a formation give us any idea of the length of time which its deposition may have consumed. It would seem that each separate formation. examining these beds. In order to get a perfect gradation between two forms in the upper and lower parts of the same formation. still standing upright as they grew. that is. within that limit of depth at which marine animals can best flourish: for we know that great geographical changes occurred in other parts of America during this space of time. had not the trees been preserved: thus Sir C. But this same movement of subsidence will tend to submerge the area whence the sediment is derived. all included within this same glacial period. and which must have required an enormous period for their accumulation. this nearly exact balancing between the supply of sediment and the amount of subsidence is probably a rare contingency.which forms only a part of one whole geological period. with ancient root-bearing strata. and on the great lapse of time. which would not have been suspected. And in the distant future. showing what wide. intervals have occurred in its accumulation. owing to the migrations of species and to geographical changes. except near their upper or lower limits. have gone on accumulating within the same area during the whole of this period. extending from before the glacial epoch to the present day. in any quarter of the world. whilst the downward movement continues. When we see. and the species undergoing change must have lived in the same district throughout the whole time. for instance. on the extreme change of climate. only a few feet in thickness. we may reasonably suspect that the process of deposition has been more or less interrupted. a geologist. and likewise to reflect on the changes of level. . yet no one ignorant of this fact would have even suspected the vast lapse of time represented by the thinner formation. sedimentary deposits. hence. instead of having been really far greater. fossiliferous throughout its entire thickness. like the whole pile of formations in any country. Lyell and Dr. the deposit must have gone on continuously accumulating during a long period. and thus diminish the supply. as is so often the case. INCLUDING FOSSIL REMAINS. It is not. one above the other. yet easily overlooked. Many cases could be given of the lower beds of a formation having been upraised. sufficient for the slow process of modification. denuded. In other cases we have the plainest evidence in great fossilised trees.

A. and they do this the more readily if the specimens come from different sub-stages of the same formation. Nor should it be forgotten. we can seldom hope to effect in any one geological section. they grant some little variability to each species. to distinct but consecutive stages of the same great formation. and top of a formation. they rank both as species. but when they meet with a somewhat greater amount of difference between any two forms. even if A were strictly intermediate between B and C. we find that the embedded fossils. and yet would not necessarily be strictly intermediate between them in all respects. as before explained. in opposition to the judgment of most naturalists. we should not recognise their blood-relationship. namely. that their varieties are generally at first local. or unless we admit. as we have formerly seen. though universally ranked as specifically different. middle. so that here again we have undoubted evidence of change in the direction required by the theory. but has disappeared and reappeared. Look again at the later tertiary deposits. and on this view we do find the kind of evidence of change which on the theory we ought to find. we have evidence of the frequent occurrence of slight modifications of the kind required. maintain that all these tertiary species are specifically distinct. It is notorious on what excessively slight differences many palaeontologists have founded their species. but to this latter subject I shall return in the following chapter. and a third. that A might be the actual progenitor of B and C. changes of form. If we look to rather wider intervals of time. unless we believe that these eminent naturalists have been misled by their imaginations. when the same species occurs at the bottom. So that we might obtain the parent-species and its several modified descendants from the lower and upper beds of the same formation. though the distinction is admitted to be very slight. but some excellent naturalists. perhaps many times. According to this .Hence. as Agassiz and Pictet. unless they are enabled to connect them together by the closest intermediate gradations. so that here. and should consequently rank them as distinct species. It is all-important to remember that naturalists have no golden rule by which to distinguish species and varieties. and that such local varieties do not spread widely and supplant their parent-form until they have been modified and perfected in some considerable degree. which include many shells believed by the majority of naturalists to be identical with existing species. Consequently if it were to undergo a considerable amount of modification during the deposition of any one geological formation. a section would not include all the fine intermediate gradations which must on our theory have existed. yet are far more closely related to each other than are the species found in more widely separated formations. and this. it would simply be ranked as a third and distinct species. the probability is that it has not lived on the same spot during the whole period of deposition. and that these late tertiary species really present no difference whatever from their living representatives. from the reasons just assigned. to be found in an older and underlying bed. during the same geological period. there is reason to suspect. unless at the same time it could be closely connected by intermediate varieties with either one or both forms. Supposing B and C to be two species. though perhaps slight. Some experienced conchologists are now sinking many of the very fine species of D'Orbigny and others into the rank of varieties. With animals and plants that propagate rapidly and do not wander much. that these tertiary species are all truly distinct from the recent. but abrupt. and unless we obtained numerous transitional gradations.

and destroy four-fifths of them. The Malay Archipelago is about the size of Europe from the North Cape to the Mediterranean. "Every species is a link between other allied forms. Most marine animals have a wide range. is certainly erroneous. and this again would greatly lessen the chance of our being able to trace the stages of transition in any one geological formation. But this ought not to be expected. so that. It may be worth while to sum up the foregoing remarks on the causes of the imperfection of the geological record under an imaginary illustration. excepting those of the United States of America. Godwin-Austen. which are ranked by some conchologists as distinct species from their European representatives. geologists at some future period will be able to prove that our different breeds of cattle. with shells and other marine animals. horses. and therefore equals all the geological formations which have been examined with any accuracy. This could be effected by the future geologist only by his discovering in a fossil state numerous intermediate gradations. I fully agree with Mr. that at the present day. and by other conchologists as only varieties. leading to the same result." If we take a genus having a score of species.view. best perceive the improbability of our being enabled to connect species by numerous. perhaps. by writers who believe in the immutability of species. as lately insisted on by Dr. that the present condition of the Malay Archipelago. no one doubts that the remainder will stand much more distinct from each other. far exceeding the limits of the known geological formations in Europe. It has been asserted over and over again. as fine as existing varieties. as we shall see in the next chapter. as it is called. it is probable that those which had the widest range. It is a more important consideration. and with fossil species this can rarely be done. If the extreme forms in the genus happen to have been thus destroyed. fossil links. that geology yields no linking forms. recent and extinct. or. and dogs are descended from a single stock or from several aboriginal stocks. that oftenest present varieties. What geological research has not revealed. or are. the chance of discovering in a formation in any one country all the early stages of transition between any two forms. Falconer. whether certain sea-shells inhabiting the shores of North America. and from Britain to Russia. and such success is improbable in the highest degree. intermediate. probably represents the former state of Europe. have oftenest given rise. first to local varieties and ultimately to new species. was probably short in comparison with that during which it remained without undergoing any change. with its numerous large islands separated by wide and shallow seas. We shall. yet this has been repeatedly advanced as a most serious objection against my views. that the period during which each species underwent modification. Lubbock has remarked. As Sir J. and thus proved to be the same species. is small. namely. This assertion. though long as measured by years. until many specimens are collected from many places. again. specifically distinct. with perfect specimens for examination. two forms can seldom be connected by intermediate varieties. is the former existence of infinitely numerous gradations. are really varieties. connecting together nearly all existing and extinct species. fine. by asking ourselves whether. for instance. for the successive changes are supposed to have been local or confined to some one spot. and we have seen that with plants it is those which have the widest range. It should not be forgotten. while most . sheep. the genus itself will stand more distinct from other allied genera.

according to the principles followed by many palaeontologists. During the periods of subsidence. which would oftenest produce new varieties. by the incessant coast-action. and these contingencies are indispensable for the preservation of all the transitional gradations between any two or more species. they would slowly spread and supplant their parent-forms. though only some of them. sedimentary beds could hardly be accumulated of great thickness during the periods of elevation. and of thickness sufficient to last to an age as distant in futurity as the secondary formations lie in the past.of our formations were accumulating. and those embedded in gravel or sand would not endure to a distant epoch. but if possessed of any decided advantage. would be embedded. no remains could be preserved. as we now see on the shores of South America. . Not many of the strictly littoral animals. and the varieties would at first be local or confined to one place. Wherever sediment did not accumulate on the bed of the sea. It is also probable that each great period of subsidence would be interrupted by oscillations of level. so as to have a good chance of enduring to a very distant future. Formations rich in fossils of many kinds. and analogy plainly leads to the belief that it would be chiefly these far-ranging species. though closely allied species. but the geological record would then be less perfect. Very many of the marine inhabitants of the archipelago now range thousands of miles beyond its confines. and that slight climatical changes would intervene during such lengthy periods. together with a contemporaneous accumulation of sediment. or when further modified and improved. there would probably be much extinction of life. and in these cases the inhabitants of the archipelago would migrate. or where it did not accumulate at a sufficient rate to protect organic bodies from decay. be ranked as new and distinct species. during the periods of elevation. though perhaps extremely slight degree. they would. during which the area would be either stationary or rising. how imperfectly would they represent the natural history of the world! But we have every reason to believe that the terrestrial productions of the archipelago would be preserved in an extremely imperfect manner in the formations which we suppose to be there accumulating. there would be much variation. yet if all the species were to be collected which have ever lived there. Even throughout the extensive and shallow seas within the archipelago. When such varieties returned to their ancient homes. These periods of subsidence would be separated from each other by immense intervals of time. and no closely consecutive record of their modifications could be preserved in any one formation. would EXCEED the average duration of the same specific forms. or become capped and protected by subsequent deposits. transitional varieties would merely appear as so many new. or of those which lived on naked submarine rocks. If such gradations were not all fully preserved. would generally be formed in the archipelago only during periods of subsidence. whilst rising. and as they would be found embedded in slightly different sub-stages of the same formation. It may be doubted whether the duration of any one great period of subsidence over the whole or part of the archipelago. the fossiliferous formations on the steeper shores would be destroyed. as they would differ from their former state in a nearly uniform. almost as soon as accumulated. The Malay Archipelago is one of the richest regions in organic beings.

as a fatal objection to the belief in the transmutation of species. for instance. have not these birds their front limbs in this precise intermediate state of "neither true arms nor true wings?" Yet these birds hold their place victoriously in the battle for life. If numerous species. In all cases positive palaeontological evidence may be implicitly trusted. But we continually overrate the perfection of the geological record. namely. I may here recall a remark formerly made. We continually forget how large the world is. Pictet. ON THE SUDDEN APPEARANCE OF WHOLE GROUPS OF ALLIED SPECIES. and taking birds as an illustration. because certain genera or families have not been found beneath a certain stage. and such assuredly we do find--some more distantly. in commenting on early transitional forms. for they exist in infinite numbers and of many kinds. that it might require a long succession of ages to adapt an organism to some new and peculiar line of life. has been urged by several palaeontologists--for instance. compared with the area over which our geological formations have been carefully examined. and falsely infer.If then there be some degree of truth in these remarks. which. the fact would be fatal to the theory of evolution through natural selection. in our geological formations. as experience has so often shown. But I do not pretend that I should ever have suspected how poor was the record in the best preserved geological sections. must have been an extremely slow process. and have slowly multiplied. related to each other. But look at the penguins of the Southern Ocean. belonging to the same genera or families. some more closely. and these links. Professor Pictet. have really started into life at once. These intervals will have given time for the multiplication of species from some one parent-form: and in the succeeding formation. in his excellent Review of this work. would. on our theory. all of which are descended from some one progenitor. We do not make due allowance for the enormous intervals of time which have elapsed between our consecutive formations. that they did not exist before that stage. before they invaded the ancient archipelagoes of Europe and the United States. and consequently that the transitional forms would often long remain confined to some one region. but that. be ranked as distinct species. The abrupt manner in which whole groups of species suddenly appear in certain formations. such groups or species will appear as if suddenly created. to fly through the air. We ought only to look for a few links. which would spread rapidly and widely throughout the world. an infinite number of those fine transitional forms. and Sedgwick. had not the absence of innumerable transitional links between the species which lived at the commencement and close of each formation. pressed so hardly on my theory. by many palaeontologists. I do not suppose that we here see the real transitional grades through which the wings of birds have passed. have connected all the past and present species of the same group into one long and branching chain of life. cannot see how the successive modifications of the anterior limbs of a supposed prototype could possibly have been of any advantage. longer perhaps in many cases than the time required for the accumulation of each formation. if found in different stages of the same formation. let them be ever so close. negative evidence is worthless. but what special difficulty is there in . by Agassiz. we have no right to expect to find. For the development by this means of a group of forms. a comparatively short time would be necessary to produce many divergent forms. when this adaptation had once been effected. and a few species had thus acquired a great advantage over other organisms. we forget that groups of species may elsewhere have long existed. and the progenitors must have lived long before their modified descendants.

some of gigantic size. Woodward in the upper chalk. and with its wings furnished with two free claws. the Archeopteryx. from the ease with which even a fragment of a valve can be recognised. from the perfect manner in which specimens are preserved in the oldest tertiary beds. adding. Even in so short an interval as that between the first and second editions of Pictet's great work on Palaeontology. M. of which not one species has as yet been found even in any tertiary stratum. the conclusions on the first appearance and disappearance of several groups of animals have been considerably modified. as I then thought. as if to make the case as striking as possible. a very common. on the authority of Professor Owen. that strange bird. In a memoir on Fossil Sessile Cirripedes. has been discovered by Mr.believing that it might profit the modified descendants of the penguin. I may give another instance. but now we know. Had it not been for the rare accident of the preservation of footsteps in the new red sandstone of the United States. Hardly any recent discovery shows more forcibly than this how little we as yet know of the former inhabitants of the world. a member of a distinct subfamily of sessile cirripedes. mammals were always spoken of as having abruptly come in at the commencement of the tertiary series. And now one of the richest known accumulations of fossil mammals belongs to the middle of the secondary series. and ultimately to rise from its surface and glide through the air? I will now give a few examples to illustrate the foregoing remarks. had sessile cirripedes existed during the secondary periods. Not long ago. published not many years ago. Cuvier used to urge that no monkey occurred in any tertiary stratum. existed during that period? Not a fragment of bone has been discovered in these beds. I concluded that this great group had been suddenly developed at the commencement of the tertiary series. This was a sore trouble to me. I may recall the well-known fact that in geological treatises. and a third edition would require still further changes. they would certainly have been preserved and discovered. and still more recently. that a bird certainly lived during the deposition of the upper greensand. I stated that. I inferred that. from having passed under my own eyes has much struck me. inhabiting various zones of depths. and true mammals have been discovered in the new red sandstone at nearly the commencement of this great series. large. one more instance of the abrupt appearance of a great group of species. as far back as the miocene stage. Still more recently. South America and in Europe. and to show how liable we are to error in supposing that whole groups of species have suddenly been produced. sent me a drawing of a perfect specimen of an unmistakable sessile cirripede. from all these circumstances. when a skilful palaeontologist. from the extraordinary abundance of the individuals of many species all over the world. and ubiquitous genus. but now extinct species have been discovered in India. published in 1844-46 and in 1853-57. so that we now have abundant evidence of the existence . palaeontologists maintained that the whole class of birds came suddenly into existence during the eocene period. has been discovered in the oolitic slates of Solenhofen. But my work had hardly been published. which he had himself extracted from the chalk of Belgium. first to become enabled to flap along the surface of the sea like the logger-headed duck. a Pyrgoma. from the upper tidal limits to fifty fathoms. from the large number of existing and extinct tertiary species. with a long lizard-like tail. who would have ventured to suppose that no less than at least thirty different bird-like animals. which. this cirripede was a Chthamalus. Bosquet. from the Arctic regions to the equator. bearing a pair of feathers on each joint. and as not one species had then been discovered in beds of this age. And.

in the Chalk period. and were enabled to double the southern capes of Africa or Australia. from our ignorance of the geology of other countries beyond the confines of Europe and the United States. unless it could likewise have been shown that at the same period the species were suddenly and simultaneously developed in other quarters of the world. Nor have we any right to suppose that the seas of the world have always been so freely open from south to north as they are at present. apply with equal force to the earliest known species. the whole interval from the Cambrian age to the present day. Even at this day. But certain Jurassic and Triassic forms are now commonly admitted to be teleostean. in which any great group of marine animals might be multiplied. I allude to the manner in which species belonging to several of the main divisions of the animal kingdom suddenly appear in the lowest known fossiliferous rocks. Lingula. From these considerations. etc. which is much more serious. and that during these vast periods the world swarmed . and it cannot on our theory be supposed. as the Nautilus. if the theory be true. and thus reach other and distant seas. the fact would have been highly remarkable. have spread widely. which must have lived long before the Cambrian age. The case most frequently insisted on by palaeontologists of the apparently sudden appearance of a whole group of species. or probably far longer than. Consequently. the teleostean fishes might formerly have had a similarly confined range. and from the revolution in our palaeontological knowledge effected by the discoveries of the last dozen years. This group includes the large majority of existing species. Some of the most ancient animals. and by running through Pictet's Palaeontology it will be seen that very few species are known from several formations in Europe. and which probably differed greatly from any known animal. for they are not in any degree intermediate in character. and here they would remain confined. it seems to me to be about as rash to dogmatize on the succession of organic forms throughout the world. until some of the species became adapted to a cooler climate. as it would be for a naturalist to land for five minutes on a barren point in Australia. according to Agassiz. and then to discuss the number and range of its productions. It is almost superfluous to remark that hardly any fossil-fish are known from south of the equator.. low down. it is indisputable that before the lowest Cambrian stratum was deposited long periods elapsed. it cannot be doubted that all the Cambrian and Silurian trilobites are descended from some one crustacean. and after having been largely developed in some one sea. but it would not have formed an insuperable difficulty.of this group of animals during the secondary period. that these old species were the progenitors of all the species belonging to the same groups which have subsequently appeared. do not differ much from living species. There is another and allied difficulty. For instance. the tropical parts of the Indian Ocean would form a large and perfectly enclosed basin. and even some palaeozoic forms have thus been classed by one high authority. if the Malay Archipelago were converted into land. is that of the teleostean fishes. as long as. Most of the arguments which have convinced me that all the existing species of the same group are descended from a single progenitor. Some few families of fish now have a confined range. If the teleosteans had really appeared suddenly in the northern hemisphere at the commencement of the chalk formation. ON THE SUDDEN APPEARANCE OF GROUPS OF ALLIED SPECIES IN THE LOWEST KNOWN FOSSILIFEROUS STRATA.

and now. even in some of the lowest azotic rocks. Thompson concludes that the consolidation of the crust can hardly have occurred less than twenty or more than four hundred million years ago. Logan states that their "united thickness may possibly far surpass that of all the succeeding rocks. but this. still lower down in the Lower Cambrian formation. and the previous one hundred and forty million years can hardly be considered as sufficient for the development of the varied forms of life which already existed during the Cambrian period. Forbes. There are three great series of strata beneath the Silurian system in Canada. The presence of phosphatic nodules and bituminous matter. however. in a fit state for the habitation of living creatures. that the appearance of the so-called primordial fauna (of Barrande) may by some be considered as a comparatively modern event. which I wrote in 1859. Mr. But the descriptions which we possess of the Silurian deposits over immense territories in Russia and in North America. and these would always have existed in a partially metamorphosed condition. beneath the then known Silurian system. it existed in countless numbers. To the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods prior to the Cambrian system. the difficulty of assigning any good reason for the absence of vast piles of strata rich in fossils beneath the Cambrian system is very great. judging from the small amount of organic change since the commencement of the Glacial epoch. were until recently convinced that we beheld in the organic remains of the lowest Silurian stratum the first dawn of life. and. with Sir R. Murchison at their head. We are thus carried back to a period so remote. appears a very short time for the many and great mutations of life. Here we encounter a formidable objection. in the lowest of which the Eozoon is found. Croll estimates that about sixty million years have elapsed since the Cambrian period. probable. from the base of the palaeozoic series to the present time. These very wide limits show how doubtful the data are. I can give no satisfactory answer. certainly preyed on other minute organic beings. but is highly organised for its class. Sir W. and other elements may have hereafter to be introduced into the problem. which have certainly occurred since the Cambrian formation. Dawson has remarked. Nevertheless. and which are almost the same with those since used by Sir W. Not very long ago M. abounding with new and peculiar species. Sir W. We should not forget that only a small portion of the world is known with accuracy. Mr Hicks has found South Wales beds rich in trilobites. have disputed this conclusion. for it seems doubtful whether the earth. about the existence of living beings long before the Cambrian period. and containing various molluscs and annelids. as Dr. Logan. Several eminent geologists. as Lyell and E. and such changes would have tended to induce changes at a corresponding rate in the organisms which then existed. It is. It does not seem probable that the most ancient beds have been quite worn away by denudation. as Sir William Thompson insists. or that their fossils have been wholly obliterated by metamorphic action. probably indicates life at these periods. but probably not less than ninety-eight or more than two hundred million years. and the existence of the Eozoon in the Laurentian formation of Canada is generally admitted. Barrande added another and lower stage. Thus the words. do not support the view that the older a formation is the more invariably it . for if this had been the case we should have found only small remnants of the formations next succeeding them in age. Other highly competent judges.with living creatures. has lasted long enough. that the world at a very early period was subjected to more rapid and violent changes in its physical conditions than those now occurring. have proved true. which must have lived in great numbers." The Eozoon belongs to the most lowly organised of all classes of animals.

led me to conclude that the great oceans are still mainly areas of subsidence. since the Cambrian period. but hardly one truly oceanic island (with the exception of New Zealand. the great archipelagoes still areas of oscillations of level. if this can be called a truly oceanic island) is as yet known to afford even a remnant of any palaeozoic or secondary formation. Our continents seem to have been formed by a preponderance. we may infer that from first to last large islands or tracts of land. occurred in the neighbourhood of the now existing continents of Europe and North America. which are thrice as extensive as the land. and would have been at least partially upheaved by the oscillations of level. But may not the areas of preponderant movement have changed in the lapse of ages? At a period long antecedent to the Cambrian epoch. The coloured map appended to my volume on Coral Reefs. namely. that where continents now exist. we see them studded with many islands. for it might well happen that strata which had subsided some miles nearer to the centre of the earth. or as a submarine surface near land. neither continents nor continental islands existed where our oceans now extend. we may infer anything from these facts. whether Europe and the United States during these intervals existed as dry land. and the continents areas of elevation.has suffered extreme denudation and metamorphism. of naked metamorphic rocks. If. during many oscillations of level. no doubt. and may be truly urged as a valid argument against the views here entertained. to great oscillations of level. though we find in . of the force of elevation. Hence. The case at present must remain inexplicable. older than the Cambrian strata. whence the sediment was derived. The several difficulties here discussed. and clear and open oceans may have existed where our continents now stand. From the nature of the organic remains which do not appear to have inhabited profound depths. we may perhaps infer. in recognisable condition. which must have intervened during these enormously long periods. for instance. oceans have extended from the remotest period of which we have any record. But we do not know what was the state of things in the intervals between the several successive formations. I will give the following hypothesis. where our oceans now extend. miles in thickness. subjected. To show that it may hereafter receive some explanation. and which had been pressed on by an enormous weight of superincumbent water. in the several formations of Europe and of the United States. palaeozoic and secondary formations would in all probability have been accumulated from sediment derived from their wear and tear. This same view has since been maintained by Agassiz and others. for had they existed. that. on which sediment was not deposited. for instance in South America. that during the palaeozoic and secondary periods. Nor should we be justified in assuming that if. we may infer that. of which the formations are composed. The immense areas in some parts of the world. continents may have existed where oceans are now spread out. supposing such to have been formerly deposited. then. which must have been heated under great pressure. might have undergone far more metamorphic action than strata which have always remained nearer to the surface. and from the amount of sediment. or as the bed of an open and unfathomable sea. have always seemed to me to require some special explanation. the bed of the Pacific Ocean were now converted into a continent we should there find sedimentary formations. large tracts of land have existed. Looking to the existing oceans. and we may perhaps believe that we see in these large areas the many formations long anterior to the Cambrian epoch in a completely metamorphosed and denuded condition. and on the other hand. But we have no reason to assume that things have thus remained from the beginning of the world.

etc. and which falsely appear to have been abruptly introduced. I look at the geological record as a history of the world imperfectly kept and written in a changing dialect. of formations rich in fossils beneath the Cambrian strata.. In some of the most recent beds. following out Lyell's metaphor. Falconer. etc. namely. maintained the immutability of species. on the face of the earth. Those who believe that the geological record is in any degree perfect.our geological formations many links between the species which now exist and which formerly existed. either locally. The sudden manner in which several groups of species first appear in our European formations. or. ON THE GEOLOGICAL SUCCESSION OF ORGANIC BEINGS. which are entombed in our consecutive formations.. one after another. only here and there a few lines. and of each page. E. and most geologists and palaeontologists are much shaken in their former belief. Of this history we possess the last volume alone. Murchison. as far as we know. Barrande. have unanimously. as Bronn has remarked. CHAPTER XI. Pictet. Sedgwick. neither the appearance nor disappearance of the many species embedded in each formation has been simultaneous. and all our greatest geologists. Cuvier. We see this in the fact that the most eminent palaeontologists. Let us now see whether the several facts and laws relating to the geological succession of organic beings accord best with the common view of the immutability of species. will undoubtedly at once reject my theory. only here and there a short chapter has been preserved. Agassiz. and every year tends to fill up the blanks between the stages. For my part. Of this volume. though undoubtedly of high antiquity if measured by years. New species have appeared very slowly. often vehemently. we do not find infinitely numerous fine transitional forms closely joining them all together. and to make the proportion between the lost and existing forms more gradual. as at present known. and only one or two are new. more or less different in the successive chapters. Forbes. as Lyell. or with that of their slow and gradual modification. having appeared there for the first time. may represent the forms of life. Each word of the slowly-changing language. both on the land and in the waters. only one or two species are extinct. On this view the difficulties above discussed are greatly diminished or even disappear. are all undoubtedly of the most serious nature. The secondary formations are more broken. the almost entire absence. Species belonging to different genera and classes have not changed at . but. But Sir Charles Lyell now gives the support of his high authority to the opposite side. relating only to two or three countries. through variation and natural selection. On the slow and successive appearance of new species--On their different rates of change--Species once lost do not reappear--Groups of species follow the same general rules in their appearance and disappearance as do single species--On extinction--On simultaneous changes in the forms of life throughout the world--On the affinities of extinct species to each other and to living species--On the state of development of ancient forms--On the succession of the same types within the same areas--Summary of preceding and present chapters. Lyell has shown that it is hardly possible to resist the evidence on this head in the case of the several tertiary stages.

may. which intrude for a period in the midst of an older formation. perhaps. Falconer has given a striking instance of a similar fact. The process of modification must be slow. but as the accumulation of enduring formations. We find similar relations between the existing inhabitants of distinct countries. the land-shells and coleopterous insects of Madeira have come to differ considerably from their nearest allies on the continent of Europe. Hence it is by no means surprising that one species should retain the same identical form much longer than others. during long and equal periods of time. as Pictet has remarked. on the immigration of new colonists. our formations . The Silurian Lingula differs but little from the living species of this genus. causing all the inhabitants of an area to change abruptly. on the slowly changing physical conditions of the country. if changing. The productions of the land seem to have changed at a quicker rate than those of the sea. change more quickly than those that are low: though there are exceptions to this rule. we see why all the species in the same region do at last. or. depends on great masses of sediment being deposited on subsiding areas. namely. whereas most of the other Silurian Molluscs and all the Crustaceans have changed greatly. rich in fossils. thus causing a greater or less amount of permanent modification.the same rate. We can perhaps understand the apparently quicker rate of change in terrestrial and in more highly organised productions compared with marine and lower productions. When a species has once disappeared from the face of the earth. In the older tertiary beds a few living shells may still be found in the midst of a multitude of extinct forms. There is some reason to believe that organisms high in the scale. which includes no fixed law of development. if we look to long enough intervals of time. for an existing crocodile is associated with many lost mammals and reptiles in the sub-Himalayan deposits. In members of the same class the average amount of change. we have no reason to believe that the same identical form ever reappears. or simultaneously. all the species will be found to have undergone some change. that it is a case of temporary migration from a distinct geographical province. and on the nature of the other inhabitants with which the varying species come into competition. and from the all-important relations of organism to organism in the struggle for life. but Lyell's explanation. and will generally affect only a few species at the same time. we can understand. on the principle of competition. for the variability of each species is independent of that of all others. for instance. should change in a less degree. seems satisfactory. Barrande. for otherwise they would become extinct. When many of the inhabitants of any area have become modified and improved. be nearly the same. as explained in a former chapter. Hence. Yet if we compare any but the most closely related formations. and then allow the pre-existing fauna to reappear. is not the same in each successive so-called formation. Whether such variations or individual differences as may arise will be accumulated through natural selection in a greater or less degree. whereas the marine shells and birds have remained unaltered. These several facts accord well with our theory. or to an equal degree. The strongest apparent exception to this latter rule is that of the so-called "colonies" of M. of which a striking instance has been observed in Switzerland. would be liable to extermination. will depend on many complex contingencies--on the variations being of a beneficial nature. or in the same degree. on the freedom of intercrossing. by the more complex relations of the higher beings to their organic and inorganic conditions of life. The amount of organic change. become modified. that any form which did not become in some degree modified and improved.

For instance. taken almost at hazard. and all from a common progenitor. yet the two forms--the old and the new--would not be identically the same. when it has once disappeared. from the lowest Silurian stratum to the present day. are the modified descendants one from the other. This gradual increase in number of the species of a group is strictly conformable with the theory. and Woodward (though all strongly opposed to such views as I maintain) admit its truth. be represented by a vertical line of varying thickness. even if the very same conditions of life. For though the offspring of one species might be adapted (and no doubt this has occurred in innumerable instances) to fill the place of another species in the economy of nature. but the exceptions are surprisingly few. which if true would be fatal to my views. but abruptly. In the genus Lingula. ascending through the successive geological formations. genera and families. never reappears. organic and inorganic. identical with the existing breed. does not mark a new and complete act of creation. follow the same general rules in their appearance and disappearance as do single species. is continuous. as long as it lasts. We have seen in the last chapter that whole groups of species sometimes falsely appear to have been abruptly developed. if all our fantail-pigeons were destroyed. consequently the amount of organic change exhibited by the fossils embedded in consecutive formations is not equal. and organisms already differing would vary in a different manner. can increase only slowly and progressively. the general rule being a gradual increase in number. and ultimately thins out in the upper beds. For all the species of the same group. Each formation. or even from any other well established race of the domestic pigeon. its existence. and the newly-formed variety would probably inherit from its progenitor some characteristic differences. but if the parent rock-pigeon were likewise destroyed. or the number of the genera within a family. so few that E. sooner or later. the species which have successively appeared at all ages must have been connected by an unbroken series of generations. the process of modification and the production of a number of allied . and under nature we have every reason to believe that parent forms are generally supplanted and exterminated by their improved offspring. and then. that is. should recur. and the rule strictly accords with the theory. changing more or less quickly. it is possible. But such cases are certainly exceptional. marking the decrease and final extinction of the species. If the number of the species included within a genus. however long it may have lasted. Forbes. and the genera of the same family. I am aware that there are some apparent exceptions to this rule. for the species of the same genus. could be raised from any other species of pigeon. Pictet. and thus supplant it. it is incredible that a fantail. that is. until the group reaches its maximum. on this view. A group. for both would almost certainly inherit different characters from their distinct progenitors. Groups of species. and in a greater or lesser degree. not in a sharp point. the line will sometimes falsely appear to begin at its lower end. for the successive variations would almost certainly be in some degree different. for instance. often keeping of equal thickness for a space. in an ever slowly changing drama. that fanciers might make a new breed hardly distinguishable from the present breed. and I have attempted to give an explanation of this fact. but only an occasional scene. a gradual decrease.have been almost necessarily accumulated at wide and irregularly intermittent intervals of time. in which the species are found. We can clearly understand why a species when once lost should never reappear. it then gradually thickens upwards.

In some cases. from the study of the tertiary formations. first from one spot. But my astonishment was groundless. I was filled with astonishment. Barrande. we have every reason to believe. some groups. for rarity is the attribute of a vast number of species of all classes. No one can have marvelled more than I have done at the extinction of species. seeing that the horse. There is reason to believe that the extinction of a whole group of species is generally a slower process than their production: if their appearance and disappearance be represented. Both single species and whole groups of species last for very unequal periods. which marks the progress of extermination. We have as yet only spoken incidentally of the disappearance of species and of groups of species. ON EXTINCTION. has run wild over the whole country and has increased in numbers at an unparalleled rate. When I found in La Plata the tooth of a horse embedded with the remains of Mastodon. The extinction of species has been involved in the most gratuitous mystery. the extinction of old forms and the production of new and improved forms are intimately connected together. we might have felt certain. and from the history of the naturalisation of the domestic horse in . in all countries. as Elie de Beaumont. which all co-existed with still living shells at a very late geological period. Some authors have even supposed that. by a vertical line of varying thickness the line is found to taper more gradually at its upper end. Murchison. In some few cases. which in their turn produce by equally slow steps other varieties and species. one species first giving rise to two or three varieties. however.forms necessarily being a slow and gradual process. as of ammonites. I asked myself what could so recently have exterminated the former horse under conditions of life apparently so favourable. On the contrary. has been wonderfully sudden. as we have seen. the process of extinction may have been rapid. for. than at its lower end. towards the close of the secondary period. some have disappeared before the close of the palaeozoic period. the extermination of whole groups. No fixed law seems to determine the length of time during which any single species or any single genus endures. and finally from the world. have endured from the earliest known dawn of life to the present day. and so on. so have species a definite duration. as by the breaking of an isthmus and the consequent irruption of a multitude of new inhabitants into an adjoining sea. one after another. then from another. from the analogy of all other mammals. On the theory of natural selection. If we ask ourselves why this or that species is rare. Megatherium. whose general views would naturally lead them to this conclusion. these being slowly converted into species. even by those geologists. but what that something is. On the supposition of the fossil horse still existing as a rare species. as before. we answer that something is unfavourable in its conditions of life. Had this horse been still living. which marks the first appearance and the early increase in number of the species. however. or by the final subsidence of an island. Toxodon and other extinct monsters. since its introduction by the Spaniards into South America. as the individual has a definite length of life. belonged to an extinct species. even of the slow-breeding elephant. The old notion of all the inhabitants of the earth having been swept away by catastrophes at successive periods is very generally given up. that species and groups of species gradually disappear. like the branching of a great tree from a single stem. Professor Owen soon perceived that the tooth. till the group becomes large. etc. but in some degree rare. we can hardly ever tell.. no naturalist would have felt the least surprise at its rarity. though so like that of the existing horse.

and we know that this has been the progress of events with those animals which have been exterminated. that to admit that species generally become rare before they become extinct--to feel no surprise at the rarity of a species. check its increase. at least during the later geological epochs. and yet to marvel greatly when the species ceases to exist. becoming less and less favourable. that I have heard surprise repeatedly expressed at such great monsters as the Mastodon and the more ancient Dinosaurians having become extinct. and in what degree they severally acted. Falconer. looking to later times. as if mere bodily strength gave victory in the battle of life. is much the same as to admit that sickness in the individual is the forerunner of death--to feel no surprise at sickness. but. and the consequent extinction of less-favoured forms almost inevitably follows. I may repeat what I published in 1845. through man's agency. namely. We see in many cases in the more recent tertiary formations that rarity precedes extinction. But we could not have told what the unfavourable conditions were which checked its increase. The theory of natural selection is grounded on the belief that each new variety and ultimately each new species. are bound together. to wonder and to suspect that he died by some deed of violence. when much improved it is transported far and near. In flourishing groups. believes that it is chiefly insects which. some cause must have checked the continued increase of the existing elephant. either locally or wholly. as has been remarked by Owen. on the contrary. and finally extinction. as formerly explained and illustrated by examples. Mere size. It is most difficult always to remember that the increase of every living creature is constantly being checked by unperceived hostile agencies. It is the same with our domestic productions: when a new and slightly improved variety has been raised. and finally extinct--its place being seized on by some more successful competitor. we assuredly should not have perceived the fact. the number of new specific forms which have been produced within a given time has at some periods probably been greater than the number of the old specific forms which have been exterminated. that under more favourable conditions it would in a very few years have stocked the whole continent. quicker extermination. from incessantly harassing and weakening the elephant in India.South America. and that these same unperceived agencies are amply sufficient to cause rarity. and this was Bruce's conclusion with respect to the African elephant in Abyssinia. but we know that species have not gone on indefinitely increasing. So little is this subject understood. whether some one or several contingencies. Before man inhabited India or Africa. The competition will generally be most severe. like our short-horn cattle. both those naturally and artificially produced. from the greater amount of requisite food. yet the fossil horse would certainly have become rarer and rarer. It is certain that insects and blood-sucking bats determine the existence of the larger naturalised quadrupeds in several parts of South America. If the conditions had gone on. Dr. and at what period of the horse's life. however slowly. we may believe that the production of new forms has caused the extinction of about the same number of old forms. and takes the place of other breeds in other countries. so that. when the sick man dies. would in some cases determine. between the forms which are most like each . Thus the appearance of new forms and the disappearance of old forms. it at first supplants the less improved varieties in the same neighbourhood. A highly capable judge. is produced and maintained by having some advantage over those with which it comes into competition.

and in these intervals there may have been much slow extermination. Thus. and of Ammonites at the close of the secondary period. ON THE FORMS OF LIFE CHANGING ALMOST SIMULTANEOUSLY THROUGHOUT THE WORLD. the organic . Hence the improved and modified descendants of a species will generally cause the extermination of the parent-species. many species of a new group have taken possession of an area. from being fitted to some peculiar line of life. If many allied forms be developed from the successful intruder. let it be at our presumption in imagining for a moment that we understand the many complex contingencies on which the existence of each species depends. the whole economy of nature will be utterly obscured. With respect to the apparently sudden extermination of whole families or orders. We need not marvel at extinction. the utter extinction of a group is generally. For instance. But whether it be species belonging to the same or to a distinct class. and thus have caused its extermination. that is a new genus. then. Whenever we can precisely say why this species is more abundant in individuals than that. we may justly feel surprise why we cannot account for the extinction of any particular species or group of species. some species of Trigonia. and the forms which thus yield their places will commonly be allied. if we must marvel. the manner in which single species and whole groups of species become extinct accords well with the theory of natural selection. namely. in Tierra del Fuego. the species of the same genus. when. But it must often have happened that a new species belonging to some one group has seized on the place occupied by a species belonging to a distinct group. and that some check is always in action. as I believe. yet seldom perceived by us. which will suffer from some inherited inferiority in common. and if many new forms have been developed from any one species. many of the older species will have been exterminated in a correspondingly rapid manner. and a few members of the great and almost extinct group of Ganoid fishes still inhabit our fresh waters. Scarcely any palaeontological discovery is more striking than the fact that the forms of life change almost simultaneously throughout the world. If we forget for an instant that each species tends to increase inordinately. For at these distant points.other in all respects. i. and in the peninsula of India. and it will generally be the allied forms. where they will have escaped severe competition. Therefore. as of Trilobites at the close of the palaeozoic period. where not a fragment of the mineral chalk itself can be found. we must remember what has been already said on the probable wide intervals of time between our consecutive formations. at the Cape of Good Hope. a number of new species descended from one species. Thus our European Chalk formation can be recognised in many distant regions. Thus. Moreover. survive in the Australian seas. many will have to yield their places. for they will partake of the same inferiority in common. belonging to the same family. a few of the sufferers may often be preserved for a long time. or from inhabiting some distant and isolated station. as it seems to me. will be the most liable to extermination. why this species and not another can be naturalised in a given country. by sudden immigration or by unusually rapid development. and not until then. as we have seen. which have yielded their places to other modified and improved species.e. a slower process than its production. in equatorial South America. a great genus of shells in the secondary formations. the nearest allies of that species. comes to supplant an old genus. under the most different climates. in North America.

in the palaeozoic and tertiary stages. So. and from not including those forms which are found only in the older underlying deposits. so it is. they add. than to the present inhabitants of Europe. we turn our attention to North America. it must not be supposed that this expression relates to the same year. without any information in regard to their geological position. and sometimes are similarly characterised in such trifling points as mere superficial sculpture. and Toxodon had been brought to Europe from La Plata. and there discover a series of analogous phenomena.remains in certain beds present an unmistakable resemblance to those of the Chalk. at distant parts of the world. The fact of the forms of life changing simultaneously in the above large sense. but they belong to the same families. Nevertheless. or even to the same century. Even if the few fossil species which are common to the Old and New Worlds were kept wholly out of view. it is evident that fossiliferous beds now deposited on the shores of North America would hereafter be liable to be classed with somewhat older European beds. and sections of genera. Moreover. After referring to the parallelism of the palaeozoic forms of life in various parts of Europe. has greatly struck those admirable observers. their . from containing fossil remains in some degree allied. the most skilful naturalist would hardly be able to say whether the present or the pleistocene inhabitants of Europe resembled most closely those of the southern hemisphere. the upper pliocene. according to Lyell. a similar parallelism in the forms of life has been observed by several authors. occur in the same order at these distant points of the world. it might at least have been inferred that they had lived during one of the later tertiary stages. including the whole glacial epoch) were compared with those now existing in South America or in Australia. again. genera. North and South America. "If struck by this strange sequence. the general parallelism in the successive forms of life. When the marine forms of life are spoken of as having changed simultaneously throughout the world. for in some cases not one species is identically the same. several highly competent observers maintain that the existing productions of the United States are more closely related to those which lived in Europe during certain late tertiary stages. These observations. which are not found in the Chalk of Europe. It is not that the same species are met with. relate to the marine inhabitants of the world: we have not sufficient data to judge whether the productions of the land and of fresh water at distant points change in the same parallel manner. for if all the marine animals now living in Europe. would still be manifest. Macrauchenia. MM. We may doubt whether they have thus changed: if the Megatherium. Western Europe and North America. Mylodon. and Australia. looking to a remotely future epoch. and all those that lived in Europe during the pleistocene period (a very remote period as measured by years. with the European and North American tertiary deposits. would be correctly ranked as simultaneous in a geological sense. de Verneuil and d'Archiac. it will appear certain that all these modifications of species. In the several successive palaeozoic formations of Russia. no one would have suspected that they had co-existed with sea-shells all still living. the pleistocene and strictly modern beds of Europe. however. but which occur in the formations either above or below. other forms. there can be little doubt that all the more modern MARINE formations. and if this be so. but as these anomalous monsters co-existed with the Mastodon and Horse. and the several formations could be easily correlated. or even that it has a very strict geological sense. namely.

a less strict degree of parallelism in the succession of the productions of the land than with those of the sea. to a certain extent. as far as fossils are concerned. indeed. and of giving rise in new countries to other new varieties and species. climate. This great fact of the parallel succession of the forms of life throughout the world. and the forms. and producing new forms. and the succession of forms everywhere tends to correspond both in their first appearance and final disappearance. old groups disappear from the world. We must. and the introduction of new ones. We shall see this more clearly when we treat of the present distribution of organic beings.extinction. occurred during the periods when the bed of the sea was either stationary or rising. and that blank intervals of vast duration. should be those which would have the best chance of spreading still further. Thus. and on the gradual acclimatization of new species to the various climates through which they might have to pass. in the plants which are dominant. or have some advantage over the other forms in their own country. During these long and blank intervals I suppose that the inhabitants of each region underwent a considerable amount of modification and extinction. New species are formed by having some advantage over older forms. The old forms which are beaten and which yield their places to the new and victorious forms. cannot be owing to mere changes in marine currents or other causes more or less local and temporary. The diffusion would. rich in fossils. We might therefore expect to find. which have already invaded. owing to their having had some advantage over their already dominant parents. Barrande has made forcible remarks to precisely the same effect. as new and improved groups spread throughout the world. It is. under the most different climates. and again spreading. the new species thus produced being themselves dominant. as Barrande has remarked. simultaneous. were deposited during periods of subsidence. that is. and. the territories of other species. be slower with the terrestrial inhabitants of distinct continents than with the marine inhabitants of the continuous sea. on strange accidents. as it seems to me. varying and far-spreading species. There is one other remark connected with this subject worth making. is explicable on the theory of natural selection. as we do find. but depend on general laws which govern the whole animal kingdom. and. and find how slight is the relation between the physical conditions of various countries and the nature of their inhabitants. It is also natural that the dominant. producing the greatest number of new varieties. quite futile to look to changes of currents. The process of diffusion would often be very slow. I have given my reasons for believing that most of our great formations. as well as over other species. succession of the same forms of life throughout the world. which are commonest and most widely diffused. taken in a large sense. look to some special law. the parallel. As we have . give birth to the greatest number of new varieties or incipient species." M. as the cause of these great mutations in the forms of life throughout the world. or other physical conditions. varying. and likewise when sediment was not thrown down quickly enough to embed and preserve organic remains. which are already dominant. it is probable. but in the course of time the dominant forms would generally succeed in spreading and would ultimately prevail. accords well with the principle of new species having been formed by dominant species spreading widely and varying. depending on climatal and geographical changes. and that there was much migration from other parts of the world. We have distinct evidence on this head. from inheriting some inferiority in common. will generally be allied in groups. therefore.

But. When two formations have been deposited in two regions during nearly. extinction. but contemporaneous faunas. All fall into a few grand classes. Another distinguished palaeontologist. it may be well to make some remarks on this subject. as a general rule. in accordance with the general succession of the forms of life. If we confine our attention either to the living or to the extinct species of the same class. although he finds in both a curious accordance in the numbers of the species belonging to the same genera. extinct species can all be classed either in still existing groups. but we are very far from having any right to conclude that this has invariably been the case. In the writings of Professor Owen we continually meet with the expression of generalised forms. unless. M. Mr. and orders. of prophetic or synthetic types. Cuvier ranked the Ruminants and Pachyderms as two of the most distinct orders of mammals. in fact. and to give some instances. and that large areas have invariably been affected by the same movements. That the extinct forms of life help to fill up the intervals between existing genera. is able to draw a close general parallelism between the successive stages in the two countries. Lyell has made similar observations on some of the later tertiary formations. and these terms imply that such forms are. but as this statement has often been ignored or even denied. from the causes explained in the foregoing paragraphs. families. but so many fossil links have been disentombed that Owen has had to alter the . the same general succession in the forms of life. has shown in the most striking manner that many of the fossil mammals discovered by him in Attica serve to break down the intervals between existing genera. as applied to extinct animals. indeed. but not exactly. as Buckland long ago remarked. it is probable that strictly contemporaneous formations have often been accumulated over very wide spaces in the same quarter of the world. we should find in both. also. ON THE AFFINITIES OF EXTINCT SPECIES TO EACH OTHER. yet the species themselves differ in a manner very difficult to account for considering the proximity of the two areas.reason to believe that large areas are affected by the same movement. but when he compares certain stages in England with those in France. in his admirable Memoirs on the eocene deposits of England and France. and immigration. Gaudry. AND TO LIVING FORMS. it differs from living forms. is certainly true. The more ancient any form is. If the several formations in these regions have not been deposited during the same exact periods--a formation in one region often corresponding with a blank interval in the other--and if in both regions the species have gone on slowly changing during the accumulation of the several formations and during the long intervals of time between them. intermediate or connecting links. for there will have been a little more time in the one region than in the other for modification. Barrande. nevertheless the species would not all be the same in the apparently corresponding stages in the two regions. and in the writings of Agassiz. Prestwich. or between them. the same period. and the order would falsely appear to be strictly parallel. shows that there is a striking general parallelism in the successive Silurian deposits of Bohemia and Scandinavia. nevertheless he finds a surprising amount of difference in the species. but the species would not exactly correspond. and this fact is at once explained on the principle of descent. I suspect that cases of this nature occur in Europe. Let us now look to the mutual affinities of extinct and living species. it be assumed that an isthmus separated two seas inhabited by distinct. the series is far less perfect than if we combine both into one general system. in this case the several formations in the two regions could be arranged in the same order. the more.

an ossified thigh-bone "articulated to a well-defined acetabulum in the pelvis. by the ostrich and extinct Archeopteryx. that he is every day taught that. although palaeozoic animals can certainly be classed under existing groups. The cetaceans or whales are widely different from all other mammals. are considered by Professor Huxley to be undoubtedly cetaceans. and it would be difficult to prove the truth of the proposition. and has placed certain Pachyderms in the same sub-order with ruminants. yet that at this ancient period the groups were not so distinctly separated from each other as they now are. The most common case. and some extinct genera between living genera. If by this term it is meant that an extinct form is directly intermediate in all its characters between two living forms or groups. the ancient members are separated by a somewhat lesser number of characters. even between genera belonging to distinct families. is discovered having affinities directed towards very distinct groups. according to Professor Flower. and the eocene Mammals. by so much the more it tends to connect by some of its characters groups now widely separated from each other. but the tertiary Zeuglodon and Squalodon. the older Cephalopods. but the Macrauchenia of South America connects to a certain extent these two grand divisions. the objection is probably valid. being considered as intermediate between any two living species. The Sirenia form a very distinct group of the mammals. But in a natural classification many fossil species certainly stand between living species. he dissolves by gradations the apparently wide interval between the pig and the camel. to which the Sirenia are in other respects allied.whole classification. as the name given to it by Professor Gervais expresses. and on the other hand by the Compsognathus. which have been placed by some naturalists in an order by themselves. and one of the most remarkable peculiarities in existing dugong and lamentin is the entire absence of hind limbs. a higher authority could not be named. . without even a rudiment being left. such as fish and reptiles. we must admit that there is truth in the remark. on the one hand. for example. with the recent members of the same classes. The Ungulata or hoofed quadrupeds are now divided into the even-toed or odd-toed divisions. Barrande asserts. and which cannot be placed in any existing order. Yet if we compare the older Reptiles and Batrachians. No one will deny that the Hipparion is intermediate between the existing horse and certain other ungulate forms. Turning to the Invertebrata." Even the wide interval between birds and reptiles has been shown by the naturalist just quoted to be partially bridged over in the most unexpected manner. the older Fish. or groups of species. Some writers have objected to any extinct species. for every now and then even a living animal. supposing them to be distinguished at the present day by a score of characters. What a wonderful connecting link in the chain of mammals is the Typotherium from South America. one of the Dinosaurians--that group which includes the most gigantic of all terrestrial reptiles. seems to be that." and it thus makes some approach to ordinary hoofed quadrupeds. This remark no doubt must be restricted to those groups which have undergone much change in the course of geological ages. so that the two groups formerly made a somewhat nearer approach to each other than they now do. "and to constitute connecting links with the aquatic carnivora. or group of species. but the extinct Halitherium had. especially with respect to very distinct groups. as the Lepidosiren. It is a common belief that the more ancient a form is.

assume that divergence of character is a necessary contingency. above No. and there would remain two families which would be less distinct from each other than they were before the discovery of the fossils. the genera a1. and the dotted lines diverging from them the species in each genus. from the continued effects of extinction and divergence of character. The three existing genera. be supposed to differ from each other by half-a-dozen important characters. If many extinct forms were to be discovered above one of the middle horizontal lines or geological formations--for instance. together with the many extinct genera on the several lines of descent diverging from the parent form (A) will form an order. as before remarked. descended from (A). q14. a third family. As the subject is somewhat complex. etc. a closely allied family or subfamily. So again. were discovered at several points low down in the series. Hence. in nearly the same manner as has occurred with ruminants and certain pachyderms. for all will have inherited something in common from their ancient progenitor. The horizontal lines may represent successive geological formations. and b14. b14 and f14. All the many forms. If. extinct and recent. would be partly justified. The diagram is much too simple. however. p14. Yet he who objected to consider as intermediate the extinct genera. the more recent any form is the more it will generally differ from its ancient progenitor. not directly. m3.) would have to be united into one. These three families. On the principle of the continued tendency to divergence of character. which thus link together the living genera of three families. but this is unimportant for us. were disinterred. This is represented in the diagram by the letter F14. then only two of the families (those on the left hand a14. and yet retain throughout a vast period the same general characteristics. one order. for they would at this early stage of descent have diverged in a less degree from their common progenitor. if the three families formed of eight genera (a14 to m14). f8. and o14. we can understand the rule that the most ancient fossils differ most from existing forms. m9.Let us see how far these several facts and inferences accord with the theory of descent with modification. i14. and this order. but only by a long and circuitous course through many widely different forms. Thus it comes that ancient and extinct genera are often in a greater or less degree intermediate . a5. on the uppermost line. too few genera and too few species being given. VI. the three existing families on the uppermost line would be rendered less distinct from each other. We may suppose that the numbered letters in italics represent genera. that a species might go on being slightly modified in relation to its slightly altered conditions of life. We must not. Therefore it is quite possible. for instance. and all the forms beneath the uppermost line may be considered as extinct. which was formerly illustrated by this diagram. etc. then the families which existed at a period marked VI would certainly have differed from each other by a less number of characters. as we have seen in the case of some Silurian forms. for they are intermediate. m6. it depends solely on the descendants from a species being thus enabled to seize on many and different places in the economy of nature. will form a small family. By looking at the diagram we can see that if many of the extinct forms supposed to be embedded in the successive formations. m14. these three families would be so closely linked together that they probably would have to be united into one great family. a14. I must request the reader to turn to the diagram in the fourth chapter. a10. and some to have endured to the present day.. some of which are supposed to have perished at different periods. make. has become divided into several sub-families and families.--but none from beneath this line.

and this by the concurrent evidence of our best palaeontologists is frequently the case. Subject to these allowances. they will have endured for extremely unequal lengths of time. and for a large amount of modification during the long and blank intervals between the successive formations. to fill up the wide intervals in the natural system. and are the parents of those which became still more modified at the seventh stage. But supposing for an instant. should in the older formations make some slight approach to each other. so that the older members should differ less from each other in some of their characters than do the existing members of the same groups. All that we have a right to expect is. in this and other such cases. For instance. when arranged by Dr. undergone much modification. It is no real objection to the truth of the statement that the fauna of each period as a whole is nearly intermediate in character between the preceding and succeeding faunas. And they are wholly inexplicable on any other view. The species extreme in character are not the oldest or the most recent. the main facts with respect to the mutual affinities of the extinct forms of life to each other and to living forms. were at once recognised by palaeontologists as intermediate in character between those of the overlying carboniferous and underlying Silurian systems. As we possess only the last volume of the geological record. hence they could hardly fail to be nearly intermediate in character between the forms of life above and below. allow for the entire extinction of some preceding forms. on the theory of descent with modification. the manner in which the fossils of the Devonian system. it is evident that the fauna during any one great period in the earth's history will be intermediate in general character between that which preceded and that which succeeded it. which is far from the case. especially in the case of terrestrial productions inhabiting separated districts. for the groups will have been more numerous. Thus. within known geological periods. But each fauna is not necessarily exactly intermediate. and will have been modified in various degrees. we have no right to expect. that those groups which have. intermediate in age. as unequal intervals of time have elapsed between consecutive formations. Falconer in two series--in the first place according to their mutual affinities. and in the second place according to their periods of existence--do not accord in arrangement. between the preceding and succeeding faunas. and thus to unite distinct families or orders. we have no reason to believe that forms successively produced necessarily endure for corresponding lengths of time. that the record of the first appearance and disappearance of the species was complete. To compare small things with . We must. A very ancient form may occasionally have lasted much longer than a form elsewhere subsequently produced. and that in a very broken condition. are explained in a satisfactory manner. namely. I need give only one instance. On this same theory. Under nature the process will be far more complicated than is represented in the diagram. or between their collateral relations. the fauna of each geological period undoubtedly is intermediate in character. and in any one region for the immigration of new forms from other regions. however. that certain genera offer exceptions to the rule. Thus the species which lived at the sixth great stage of descent in the diagram are the modified offspring of those which lived at the fifth stage. nor are those which are intermediate in character. except in rare cases.in character between their modified descendants. when this system was first discovered. the species of mastodons and elephants.

and as long blank intervals have intervened between successive formations. though ranked as distinct species. as I attempted to show in the last chapter. In another and more general manner. have changed almost simultaneously throughout the world. though the species are distinct in each stage. Consider the prodigious vicissitudes of climate during the pleistocene period. ON THE STATE OF DEVELOPMENT OF ANCIENT COMPARED WITH LIVING FORMS. as the specialisation of parts is an advantage to each being. the full meaning of the fossil remains from closely consecutive formations. at least those inhabiting the sea. On the theory of descent. is the fact. we ought not to expect to find. seems to have shaken Professor Pictet in his belief in the immutability of species. Pictet gives as a well-known instance. Closely connected with the statement. or. We have also seen that. is the best standard.great. and in some cases will even degrade or simplify the organisation. representative species. and many varieties between the rock-pigeon and the carrier have become extinct. and therefore under the most different climates and conditions. and these assuredly we do find. not but that it may leave many creatures with simple and unimproved structures fitted for simple conditions of life. with which they come into close competition. for the parent rock-pigeon still lives. We find. all the intermediate varieties between the species which appeared at the commencement and close of these periods: but we ought to find after intervals. is obvious. As the accumulation of each formation has often been interrupted. which includes the whole glacial epoch. in short. yet leaving such degraded beings better fitted for their new walks of life. this arrangement would not closely accord with the order in time of their production. and note how little the specific forms of the inhabitants of the sea have been affected. and carriers which are extreme in the important character of length of beak originated earlier than short-beaked tumblers. new species become superior to their predecessors. if the principal living and extinct races of the domestic pigeon were arranged in serial affinity. as yet suggested. closely allied forms. This fact alone. very long as measured by years. for they have to beat in the struggle for life all the older forms. that the organic remains from an intermediate formation are in some degree intermediate in character. He who is acquainted with the distribution of existing species over the globe. that fossils from two consecutive formations are far more closely related to each other. such evidence of the slow and scarcely sensible mutations of specific forms. as we have the right to expect. which are at the opposite end of the series in this respect. by the physical conditions of the ancient areas having remained nearly the same. being closely related. and in this sense higher. and even less with the order of their disappearance. We have seen in the fourth chapter that the degree of differentiation and specialisation of the parts in organic beings. but only moderately long as measured geologically. from its generality. We may therefore conclude that if under a nearly similar climate the eocene . the general resemblance of the organic remains from the several stages of the Chalk formation. in any one or in any two formations. of their degree of perfection or highness. so natural selection will tend to render the organisation of each being more specialised and perfect. insisted on by all palaeontologists. will not attempt to account for the close resemblance of distinct species in closely consecutive formations. when arrived at maturity. than are the fossils from two remote formations. as they have been called by some authors. Let it be remembered that the forms of life.

at all times imperfect. for some organisms would have to remain fitted for simple conditions of life. but formerly selaceans and ganoids alone existed. have to be slightly modified. It is not an insuperable difficulty that Foraminifera have not. and such crustaceans. looking to members of the same class. for in this case. though difficult of proof. others look at the teleosteans as the highest. for instance. as far as is known. Carpenter. the former would be beaten and exterminated by the latter. and it seems that this answer must be admitted as true. although upon another type?" In the complex struggle for life it is quite credible that crustaceans. so as to hold their places in relation to slight changes in their conditions. who will decide whether a cuttle-fish be higher than a bee--that insect which the great Von Baer believed to be "in fact more highly organised than a fish. from their approach in some important points of structure to reptiles. could be proved to have first come into existence during the Laurentian epoch. When advanced up to any given point. if it included advance in organisation as a necessary contingent. high . on the theory of natural selection. progressed in organisation since even the Laurentian epoch. The ganoids stand intermediate between the selaceans and teleosteans. and the palaeozoic by the secondary forms. They would likewise be fatal. Is this the case? A large majority of palaeontologists would answer in the affirmative. that certain Brachiopods have been but slightly modified from an extremely remote geological epoch. the latter at the present day are largely preponderant in number. as would the secondary by the eocene. not very high in their own class. as well as by the standard of the specialisation of organs. to stand higher than ancient forms. modern forms ought. if judged by the most decisive of all trials--the law of battle. So that by this fundamental test of victory in the battle for life. naturalists are not unanimous which forms ought to be ranked as highest: thus. and at what period the various forms of life first appeared. there would not have been time sufficient for the development of these organisms up to the standard which they had then reached. might beat cephalopods. we ought not solely to compare the highest members of a class at any two periods--though undoubtedly this is one and perhaps the most important element in striking a balance--but we ought to compare all the members. The geological record. and this may well be disputed. for their further continued process. if the above Foraminifera. though not highly developed. Beside these inherent difficulties in deciding which forms are the most advanced in organisation. though they will. The problem whether organisation on the whole has advanced is in many ways excessively intricate. and that certain land and fresh-water shells have remained nearly the same. some look at the selaceans or sharks. the highest molluscs. as insisted on by Dr. there is no necessity. from the time when. on the theory of natural selection. according to the standard of highness chosen. so will it be said that fishes have advanced or retrograded in organisation. or the above Brachiopods during the Cambrian formation. would stand very high in the scale of invertebrate animals. The foregoing objections hinge on the question whether we really know how old the world is. does not extend far enough back to show with unmistakable clearness that within the known history of the world organisation has largely advanced. they first appeared. Even at the present day.inhabitants of the world could be put into competition with the existing inhabitants. It is no valid objection to this conclusion. and what could be better fitted for this end than these lowly organised Protozoa? Such objections as the above would be fatal to my view. as the highest fish. To attempt to compare members of distinct types in the scale of highness seems hopeless. during each successive age. and in this case.

by considering the vast reduction of brachiopods. the standard of organisation of the imperfectly-known faunas of successive periods. At an ancient epoch the highest and lowest molluscoidal animals. of the former and less modified condition of the species. for instance. at the present day fifty thousand kinds of vertebrate animals exist. a multitude of British forms would in the course of time become thoroughly naturalized there. that if all the animals and plants of Great Britain were set free in New Zealand. namely. consequently some naturalists maintain that molluscs were formerly more highly developed than at present. From the extraordinary manner in which European productions have recently spread over New Zealand. as a decided advance in the organisation of the world. DURING THE LATER TERTIARY PERIODS. and having been inherited at a corresponding age. from the fact that hardly a single inhabitant of the southern hemisphere has become wild in any part of Europe. that the oldest known mammals. intermediate in organisation. reptiles. at the present time both groups are greatly reduced. we may well doubt whether. of the high and low classes throughout the world: if. are more highly organised than their ancient representatives. if all the productions of New Zealand were set free in Great Britain. more and more difference to the adult. and the fact that our existing cephalopods. while others. continually adds. we ought to look at this increase in number in the highest class. which implies a great displacement of lower forms. owing to variations having supervened at a not early age. until beds rich in fossils are discovered far beneath the lowest Cambrian strata--a discovery of which the chance is small. have largely increased. and fishes strictly belong to their proper classes. ON THE SUCCESSION OF THE SAME TYPES WITHIN THE SAME AREAS. and have seized on places which must have been previously occupied by the indigenes. Yet the most skilful naturalist. and that the geological succession of extinct forms is nearly parallel with the embryological development of existing forms. and if we knew that at some former period only ten thousand kinds existed. from an examination of the species of the two countries. and yet may never be capable of proof. whilst it leaves the embryo almost unaltered. On the other hand. We thus see how hopelessly difficult it is to compare with perfect fairness. In a future chapter I shall attempt to show that the adult differs from its embryo. though few in number. swarmed in numbers. Under this point of view. This process. cephalopods and brachiopods. Seeing.and low. at any two periods. This view accords admirably well with our theory. We ought also to compare the relative proportional numbers. This view may be true. and would exterminate many of the natives. Thus the embryo comes to be left as a sort of picture. . we must believe. at two periods. though some of these old forms are in a slight degree less distinct from each other than are the typical members of the same groups at the present day. for instance. under such extremely complex relations. in the course of successive generations. could not have foreseen this result. but a stronger case can be made out on the opposite side. We shall appreciate this difficulty more clearly by looking to certain existing faunas and floras. any considerable number would be enabled to seize on places now occupied by our native plants and animals. the productions of Great Britain stand much higher in the scale than those of New Zealand. preserved by nature. it would be vain to look for animals having the common embryological character of the Vertebrata. Agassiz and several other highly competent judges insist that ancient animals resemble to a certain extent the embryos of recent animals belonging to the same classes.

for the dissimilarity of the inhabitants of these two continents. buried there in such numbers. the great law of the long enduring. a similar relationship is manifest. and Professor Owen has shown in the most striking manner that most of the fossil mammals. even to an uneducated eye. Analogous facts could be given in relation to the distribution of marine animals. Nor can it be pretended that it is an immutable law that marsupials should have been chiefly or solely produced in Australia. and between the extinct and living brackish water-shells of the Aralo-Caspian Sea. after comparing the present climate of Australia and of parts of South America. for the inhabitants of each quarter of the world will obviously tend to leave in that quarter. so will their modified descendants still differ in nearly the same manner and degree. but. permitting much intermigration. Lund and Clausen in the caves of Brazil. succession of the same types within the same areas. have . what does this remarkable law of the succession of the same types within the same areas mean? He would be a bold man who. and after great geographical changes. but not immutable. the feebler will yield to the more dominant forms. that in America the law of distribution of terrestrial mammals was formerly different from what it now is. for the uniformity of the same types in each continent during the later tertiary periods. For we know that Europe in ancient times was peopled by numerous marsupials. found in several parts of La Plata. Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that continent. I was so much impressed with these facts that I strongly insisted. We see the same law in this author's restorations of the extinct and gigantic birds of New Zealand. In South America. and. In a similar manner we know. like those of the armadillo. are related to South American types. and I have shown in the publications above alluded to. that Northern India was formerly more closely related in its mammals to Africa than it is at the present time. as the relation between the extinct and living land-shells of Madeira. from Falconer and Cautley's discoveries. to the northern half. closely allied though in some degree modified descendants. and the southern half was formerly more closely allied. on the one hand through dissimilar physical conditions.Mr. and there will be nothing immutable in the distribution of organic beings. But after very long intervals of time. than it is at present. on this "law of the succession of types. would attempt to account. which formerly lived in South America. This relationship is even more clearly seen in the wonderful collection of fossil bones made by MM. from the wide distribution of most molluscs."--on "this wonderful relationship in the same continent between the dead and the living. during the next succeeding period of time. It may be asked in ridicule whether I suppose that the megatherium and other allied huge monsters. it is not well displayed by them. Mr." Professor Owen has subsequently extended the same generalisation to the mammals of the Old World. Now. in 1839 and 1845. is at once explained. Woodward has shown that the same law holds good with sea-shells. If the inhabitants of one continent formerly differed greatly from those of another continent. or that Edentata and other American types should have been solely produced in South America. in the gigantic pieces of armour. North America formerly partook strongly of the present character of the southern half of the continent. Other cases could be added. on the other hand through similarity of conditions. under the same latitude. We see it also in the birds of the caves of Brazil. On the theory of descent with modification.

that each single formation has not been continuously deposited. each with the same number of species. although each species must have passed through numerous transitional stages.left behind them the sloth. and have oftenest given rise to new species. unless the whole chain could be perfectly restored. and in a succeeding formation there be six other allied or representative genera. that only a small portion of the globe has been geologically explored with care. on our theory. armadillo. connecting together all extinct and existing forms by the finest graduated steps. and more variation during the periods of elevation. and during the latter the record will have been least perfectly kept. that widely ranging species are those which have varied most frequently. that the number both of specimens and of species. For he may ask in vain where are the numberless transitional links which must formerly have connected the . during which each underwent modification. I have attempted to show that the geological record is extremely imperfect. it is probable that the periods. These causes. that varieties have at first been local. In failing orders. and some of these fossils may have been the actual progenitors of the living species. for it is not pretended that we have any sure criterion by which species and varieties can be discriminated. It must not be forgotten that. and anteater. if six genera. that only certain classes of organic beings have been largely preserved in a fossil state. still fewer genera and species will leave modified blood-descendants. with the genera and species decreasing in numbers as is the case with the Edentata of South America. then we may conclude that generally only one species of each of the older genera has left modified descendants. that. which might be found. that there has probably been more extinction during the periods of subsidence. so that. which constitute the new genera containing the several species. and thick enough to outlast future degradation. preserved in our museums. It should also be constantly borne in mind that any linking variety between two forms. as a new and distinct species. and lastly. will to a large extent explain why--though we do find many links--we do not find interminable varieties. and have left no progeny. two or three species in two or three alone of the six older genera will be the parents of the new genera: the other species and the other old genera having become utterly extinct. But in the caves of Brazil there are many extinct species which are closely allied in size and in all other characters to the species still living in South America. This cannot for an instant be admitted. Or. taken conjointly. and this will be a far commoner case. great intervals of time must have elapsed between most of our successive formations. as their degenerate descendants. that migration has played an important part in the first appearance of new forms in any one area and formation. each having eight species. will rightly reject the whole theory. the other seven species of each old genus having died out and left no progeny. would be ranked. SUMMARY OF THE PRECEDING AND PRESENT CHAPTERS. owing to subsidence being almost necessary for the accumulation of deposits rich in fossil species of many kinds. have been short in comparison with the periods during which each remained in an unchanged condition. though many and long as measured by years. These huge animals have become wholly extinct. be found in one geological formation. that the duration of each formation is probably short compared with the average duration of specific forms. He who rejects this view of the imperfection of the geological record. all the species of the same genus are the descendants of some one species. is absolutely as nothing compared with the number of generations which must have passed away even during a single formation.

the more it generally differs from those now living. The extinction of old forms is the almost inevitable consequence of the production of new forms. Why ancient and extinct forms often tend to fill up gaps between existing forms. the other great leading facts in palaeontology agree admirably with the theory of descent with modification through variation and natural selection. the species of the less vigorous groups. Passing from these difficulties. and where our oscillating continents now stand they have stood since the commencement of the Cambrian system. from the survival of a few descendants. Groups of species increase in numbers slowly. We can understand why. the world presented a widely different aspect. and consequently resemble. the more often it stands in some degree intermediate between groups now distinct. from their inferiority inherited from a common progenitor. or lie still buried under the ocean. the more nearly it will be related to. and depends on many complex contingencies. he may overlook how important a part migration has played. exist now only as remnants in a metamorphosed condition. and these will generally succeed in displacing the groups which are their inferiors in the struggle for existence. the common progenitor of groups. The more ancient a form is. after long intervals of time. But the utter extinction of a whole group of species has sometimes been a slow process. which form new sub-groups and groups. but more commonly bringing them only a little closer together. but often falsely apparent. tend to become extinct together. it never reappears. when a species has once disappeared. We can understand how it is that all the forms of life. make together a few grand classes. or at the same rate. and endure for unequal periods of time. for the link of generation has been broken. since become widely divergent. We can thus understand how it is that new species come in slowly and successively. from the continued tendency to divergence of character. When a group has once wholly disappeared. but are intermediate only by a long and circuitous course through other extinct . how species of different classes do not necessarily change together. We can understand how it is that dominant forms which spread widely and yield the greatest number of varieties tend to people the world with allied. The dominant species belonging to large and dominant groups tend to leave many modified descendants. Hence. but modified. are considered. long before that epoch. We can understand. yet in the long run that all undergo modification to some extent. but I can answer this last question only by supposing that where our oceans now extend they have extended for an enormous period. Extinct forms are seldom directly intermediate between existing forms. He may ask where are the remains of those infinitely numerous organisms which must have existed long before the Cambrian system was deposited? We now know that at least one animal did then exist. As these are formed. formed of formations older than any known to us. sudden coming in of whole groups of species. and that the older continents. for the more ancient a form is. sometimes blending two groups. he may urge the apparent. found in the successive stages of the same great formation? He may disbelieve in the immense intervals of time which must have elapsed between our consecutive formations. descendants. why the more ancient a form is. previously classed as distinct into one.closely allied or representative species. lingering in protected and isolated situations. it does not reappear. but that. the productions of the world appear to have changed simultaneously. as those of Europe. ancient and recent. for the process of modification is necessarily slow. or in the same degree. and to leave no modified offspring on the face of the earth. when the formations of any one great region.

GEOGRAPHICAL DISTRIBUTION. humid districts. marshes. in so far. from the central parts of the United States to its extreme southern point. we meet with the most diversified conditions. grassy plains. forests. of which the conditions are peculiar in only a slight degree. for they are closely linked together by generation. the products of variation and the survival of the fittest. Notwithstanding this general parallelism in the conditions of Old and . but these are not inhabited by a fauna different from that of the surrounding districts. The succession of the same types of structure within the same areas during the later geological periods ceases to be mysterious. as it seems to me. all authors agree that one of the most fundamental divisions in geographical distribution is that between the New and Old Worlds. higher in the scale. that species have been produced by ordinary generation: old forms having been supplanted by new and improved forms of life. There is hardly a climate or condition in the Old World which cannot be paralleled in the New--at least so closely as the same species generally require. yet if we travel over the vast American continent. The case of America alone would almost suffice to prove its truth. under almost every temperature. Present distribution cannot be accounted for by differences in physical conditions--Importance of barriers--Affinity of the productions of the same continent--Centres of creation--Means of dispersal by changes of climate and of the level of the land. If. The inhabitants of the world at each successive period in its history have beaten their predecessors in the race for life. Of late.and different forms. and by occasional means--Dispersal during the Glacial period--Alternate Glacial periods in the North and South. the geological record be as imperfect as many believe. the first great fact which strikes us is. arid deserts. and their structure has generally become more specialised. No doubt small areas can be pointed out in the Old World hotter than any in the New World. Extinct and ancient animals resemble to a certain extent the embryos of the more recent animals belonging to the same classes. and this may account for the common belief held by so many palaeontologists. and is intelligible on the principle of inheritance. lakes and great rivers. In considering the distribution of organic beings over the face of the globe. and are. CHAPTER XII. the main objections to the theory of natural selection are greatly diminished or disappear. We can clearly see why the remains of an intermediate formation are intermediate in character. and this wonderful fact receives a simple explanation according to our views. that organisation on the whole has progressed. On the other hand. almost every author who has studied the subject has come to this conclusion. then. lofty mountains. all the chief laws of palaeontology plainly proclaim. that neither the similarity nor the dissimilarity of the inhabitants of various regions can be wholly accounted for by climatal and other physical conditions. and it may at least be asserted that the record cannot be proved to be much more perfect. for it is rare to find a group of organisms confined to a small area. We can clearly see why the organic remains of closely consecutive formations are closely allied. for if we exclude the arctic and northern temperate parts.

we find different productions. and this fact has led naturalists to believe that the isthmus was formerly open. which consequently are separated by a space of ten degrees of latitude. after travelling over a hemisphere. yet they are incomparably more closely related to each other than they are to the productions of Australia or Africa under nearly the same climate. A second great fact which strikes us in our general review is. until. South Africa. where the land almost joins. with extremely few shells. A third great fact. or echinodermata in common. yet it would not be possible to point out three faunas and floras more utterly dissimilar. The marine inhabitants of the eastern and western shores of South America are very distinct. and western South America. Africa. how widely different are their living productions! In the southern hemisphere. for on the opposite sides of lofty and continuous mountain-ranges. and of great deserts and even of large rivers. and where.New Worlds. a wide space of open ocean extends. On the other hand. yet many fishes range from the Pacific into the Indian Ocean. partly included in the foregoing statement. So that three marine faunas range northward and southward in parallel lines not far from each other. again. that barriers of any kind. etc. crustacea. under a slightly different climate. deserts. Analogous facts could be given with respect to the inhabitants of the sea. Although so few marine animals are common to the above-named three approximate faunas of Eastern and Western America and the eastern Pacific islands. We see the same fact in the great difference between the inhabitants of Australia. and South America under the same latitude. with not an island as a halting-place for emigrants. or continuous coasts. and many shells are common to the eastern islands of the Pacific and the eastern shores of Africa on almost exactly opposite meridians of longitude. also. either of land or open sea. we shall find parts extremely similar in all their conditions. and over this vast space we meet with no well-defined and distinct marine faunas. excepting in the northern parts. though as mountain chains. as the oceans separating continents. they are almost wholly distinct. or obstacles to free migration. but Dr. here we have a barrier of another kind.. or likely to have endured so long. there might have been free migration for the northern temperate forms. proceeding still further westward from the eastern islands of the tropical parts of the Pacific. and are exposed to considerably different conditions. we come to the shores of Africa. we encounter no impassable barriers. Turning to the sea. We see this in the great difference in nearly all the terrestrial productions of the New and Old Worlds. as there now is for the strictly arctic productions. and we have innumerable islands as halting-places. On each continent. are not as impassable. Or. are related in a close and important manner to the differences between the productions of various regions. is the . but from being separated from each other by impassable barriers. Westward of the shores of America. if we compare large tracts of land in Australia. for these countries are almost as much isolated from each other as is possible. and as soon as this is passed we meet in the eastern islands of the Pacific with another and totally distinct fauna. between latitudes 25 and 35 degrees. under corresponding climate. the differences are very inferior in degree to those characteristic of distinct continents. we see the same fact. we find the same law. Gunther has recently shown that about thirty per cent of the fishes are the same on the opposite sides of the isthmus of Panama. we may compare the productions of South America south of latitude 35 degrees with those north of 25 degrees.

from north to south. rodents of the South American type. We may look back to past ages.affinity of the productions of the same continent or of the same sea. but not quite alike. as far as we positively know. replace each other. and we find an alpine species of bizcacha. If we look to the islands off the American shore. over the same areas of land and water. we look to the waters. As the variability of each species is an independent property. and belonging to the same order of Rodents. though the species themselves are distinct at different points and stations. He hears from closely allied. that cause which alone. the naturalist. and thus they will become still further victorious. and we do not find the beaver or muskrat. abounding in individuals. when they spread out into new countries. animals having nearly the same habits as our hares and rabbits. throughout space and time. as shown in the last chapter. and we find American types then prevailing on the American continent and in the American seas. will have the best chance of seizing on new places. nearly alike. On this principle of inheritance with modification we can understand how it is that sections of genera. produces organisms quite like each other. and northward the plains of La Plata by another species of the same genus. the relation of organism to organism in the struggle for life being. however much they may differ in geological structure. only so far as it profits each individual in its . but they plainly display an American type of structure. and every continent offers innumerable instances. The naturalist must be dull who is not led to inquire what this bond is. We ascend the lofty peaks of the Cordillera. independently of physical conditions. and will produce groups of modified descendants. though nearly related. The bond is simply inheritance. and probably in a subordinate degree to the definite influence of different physical conditions. as was remarked in the last chapter. for instance. specifically distinct. though they may be all peculiar species. with eggs coloured in nearly the same manner. and not by a true ostrich or emu. as we see in the case of varieties. never fails to be struck by the manner in which successive groups of beings. at periods more or less remote--on the nature and number of the former immigrants--and on the action of the inhabitants on each other in leading to the preservation of different modifications. the inhabitants are essentially American. or. and will be taken advantage of by natural selection. which have already triumphed over many competitors in their own widely-extended homes. It is a law of the widest generality. are confined to the same areas. yet distinct kinds of birds. the most important of all relations. but the coypu and capybara. in travelling. In their new homes they will be exposed to new conditions. as is so commonly and notoriously the case. Innumerable other instances could be given. and even families. The degrees of dissimilarity will depend on the migration of the more dominant forms of life from one region into another having been more or less effectually prevented. The dissimilarity of the inhabitants of different regions may be attributed to modification through variation and natural selection. We see in these facts some deep organic bond. There is no evidence. and sees their nests similarly constructed. Widely-ranging species. and will frequently undergo further modification and improvement. whole genera. like those inhabiting Africa and Australia under the same latitude. notes nearly similar. as I have already often remarked. as does time for the slow process of modification through natural selection. of the existence of any law of necessary development. On these same plains of La Plata we see the agouti and bizcacha. Thus the high importance of barriers comes into play by checking migration. Nevertheless. The plains near the Straits of Magellan are inhabited by one species of Rhea (American ostrich).

they would be little liable to modification. namely. for neither migration nor isolation in themselves effect anything. so the amount of modification in different species will be no uniform quantity. where their parents were first produced: for. almost any amount of migration is possible. as they are descended from the same progenitor. where now found. He who rejects it. that the space could not have been easily passed over by migration. Nevertheless the simplicity of the view that each species was first produced within a single region captivates the mind. from their varied means of dispersal. As we have seen in the last chapter that some forms have retained nearly the same character from an enormously remote geological period. is. No geologist feels any difficulty in Great Britain possessing the same quadrupeds with the rest of Europe. rejects the vera causa of ordinary generation with subsequent migration. little modification. and that when a plant or animal inhabits two points so distant from each other. and. It is universally admitted. or with an interval of such a nature. whether species have been created at one or more points of the earth's surface. must have proceeded from one spot. must originally have proceeded from the same source. it is incredible that individuals identically the same should have been produced from parents specifically distinct. that in most cases the area inhabited by a species is continuous. though now inhabiting distant and isolated regions. In the case of those species which have undergone. there is not much difficulty in believing that they have migrated from the same region. The incapacity of migrating across a wide sea is more clear in the case of terrestrial mammals than perhaps with any other organic beings. in which we have reason to believe that the species of a genus have been produced within comparatively recent times. whereas some plants. there is great difficulty on this head.complex struggle for life. and have not become greatly or at all modified. it is obvious that the several species of the same genus. for they were no doubt once united. as I believe. were to migrate in a body into a new and afterwards isolated country. These principles come into play only by bringing organisms into new relations with each other and in a lesser degree with the surrounding physical conditions. accordingly. we find no inexplicable instances of the same mammals inhabiting distant points of the world. though inhabiting the most distant quarters of the world. It is also obvious that the individuals of the same species. so that a multitude of European animals and plants have become naturalised in America and Australia. after having long competed with each other in their old home. We are thus brought to the question which has been largely discussed by naturalists. and some of the aboriginal plants are identically the same at these distant points of the northern and southern hemispheres? The answer. so certain species have migrated over vast spaces. and calls in the agency of a miracle. SINGLE CENTRES OF SUPPOSED CREATION. as has been explained. The . If a number of species. for during the vast geographical and climatical changes which have supervened since ancient times. But in many other cases. the fact is given as something remarkable and exceptional. Undoubtedly there are many cases of extreme difficulty in understanding how the same species could possibly have migrated from some one point to the several distant and isolated points. have migrated across the wide and broken interspaces. According to these views. during whole geological periods. why do we not find a single mammal common to Europe and Australia or South America? The conditions of life are nearly the same. But if the same species can be produced at two separate points. that mammals have not been able to migrate.

and to the various occasional means of transport. inexplicable on the theory of independent creation. as it has to many other naturalists. But the geographical and climatical changes which have certainly occurred within recent geological times. would still be related by inheritance to the inhabitants of that continent. that the view of each species having been produced in one area alone. can have migrated. confined to some one region! Hence. when most of the species inhabiting one region are different from those of another region. namely. What a strange anomaly it would be if a directly opposite rule were to prevail when we go down one step lower in the series. would probably receive from it in the course of time a few colonists. and at distant points in the Arctic and Antarctic regions. It would be hopelessly tedious to discuss all the exceptional cases of the same species. considering our ignorance with respect to former climatical and geographical changes. and of so grave a nature. a still greater number of sections of genera. namely. that each species has been produced within one area. nor do I for a moment pretend that any explanation could be offered of many instances. for the explanation is obvious on the principle of descent with modification. now living at distant and separated points. for instance. and these had not been.great and striking influence of barriers of all kinds. are generally confined to the same country. though separated by hundreds of miles of open sea. In discussing this subject we shall be enabled at the same time to consider a point equally important for us. If. is the most probable. very many genera. and their descendants. does not differ much from . upheaved and formed at the distance of a few hundreds of miles from a continent. and it has been observed by several naturalists that the most natural genera. rendered probable by general considerations. and thirdly. or those genera in which the species are most closely related to each other. and have not been able to migrate to the opposite side. though closely allied to them. and having subsequently migrated from that area as far as its powers of migration and subsistence under past and present conditions permitted. though modified. many subfamilies. So that we are reduced to consider whether the exceptions to continuity of range are so numerous. whether the several species of a genus which must on our theory all be descended from a common progenitor. as we shall hereafter see. the wide distribution of fresh water productions. are confined to a single region. Undoubtedly many cases occur in which we cannot explain how the same species could have passed from one point to the other. Cases of this nature are common. and are. and has migrated thence as far as it could. at least at first. our general view will be much strengthened. the occurrence of the same terrestrial species on islands and on the nearest mainland. Some few families. namely to the individuals of the same species. If the existence of the same species at distant and isolated points of the earth's surface can in many instances be explained on the view of each species having migrated from a single birthplace. the existence of the same species on the summits of distant mountain ranges. or if they have a wide range that their range is continuous. the belief that a single birthplace is the law seems to me incomparably the safest. But. undergoing modification during their migration from some one area. A volcanic island. is intelligible only on the view that the great majority of species have been produced on one side. must have rendered discontinuous the formerly continuous range of many species. I will discuss a few of the most striking classes of facts. it can be shown that migration from the one region to the other has probably occurred at some former period. it seems to me. then. after some preliminary remarks. and secondly (in the following chapter). that we ought to give up the belief. This view of the relation of the species of one region to those of another.

that advanced by Mr. as some authors suppose. might have been a high road for migration. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two marine faunas. however. when the climate was different. and removes many a difficulty. A region now impassable to certain organisms from the nature of its climate. but have never blended with other individuals or varieties of the same species. I shall. who concludes that "every species has come into existence coincident both in space and time with a pre-existing closely allied species. Wallace. Sir C. presently have to discuss this branch of the subject in some detail. which I have selected as presenting the greatest amount of difficulty on the theory of "single centres of creation. No geologist disputes that great mutations of level have occurred within the period of existing organisms. as to have united them within the recent period to each other and to the several intervening oceanic . but to continued care in the selecting and training of many individuals during each generation. at each successive stage of modification. This view cuts the Gordian knot of the dispersal of the same species to the most distant points. but not of such vast changes in the position and extension of our continents. so that. or single hermaphrodite. If. must be descended from a succession of modified varieties. if such exist. submerge it. The question of single or multiple centres of creation differs from another though allied question. that have supplanted each other. or which occasionally intercross. and the two faunas will now blend together. Edward Forbes insisted that all the islands in the Atlantic must have been recently connected with Europe or Africa. each species. and united almost every island with some mainland. To illustrate what I mean: our English race-horses differ from the horses of every other breed. Lyell and other authors have ably treated this subject. it must be admitted that scarcely a single island exists which has not recently been united to some continent. with all organisms which habitually unite for each birth. and Europe likewise with America. but to the best of my judgment we are not authorized in admitting such enormous geographical changes within the period of existing species. and the whole amount of modification at each stage will not be due to descent from a single parent. the individuals of the same species inhabiting the same area will be kept nearly uniform by intercrossing. land may at a former period have connected islands or possibly even continents together. but they do not owe their difference and superiority to descent from any single pair. Where the sea now extends. the arguments used by Forbes are to be trusted. Other authors have thus hypothetically bridged over every ocean. from many individuals simultaneously created." And it is now well known that he attributes this coincidence to descent with modification. I can give here only the briefest abstract of the more important facts. With organic beings which never intercross. or may formerly have blended. and thus have allowed terrestrial productions to pass from one to the other. so that many individuals will go on simultaneously changing. or let it formerly have been submerged. Before discussing the three classes of facts. MEANS OF DISPERSAL. Change of climate must have had a powerful influence on migration." I must say a few words on the means of dispersal. But in the great majority of cases. namely. indeed. all the individuals of the same form will be descended from a single parent. whether all the individuals of the same species are descended from a single pair. or whether. It seems to me that we have abundant evidence of great oscillations in the level of the land or sea. namely.

I shall here confine myself to plants. Afterwards I tried some larger fruits. which may have served as halting places for plants and for many animals during their migration. capsules. but the greater or less facilities for transport across the sea may be said to be almost wholly unknown. The nature and relative proportions of the inhabitants of oceanic islands are likewise opposed to the belief of their former continuity of continents. ripe hazel-nuts sank immediately. whilst green. when . and afterwards when planted germinated. It is well known what a difference there is in the buoyancy of green and seasoned timber. whether or not they were injured by salt water. this or that plant is often stated to be ill adapted for wide dissemination. with one exception. and. being in part determined (as we shall hereafter see) by the depth of the intervening ocean--these and other such facts are opposed to the admission of such prodigious geographical revolutions within the recent period. etc. and it occurred to me that floods would often wash into the sea dried plants or branches with seed-capsules or fruit attached to them. I must now say a few words on what are called accidental means. now buried beneath the sea. that each species has proceeded from a single birthplace. it was not even known how far seeds could resist the injurious action of sea-water. they resisted the salt-water badly. were all killed by a month's immersion. The majority sank quickly. Several facts in distribution--such as the great difference in the marine faunas on the opposite sides of almost every continent--the close relation of the tertiary inhabitants of several lands and even seas to their present inhabitants--the degree of affinity between the mammals inhabiting islands with those of the nearest continent. I freely admit the former existence of many islands. Nor does the almost universally volcanic composition of such islands favour the admission that they are the wrecks of sunken continents. and as all of these sank in a few days. but when dried they floated for ninety days.. some at least of the islands would have been formed. To my surprise I found that out of eighty-seven kinds. an asparagus plant with ripe berries floated for twenty-three days. as are necessary on the view advanced by Forbes and admitted by his followers. metamorphic schists. for instance. and the seeds afterwards germinated: the ripe seeds of Helosciadium sank in two days. but some which. but which more properly should be called occasional means of distribution. In botanical works. Berkeley's aid. and a few survived an immersion of 137 days. floated for a very short time. we shall be enabled to speculate with security on the former extension of the land. or almost continuously united with each other. and some of these floated for a long time. sixty-four germinated after an immersion of twenty-eight days. with Mr. a few experiments. if they had originally existed as continental mountain ranges. when dried floated much longer.islands. when dried it floated for eighty-five days. of granite. In the coral-producing oceans such sunken islands are now marked by rings of coral or atolls standing over them. instead of consisting of mere piles of volcanic matter. and when in the course of time we know something definite about the means of distribution. Hence I was led to dry the stems and branches of ninety-four plants with ripe fruit. they could not have been floated across wide spaces of the sea. It deserves notice that certain orders were far more injured than others: nine Leguminosae were tried. like other mountain summits. old fossiliferous and other rocks. Until I tried. seven species of the allied orders. For convenience sake I chiefly tried small seeds without the capsules or fruit. as it will some day be. But I do not believe that it will ever be proved that within the recent period most of our continents which now stand quite separate. Hydrophyllaceae and Polemoniaceae. and to place them on sea-water. Whenever it is fully admitted. and with the many existing oceanic islands. have been continuously.

to my surprise nearly all germinated. when floating on the sea. The fact of the larger fruits often floating longer than the small. Seeds may be occasionally transported in another manner. so that they were alternately wet and exposed to the air like really floating plants. and afterwards germinated. as we have seen. for he placed the seeds in a box in the actual sea. and would retain their power of germination. and some of the eighteen floated for a very much longer period. I could give many facts showing how frequently birds of many kinds are blown by gales to vast distances across the . even on those in the midst of the widest oceans. but some taken out of the crop of a pigeon. But I do not doubt that plants exposed to the waves would float for a less time than those protected from violent movement as in our experiments. out of the ninety-four dried plants. de Candolle has shown. and likewise seeds. after being dried. mostly different from mine. that the seeds of 14/100 kinds of plants of any country might be floated by sea-currents during twenty-eight days. The result was that 18/98 of his seeds of different kinds floated for forty-two days. which had floated on artificial sea-water for thirty days. On the other hand. if blown by an inland gale to a favourable spot. and when stranded. these stones being a valuable royal tax. Drift timber is thrown up on most islands. and were then capable of germination. sometimes escape being immediately devoured. so perfectly that not a particle could be washed away during the longest transport: out of one small portion of earth thus COMPLETELY enclosed by the roots of an oak about fifty years old. and as 18/94 distinct species with ripe fruit (but not all the same species as in the foregoing experiment) floated. the seeds of 14/100 plants belonging to one country might be floated across 924 miles of sea to another country. I find that when irregularly shaped stones are embedded in the roots of trees. Martens tried similar ones. for above twenty-eight days. three dicotyledonous plants germinated: I am certain of the accuracy of this observation. and would then germinate. solely from the roots of drifted trees. on this average. as Alph. and the natives of the coral islands in the Pacific procure stones for their tools.dried they floated for above ninety days. as far as anything can be inferred from these scanty facts. Therefore. and this. and many kinds of seeds in the crops of floating birds long retain their vitality: peas and vetches. are killed by even a few days' immersion in sea-water. Altogether. he did not previously dry the plants or branches with the fruit. after having been dried. small parcels of earth are very frequently enclosed in their interstices and behind them. Living birds can hardly fail to be highly effective agents in the transportation of seeds. In Johnston's Physical Atlas. He tried ninety-eight seeds. could be floated across a space of sea 900 miles in width. could hardly be transported by any other means. Again. eighteen floated for above twenty-eight days. and this would have favoured both the average length of their flotation and their resistance to the injurious action of the salt-water. would have caused some of them to have floated much longer. I can show that the carcasses of birds. is interesting. but in a much better manner. So that as 64/87 kinds of seeds germinated after an immersion of twenty-eight days. as plants with large seeds or fruit which. the average rate of the several Atlantic currents is thirty-three miles per diem (some currents running at the rate of sixty miles per diem). generally have restricted ranges. it would perhaps be safer to assume that the seeds of about 10/100 plants of a flora. we may conclude. from plants which live near the sea. Subsequently to my experiments. M. but he chose many large fruits. for instance. would germinate.

ocean. We may safely assume that under such circumstances their rate of flight would often be thirty-five miles an hour; and some authors have given a far higher estimate. I have never seen an instance of nutritious seeds passing through the intestines of a bird; but hard seeds of fruit pass uninjured through even the digestive organs of a turkey. In the course of two months, I picked up in my garden twelve kinds of seeds, out of the excrement of small birds, and these seemed perfect, and some of them, which were tried, germinated. But the following fact is more important: the crops of birds do not secrete gastric juice, and do not, as I know by trial, injure in the least the germination of seeds; now, after a bird has found and devoured a large supply of food, it is positively asserted that all the grains do not pass into the gizzard for twelve or even eighteen hours. A bird in this interval might easily be blown to the distance of five hundred miles, and hawks are known to look out for tired birds, and the contents of their torn crops might thus readily get scattered. Some hawks and owls bolt their prey whole, and after an interval of from twelve to twenty hours, disgorge pellets, which, as I know from experiments made in the Zoological Gardens, include seeds capable of germination. Some seeds of the oat, wheat, millet, canary, hemp, clover, and beet germinated after having been from twelve to twenty-one hours in the stomachs of different birds of prey; and two seeds of beet grew after having been thus retained for two days and fourteen hours. Fresh-water fish, I find, eat seeds of many land and water plants; fish are frequently devoured by birds, and thus the seeds might be transported from place to place. I forced many kinds of seeds into the stomachs of dead fish, and then gave their bodies to fishing-eagles, storks, and pelicans; these birds, after an interval of many hours, either rejected the seeds in pellets or passed them in their excrement; and several of these seeds retained the power of germination. Certain seeds, however, were always killed by this process. Locusts are sometimes blown to great distances from the land. I myself caught one 370 miles from the coast of Africa, and have heard of others caught at greater distances. The Rev. R.T. Lowe informed Sir C. Lyell that in November, 1844, swarms of locusts visited the island of Madeira. They were in countless numbers, as thick as the flakes of snow in the heaviest snowstorm, and extended upward as far as could be seen with a telescope. During two or three days they slowly careered round and round in an immense ellipse, at least five or six miles in diameter, and at night alighted on the taller trees, which were completely coated with them. They then disappeared over the sea, as suddenly as they had appeared, and have not since visited the island. Now, in parts of Natal it is believed by some farmers, though on insufficient evidence, that injurious seeds are introduced into their grass-land in the dung left by the great flights of locusts which often visit that country. In consequence of this belief Mr. Weale sent me in a letter a small packet of the dried pellets, out of which I extracted under the microscope several seeds, and raised from them seven grass plants, belonging to two species, of two genera. Hence a swarm of locusts, such as that which visited Madeira, might readily be the means of introducing several kinds of plants into an island lying far from the mainland. Although the beaks and feet of birds are generally clean, earth sometimes adheres to them: in one case I removed sixty-one grains, and in another case twenty-two grains of dry argillaceous earth from the foot of a partridge, and in the earth there was a pebble as large as the seed of a vetch. Here is a better case: the leg of a woodcock was sent to me by a friend, with a little cake of dry earth attached to the shank, weighing only nine grains; and this contained a seed of the toad-rush (Juncus bufonius) which germinated and flowered. Mr.

Swaysland, of Brighton, who during the last forty years has paid close attention to our migratory birds, informs me that he has often shot wagtails (Motacillae), wheatears, and whinchats (Saxicolae), on their first arrival on our shores, before they had alighted; and he has several times noticed little cakes of earth attached to their feet. Many facts could be given showing how generally soil is charged with seeds. For instance, Professor Newton sent me the leg of a red-legged partridge (Caccabis rufa) which had been wounded and could not fly, with a ball of hard earth adhering to it, and weighing six and a half ounces. The earth had been kept for three years, but when broken, watered and placed under a bell glass, no less than eighty-two plants sprung from it: these consisted of twelve monocotyledons, including the common oat, and at least one kind of grass, and of seventy dicotyledons, which consisted, judging from the young leaves, of at least three distinct species. With such facts before us, can we doubt that the many birds which are annually blown by gales across great spaces of ocean, and which annually migrate--for instance, the millions of quails across the Mediterranean--must occasionally transport a few seeds embedded in dirt adhering to their feet or beaks? But I shall have to recur to this subject. As icebergs are known to be sometimes loaded with earth and stones, and have even carried brushwood, bones, and the nest of a land-bird, it can hardly be doubted that they must occasionally, as suggested by Lyell, have transported seeds from one part to another of the arctic and antarctic regions; and during the Glacial period from one part of the now temperate regions to another. In the Azores, from the large number of plants common to Europe, in comparison with the species on the other islands of the Atlantic, which stand nearer to the mainland, and (as remarked by Mr. H.C. Watson) from their somewhat northern character, in comparison with the latitude, I suspected that these islands had been partly stocked by ice-borne seeds during the Glacial epoch. At my request Sir C. Lyell wrote to M. Hartung to inquire whether he had observed erratic boulders on these islands, and he answered that he had found large fragments of granite and other rocks, which do not occur in the archipelago. Hence we may safely infer that icebergs formerly landed their rocky burdens on the shores of these mid-ocean islands, and it is at least possible that they may have brought thither the seeds of northern plants. Considering that these several means of transport, and that other means, which without doubt remain to be discovered, have been in action year after year for tens of thousands of years, it would, I think, be a marvellous fact if many plants had not thus become widely transported. These means of transport are sometimes called accidental, but this is not strictly correct: the currents of the sea are not accidental, nor is the direction of prevalent gales of wind. It should be observed that scarcely any means of transport would carry seeds for very great distances; for seeds do not retain their vitality when exposed for a great length of time to the action of sea water; nor could they be long carried in the crops or intestines of birds. These means, however, would suffice for occasional transport across tracts of sea some hundred miles in breadth, or from island to island, or from a continent to a neighbouring island, but not from one distant continent to another. The floras of distant continents would not by such means become mingled; but would remain as distinct as they now are. The currents, from their course, would never bring seeds from North America to Britain, though they might and do bring seeds from the West Indies to our western shores, where, if not killed by their very long immersion in salt water, they could not endure our climate. Almost every year, one or two

land-birds are blown across the whole Atlantic Ocean, from North America to the western shores of Ireland and England; but seeds could be transported by these rare wanderers only by one means, namely, by dirt adhering to their feet or beaks, which is in itself a rare accident. Even in this case, how small would be the chance of a seed falling on favourable soil, and coming to maturity! But it would be a great error to argue that because a well-stocked island, like Great Britain, has not, as far as is known (and it would be very difficult to prove this), received within the last few centuries, through occasional means of transport, immigrants from Europe or any other continent, that a poorly-stocked island, though standing more remote from the mainland, would not receive colonists by similar means. Out of a hundred kinds of seeds or animals transported to an island, even if far less well-stocked than Britain, perhaps not more than one would be so well fitted to its new home, as to become naturalised. But this is no valid argument against what would be effected by occasional means of transport, during the long lapse of geological time, whilst the island was being upheaved, and before it had become fully stocked with inhabitants. On almost bare land, with few or no destructive insects or birds living there, nearly every seed which chanced to arrive, if fitted for the climate, would germinate and survive. DISPERSAL DURING THE GLACIAL PERIOD. The identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where Alpine species could not possibly exist, is one of the most striking cases known of the same species living at distant points, without the apparent possibility of their having migrated from one point to the other. It is indeed a remarkable fact to see so many plants of the same species living on the snowy regions of the Alps or Pyrenees, and in the extreme northern parts of Europe; but it is far more remarkable, that the plants on the White Mountains, in the United States of America, are all the same with those of Labrador, and nearly all the same, as we hear from Asa Gray, with those on the loftiest mountains of Europe. Even as long ago as 1747, such facts led Gmelin to conclude that the same species must have been independently created at many distinct points; and we might have remained in this same belief, had not Agassiz and others called vivid attention to the Glacial period, which, as we shall immediately see, affords a simple explanation of these facts. We have evidence of almost every conceivable kind, organic and inorganic, that, within a very recent geological period, central Europe and North America suffered under an Arctic climate. The ruins of a house burnt by fire do not tell their tale more plainly than do the mountains of Scotland and Wales, with their scored flanks, polished surfaces, and perched boulders, of the icy streams with which their valleys were lately filled. So greatly has the climate of Europe changed, that in Northern Italy, gigantic moraines, left by old glaciers, are now clothed by the vine and maize. Throughout a large part of the United States, erratic boulders and scored rocks plainly reveal a former cold period. The former influence of the glacial climate on the distribution of the inhabitants of Europe, as explained by Edward Forbes, is substantially as follows. But we shall follow the changes more readily, by supposing a new glacial period slowly to come on, and then pass away, as formerly occurred. As the cold came on, and as each more southern zone became fitted for the inhabitants of the north, these would take the places of the former inhabitants of the temperate regions. The latter, at the same time would travel further and further southward, unless they were stopped by barriers, in which case they would perish. The mountains

would become covered with snow and ice, and their former Alpine inhabitants would descend to the plains. By the time that the cold had reached its maximum, we should have an arctic fauna and flora, covering the central parts of Europe, as far south as the Alps and Pyrenees, and even stretching into Spain. The now temperate regions of the United States would likewise be covered by arctic plants and animals and these would be nearly the same with those of Europe; for the present circumpolar inhabitants, which we suppose to have everywhere travelled southward, are remarkably uniform round the world. As the warmth returned, the arctic forms would retreat northward, closely followed up in their retreat by the productions of the more temperate regions. And as the snow melted from the bases of the mountains, the arctic forms would seize on the cleared and thawed ground, always ascending, as the warmth increased and the snow still further disappeared, higher and higher, whilst their brethren were pursuing their northern journey. Hence, when the warmth had fully returned, the same species, which had lately lived together on the European and North American lowlands, would again be found in the arctic regions of the Old and New Worlds, and on many isolated mountain-summits far distant from each other. Thus we can understand the identity of many plants at points so immensely remote as the mountains of the United States and those of Europe. We can thus also understand the fact that the Alpine plants of each mountain-range are more especially related to the arctic forms living due north or nearly due north of them: for the first migration when the cold came on, and the re-migration on the returning warmth, would generally have been due south and north. The Alpine plants, for example, of Scotland, as remarked by Mr. H.C. Watson, and those of the Pyrenees, as remarked by Ramond, are more especially allied to the plants of northern Scandinavia; those of the United States to Labrador; those of the mountains of Siberia to the arctic regions of that country. These views, grounded as they are on the perfectly well-ascertained occurrence of a former Glacial period, seem to me to explain in so satisfactory a manner the present distribution of the Alpine and Arctic productions of Europe and America, that when in other regions we find the same species on distant mountain-summits, we may almost conclude, without other evidence, that a colder climate formerly permitted their migration across the intervening lowlands, now become too warm for their existence. As the arctic forms moved first southward and afterwards backward to the north, in unison with the changing climate, they will not have been exposed during their long migrations to any great diversity of temperature; and as they all migrated in a body together, their mutual relations will not have been much disturbed. Hence, in accordance with the principles inculcated in this volume, these forms will not have been liable to much modification. But with the Alpine productions, left isolated from the moment of the returning warmth, first at the bases and ultimately on the summits of the mountains, the case will have been somewhat different; for it is not likely that all the same arctic species will have been left on mountain ranges far distant from each other, and have survived there ever since; they will also, in all probability, have become mingled with ancient Alpine species, which must have existed on the mountains before the commencement of the Glacial epoch, and which during the coldest period will have been temporarily driven down to the plains; they will, also, have been subsequently exposed to somewhat different climatical influences. Their mutual relations will thus have been in some degree disturbed; consequently

they will have been liable to modification; and they have been modified; for if we compare the present Alpine plants and animals of the several great European mountain ranges, one with another, though many of the species remain identically the same, some exist as varieties, some as doubtful forms or sub-species and some as distinct yet closely allied species representing each other on the several ranges. In the foregoing illustration, I have assumed that at the commencement of our imaginary Glacial period, the arctic productions were as uniform round the polar regions as they are at the present day. But it is also necessary to assume that many sub-arctic and some few temperate forms were the same round the world, for some of the species which now exist on the lower mountain slopes and on the plains of North America and Europe are the same; and it may be asked how I account for this degree of uniformity of the sub-arctic and temperate forms round the world, at the commencement of the real Glacial period. At the present day, the sub-arctic and northern temperate productions of the Old and New Worlds are separated from each other by the whole Atlantic Ocean and by the northern part of the Pacific. During the Glacial period, when the inhabitants of the Old and New Worlds lived further southwards than they do at present, they must have been still more completely separated from each other by wider spaces of ocean; so that it may well be asked how the same species could then or previously have entered the two continents. The explanation, I believe, lies in the nature of the climate before the commencement of the Glacial period. At this, the newer Pliocene period, the majority of the inhabitants of the world were specifically the same as now, and we have good reason to believe that the climate was warmer than at the present day. Hence, we may suppose that the organisms which now live under latitude 60 degrees, lived during the Pliocene period further north, under the Polar Circle, in latitude 66-67 degrees; and that the present arctic productions then lived on the broken land still nearer to the pole. Now, if we look at a terrestrial globe, we see under the Polar Circle that there is almost continuous land from western Europe through Siberia, to eastern America. And this continuity of the circumpolar land, with the consequent freedom under a more favourable climate for intermigration, will account for the supposed uniformity of the sub-arctic and temperate productions of the Old and New Worlds, at a period anterior to the Glacial epoch. Believing, from reasons before alluded to, that our continents have long remained in nearly the same relative position, though subjected to great oscillations of level, I am strongly inclined to extend the above view, and to infer that during some earlier and still warmer period, such as the older Pliocene period, a large number of the same plants and animals inhabited the almost continuous circumpolar land; and that these plants and animals, both in the Old and New Worlds, began slowly to migrate southwards as the climate became less warm, long before the commencement of the Glacial period. We now see, as I believe, their descendants, mostly in a modified condition, in the central parts of Europe and the United States. On this view we can understand the relationship with very little identity, between the productions of North America and Europe--a relationship which is highly remarkable, considering the distance of the two areas, and their separation by the whole Atlantic Ocean. We can further understand the singular fact remarked on by several observers that the productions of Europe and America during the later tertiary stages were more closely related to each other than they are at the present time; for during these warmer periods the northern parts of the Old and New Worlds will have been almost continuously united by land, serving as a bridge, since rendered impassable by cold, for the intermigration of their inhabitants.

During the slowly decreasing warmth of the Pliocene period, as soon as the species in common, which inhabited the New and Old Worlds, migrated south of the Polar Circle, they will have been completely cut off from each other. This separation, as far as the more temperate productions are concerned, must have taken place long ages ago. As the plants and animals migrated southward, they will have become mingled in the one great region with the native American productions, and would have had to compete with them; and in the other great region, with those of the Old World. Consequently we have here everything favourable for much modification--for far more modification than with the Alpine productions, left isolated, within a much more recent period, on the several mountain ranges and on the arctic lands of Europe and North America. Hence, it has come, that when we compare the now living productions of the temperate regions of the New and Old Worlds, we find very few identical species (though Asa Gray has lately shown that more plants are identical than was formerly supposed), but we find in every great class many forms, which some naturalists rank as geographical races, and others as distinct species; and a host of closely allied or representative forms which are ranked by all naturalists as specifically distinct. As on the land, so in the waters of the sea, a slow southern migration of a marine fauna, which, during the Pliocene or even a somewhat earlier period, was nearly uniform along the continuous shores of the Polar Circle, will account, on the theory of modification, for many closely allied forms now living in marine areas completely sundered. Thus, I think, we can understand the presence of some closely allied, still existing and extinct tertiary forms, on the eastern and western shores of temperate North America; and the still more striking fact of many closely allied crustaceans (as described in Dana's admirable work), some fish and other marine animals, inhabiting the Mediterranean and the seas of Japan--these two areas being now completely separated by the breadth of a whole continent and by wide spaces of ocean. These cases of close relationship in species either now or formerly inhabiting the seas on the eastern and western shores of North America, the Mediterranean and Japan, and the temperate lands of North America and Europe, are inexplicable on the theory of creation. We cannot maintain that such species have been created alike, in correspondence with the nearly similar physical conditions of the areas; for if we compare, for instance, certain parts of South America with parts of South Africa or Australia, we see countries closely similar in all their physical conditions, with their inhabitants utterly dissimilar. ALTERNATE GLACIAL PERIODS IN THE NORTH AND SOUTH. But we must return to our more immediate subject. I am convinced that Forbes's view may be largely extended. In Europe we meet with the plainest evidence of the Glacial period, from the western shores of Britain to the Ural range, and southward to the Pyrenees. We may infer from the frozen mammals and nature of the mountain vegetation, that Siberia was similarly affected. In the Lebanon, according to Dr. Hooker, perpetual snow formerly covered the central axis, and fed glaciers which rolled 4,000 feet down the valleys. The same observer has recently found great moraines at a low level on the Atlas range in North Africa. Along the Himalaya, at points 900 miles apart, glaciers have left the marks of their former low descent; and in Sikkim, Dr. Hooker saw maize growing on ancient and gigantic moraines. Southward of the Asiatic continent, on the opposite side of the equator, we know, from the excellent researches

of Dr. J. Haast and Dr. Hector, that in New Zealand immense glaciers formerly descended to a low level; and the same plants, found by Dr. Hooker on widely separated mountains in this island tell the same story of a former cold period. From facts communicated to me by the Rev. W.B. Clarke, it appears also that there are traces of former glacial action on the mountains of the south-eastern corner of Australia. Looking to America: in the northern half, ice-borne fragments of rock have been observed on the eastern side of the continent, as far south as latitude 36 and 37 degrees, and on the shores of the Pacific, where the climate is now so different, as far south as latitude 46 degrees. Erratic boulders have, also, been noticed on the Rocky Mountains. In the Cordillera of South America, nearly under the equator, glaciers once extended far below their present level. In central Chile I examined a vast mound of detritus with great boulders, crossing the Portillo valley, which, there can hardly be a doubt, once formed a huge moraine; and Mr. D. Forbes informs me that he found in various parts of the Cordillera, from latitude 13 to 30 degrees south, at about the height of 12,000 feet, deeply-furrowed rocks, resembling those with which he was familiar in Norway, and likewise great masses of detritus, including grooved pebbles. Along this whole space of the Cordillera true glaciers do not now exist even at much more considerable heights. Further south, on both sides of the continent, from latitude 41 degrees to the southernmost extremity, we have the clearest evidence of former glacial action, in numerous immense boulders transported far from their parent source. From these several facts, namely, from the glacial action having extended all round the northern and southern hemispheres--from the period having been in a geological sense recent in both hemispheres--from its having lasted in both during a great length of time, as may be inferred from the amount of work effected--and lastly, from glaciers having recently descended to a low level along the whole line of the Cordillera, it at one time appeared to me that we could not avoid the conclusion that the temperature of the whole world had been simultaneously lowered during the Glacial period. But now, Mr. Croll, in a series of admirable memoirs, has attempted to show that a glacial condition of climate is the result of various physical causes, brought into operation by an increase in the eccentricity of the earth's orbit. All these causes tend towards the same end; but the most powerful appears to be the indirect influence of the eccentricity of the orbit upon oceanic currents. According to Mr. Croll, cold periods regularly recur every ten or fifteen thousand years; and these at long intervals are extremely severe, owing to certain contingencies, of which the most important, as Sir C. Lyell has shown, is the relative position of the land and water. Mr. Croll believes that the last great glacial period occurred about 240,000 years ago, and endured, with slight alterations of climate, for about 160,000 years. With respect to more ancient glacial periods, several geologists are convinced, from direct evidence, that such occurred during the miocene and eocene formations, not to mention still more ancient formations. But the most important result for us, arrived at by Mr. Croll, is that whenever the northern hemisphere passes through a cold period the temperature of the southern hemisphere is actually raised, with the winters rendered much milder, chiefly through changes in the direction of the ocean currents. So conversely it will be with the northern hemisphere, while the southern passes through a glacial period. This conclusion throws so much light on geographical distribution that I am strongly inclined to trust in it; but I will first give the facts which demand an explanation.

Some of these Australian forms. Hooker. A list of the genera of plants collected on the loftier peaks of Java. Still more striking is the fact that peculiar Australian forms are represented by certain plants growing on the summits of the mountains of Borneo. Dr. almost under the equator. Hooker has shown that besides many closely allied species. Dr. H. of European genera. and are thinly scattered on the one hand over India. Hooker. Watson has remarked. On the southern mountains of Australia. Muller has discovered several European species.T. but not in the intermediate torrid regions. on the heights of Ceylon. are closely related to those on the mountains of Abyssinia. On the Silla of Caraccas the illustrious Humboldt long ago found species belonging to genera characteristic of the Cordillera.C. many species inhabiting the same widely sundered areas. across the whole continent of Africa and to the mountains of the Cape Verde archipelago. belong to genera not now found in the intermediate tropical lowlands. however. and at the same time representing plants of Europe not found in the intervening hot lowlands. as I hear from Dr. Lowe on the mountains of the Cape Verde Islands. Hooker. and on the volcanic cones of Java. and on the neighbouring Cameroon Mountains. not introduced by man. and some few representatives of the flora of the Cape of Good Hope. between forty and fifty of the flowering plants of Tierra del Fuego. found in Australia. It should. occur on the lowlands. and on the isolated mountain ranges of the peninsula of India. This extension of the same temperate forms. are common to North America and Europe. forming no inconsiderable part of its scanty flora.In South America. in the Gulf of Guinea. as Mr. R. In the admirable "Introduction to the Flora of New Zealand. believed not to have been introduced by man. Hooker. On the Organ Mountains of Brazil some few temperate European. are either the same species or varieties of the same species. It now also appears. some Antarctic and some Andean genera were found by Gardner which do not exist in the low intervening hot countries. other species. enormously remote as these areas in opposite hemispheres are from each other. is one of the most astonishing facts ever recorded in the distribution of plants. be observed that these plants are not strictly arctic forms. occur on the mountains of Abyssinia. we see that certain plants growing on the more lofty mountains of the tropics in all parts of the world. that some of these same temperate plants have been discovered by the Rev. analogous and striking facts are given in regard to the plants of that large island. extend along the heights of the peninsula of Malacca. and on the other hand as far north as Japan. "in receding from polar toward equatorial latitudes. and on the temperate plains of the north and south." Besides these identical and closely allied forms. but some few analogous facts . and on the mountains several representative European forms are found which have not been discovered in the intertropical parts of Africa. Dr. In Africa. At the Cape of Good Hope a very few European species. and a long list can be given. raises a picture of a collection made on a hillock in Europe. many plants occur either identically the same or representing each other. Hence. several forms characteristic of Europe. as I am informed by Dr. On the lofty mountains of equatorial America a host of peculiar species belonging to European genera occur. F. the Alpine or mountain flora really become less and less Arctic. These brief remarks apply to plants alone. as I hear from Dr. and likewise to those of temperate Europe. Hooker has also lately shown that several of the plants living on the upper parts of the lofty island of Fernando Po. for. On the Himalaya." by Dr.

" Now let us see whether Mr." Sir J. Croll's conclusion that when the northern hemisphere suffered from the extreme cold of the great Glacial period. as measured by years. must have been very long.. They might have survived. Prof. In marine productions. As the cold became more and more intense. and when we remember over what vast spaces some naturalised plants and animals have spread within a few centuries. of the northern temperate forms would almost certainly have ascended any adjoining high land. The Glacial period. Dana. dominant and widest-spreading temperate forms invaded the equatorial lowlands. and on the mountains of the tropics. like that described by Hooker as growing luxuriantly at the height of from four to five thousand feet on the lower slopes of the Himalaya. no doubt during the most severe part of a Glacial period. On the mountains of Panama. etc. similar cases likewise occur. Mr. and along the Peninsula of India. Hooker informs me that twenty-five species of Algae are common to New Zealand and to Europe. During this. at the height of only two thousand feet. would have been driven to their former homes or have been destroyed. the presence of temperate forms on the highlands across the whole of equatorial Africa. its antipode. but have not been found in the intermediate tropical seas. throws any clear light on the present apparently inexplicable distribution of various organisms in the temperate parts of both hemispheres. . for the change of temperature must have been very slow. the coldest period. namely. Seemann found the vegetation like that of Mexico. Some. as an example. than to any other part of the world. there can hardly be a doubt that some of the more vigorous. of northern forms of fish. where. in the Gulf of Guinea.could be given in regard to terrestrial animals. So again in the mountainous island of Fernando Po. the southern hemisphere was actually warmer. Richardson. Dr. the northern temperate forms living on the lowlands under the equator. or perhaps even rather cooler. this period will have been ample for any amount of migration. At this period the equatorial climate at the level of the sea was probably about the same with that now experienced at the height of from five to six thousand feet under the same latitude. to Ceylon and the Malay Archipelago. and plants undoubtedly possess a certain capacity for acclimatisation. the lowlands of these great continents were everywhere tenanted under the equator by a considerable number of temperate forms. if sufficiently lofty. as shown by their transmitting to their offspring different constitutional powers of resisting heat and cold. On the decline of the Glacial period. that "it is certainly a wonderful fact that New Zealand should have a closer resemblance in its crustacea to Great Britain. however. Mann found temperate European forms beginning to appear at the height of about five thousand feet. also. The inhabitants of these hot lowlands would at the same time have migrated to the tropical and subtropical regions of the south. speaks of the reappearance on the shores of New Zealand. "with forms of the torrid zone harmoniously blended with those of the temperate. I may quote a statement by the highest authority. and in a less well-marked manner across the wide expanse of tropical South America. Dr. the lowlands under the equator must have been clothed with a mingled tropical and temperate vegetation. even if the climate was not perfectly fitted for them. but with perhaps a still greater preponderance of temperate forms. as both hemispheres gradually recovered their former temperature. and from the facts just given. Tasmania. it appears almost certain that at some former period. for the southern hemisphere was at this period warmer. they would have long survived like the Arctic forms on the mountains of Europe. we know that Arctic forms invaded the temperate regions. From the foregoing facts. being replaced by the equatorial forms returning from the south.

during the alternations of the Glacial periods. Mountains are islands on the land. and this is the first stage towards their extinction. we see that very many European productions cover the ground in La Plata. and having consequently been advanced through natural selection and competition to a higher stage of perfection. I suspect that this preponderant migration from the north to the south is due to the greater extent of land in the north. The northern forms which had before been left on the mountains would now descend and mingle with the southern forms. would have to compete with many new forms and would be exposed to somewhat different physical conditions. Hooker. The same principles apply to the distribution of terrestrial animals and of marine productions. for here. with the northern hemisphere rendered warmer. de Candolle in regard to Australia. that many more identical or slightly modified species have migrated from the north to the south. would return to their former homes.In the regular course of events the southern hemisphere would in its turn be subjected to a severe Glacial period. however. just in the same way as the inhabitants of real islands have everywhere yielded and are still yielding to continental forms naturalised through man's agency. and this is the case. wool. at the present day. or dominating power. and have beaten the natives. and belonging to genera not now found in the intermediate torrid zones. a few southern forms on the mountains of Borneo and Abyssinia. We see. And thus. or in opposite hemispheres. But the species left during a long time on these mountains. and then the southern temperate forms would invade the equatorial lowlands. in the northern and southern temperate zones. than the southern forms. and . and to a lesser degree in Australia. when the warmth returned. the northern forms were the more powerful and were able to hold their places on the mountains. but not so the southern in regard to the northern forms. and by Alph. We must. for the many quite distinct species inhabiting the same widely separated areas. and afterwards migrate southward with the southern forms. for these will account. and would generally now exist as varieties or as representative species. Australian forms are rapidly sowing themselves and becoming naturalised. bear in mind the occurrence in both hemispheres of former Glacial periods. and other objects likely to carry seeds have been largely imported into Europe during the last two or three centuries from La Plata and during the last forty or fifty years from Australia. by those which have become naturalised. or even outnumbered. It is a remarkable fact. The Neilgherrie Mountains in India. hence. as I hear from Dr. New Zealand. whereas extremely few southern forms have become naturalised in any part of the northern hemisphere. however. and carrying southward with them some of the northern temperate forms which had descended from their mountain fastnesses. also. and to the northern forms having existed in their own homes in greater numbers. In many islands the native productions are nearly equalled. when the two sets became commingled in the equatorial regions. strongly insisted on by Hooker in regard to America. offer a partial exception. we should have some few species identically the same in the northern and southern temperate zones and on the mountains of the intermediate tropical regions. Thus. though hides. but these have almost everywhere yielded to the more dominant forms generated in the larger areas and more efficient workshops of the north. Before the last great Glacial period. In the same manner. no doubt the intertropical mountains were stocked with endemic Alpine forms. in accordance with the same principles. they would be eminently liable to modification. These latter. and their inhabitants have yielded to those produced within the larger areas of the north. leaving some few species on the mountains. than in a reversed direction.

explains a multitude of facts in the distribution of the same and of the allied forms of life in all parts of the globe. until we can say why one species and not another becomes naturalised by man's agency in a foreign land. Australia. may have been concerned in their dispersal. rising higher on the shores where the tide rises highest. It may be suspected that before this flora was exterminated during the last Glacial epoch. as halting-places. Hooker. in nearly the same manner as. The living waters have flowed during one period from the north and during another from the south. the ocean-currents were widely different to what they now are. and by the aid. on the effects of great alternations of climate throughout the world on geographical distribution. why certain species have been modified and have given rise to new forms. a few forms had been already widely dispersed to various points of the southern hemisphere by occasional means of transport. to a former and warmer period. during the height of the Glacial period. some of the inhabitants of the temperate seas might have reached the equator. Various special difficulties also remain to be solved. As the tide leaves its drift in horizontal lines. now covered with ice. We cannot say why certain species and not others have migrated. though distinct. as shown by Dr. Croll's conclusion that successive Glacial periods in the one hemisphere coincide with warmer periods in the opposite hemisphere. Sir C. while others might remain and survive in the colder depths until the southern hemisphere was in its turn subjected to a glacial climate and permitted their further progress. together with the admission of the slow modification of species. which. of now sunken islands. when the Antarctic lands. as another species within their own homes. New Zealand. Lyell in a striking passage has speculated. by keeping to the cooler currents. while others have remained unaltered. and I am inclined to look in the southern. and in both cases have reached the equator. is a more remarkable case. the occurrence. are removed on the views above given. but the stream of life has flowed with greater force from the north than in the opposite direction. in language almost identical with mine. as in the northern hemisphere. I am far from supposing that all the difficulties in regard to the distribution and affinities of the identical and allied species. of the same plants at points so enormously remote as Kerguelen Land. before the commencement of the last Glacial period. as suggested by Lyell. that we cannot suppose that there has been time since the commencement of the last Glacial period for their migration and subsequent modification to the necessary degree. Some of these species are so distinct. and New Zealand may have become slightly tinted by the same peculiar forms of life. The facts seem to indicate that distinct species belonging to the same genera have migrated in radiating lines from a common centre. The exact lines of migration cannot be indicated. for instance. according to Forbes. and Fuegia. but icebergs. of species. and has consequently more freely inundated the south. and is twice or thrice as common. why one species ranges twice or thrice as far. The existence at these and other distant points of the southern hemisphere. isolated spaces inhabited by Arctic productions exist to the present day in the deeper parts of the northern temperate seas. We cannot hope to explain such facts. And we have now seen that Mr. supported a highly peculiar and isolated flora. so have the living waters left their . of these a few would perhaps at once be able to migrate southwards. and sometimes on the intermediate mountain ranges. which now live so widely separated in the north and south. belong to genera exclusively confined to the south. When. Thus the southern shores of America.on the intertropical mountains.

compared with those of Britain. be mainly attributed to changes in the level of the land within the recent period. some of the most difficult to explain are presented by fish. This is a wonderful case. New Zealand. and as the sea is apparently a still more formidable barrier. of the former inhabitants of the surrounding lowlands. and liability to wide dispersal would follow from this capacity as an almost necessary consequence. or from stream to stream. causing rivers to flow . etc. The various beings thus left stranded may be compared with savage races of man. But the case is exactly the reverse. When first collecting in the fresh waters of Brazil. As lakes and river-systems are separated from each other by barriers of land. which serves as a record. It is probable that they are occasionally transported by what may be called accidental means. driven up and surviving in the mountain fastnesses of almost every land. But Dr. It was formerly believed that the same fresh-water species never existed on two continents distant from each other. within their own countries. and probably indicates dispersal from an Antarctic centre during a former warm period. I think. We can here consider only a few cases. for short and frequent migrations from pond to pond. however. Not only have many fresh-water species. for in two adjoining river systems some of the species may be the same and some wholly different.. in a line gently rising from the Arctic lowlands to a great latitude under the equator. it might have been thought that fresh-water productions would not have ranged widely within the same country. I well remember feeling much surprise at the similarity of the fresh-water insects. and it is known that the ova retain their vitality for a considerable time after removal from the water. belonging to different classes. shells. Their dispersal may. in a manner highly useful to them. an enormous range. though separated by a distance of about 230 miles. that they would never have extended to distant countries. the Falkland Islands and the mainland of South America. full of interest to us. of these. however. Distribution of fresh-water productions--On the inhabitants of oceanic islands--Absence of Batrachians and of terrestrial Mammals--On the relation of the inhabitants of islands to those of the nearest mainland--On colonisation from the nearest source with subsequent modification--Summary of the last and present chapters. On the same continent fresh-water fish often range widely. is rendered in some degree less surprising by the species of this genus having the power of crossing by some unknown means considerable spaces of open ocean: thus there is one species common to New Zealand and to the Auckland Islands. FRESH-WATER PRODUCTIONS.living drift on our mountain summits. but allied species prevail in a remarkable manner throughout the world. Gunther has lately shown that the Galaxias attenuatus inhabits Tasmania. and as if capriciously. in most cases be explained by their having become fitted. Thus fishes still alive are not very rarely dropped at distant points by whirlwinds. GEOGRAPHICAL DISTRIBUTION--continued. CHAPTER XIII. and at the dissimilarity of the surrounding terrestrial beings. But the wide ranging power of fresh-water productions can. This case.

prevail throughout the world. and allied species which. once flew on board the "Beagle. so that any . as their ova are not likely to be transported by birds. leads to the same conclusion. and the ova. survived on the duck's feet. though aquatic in their nature. according to Alph. With respect to plants. according to Valenciennes. on our theory. Gunther has recently been led by several considerations to infer that with fishes the same forms have a long endurance. and a water-beetle of the same family. could be given of this having occurred during floods. Moreover.into each other. where many ova of fresh-water shells were hatching. as well as the adults. and even marsh-species. for the latter seem immediately to acquire. These just-hatched molluscs. The wide differences of the fish on the opposite sides of most mountain-ranges. in removing a little duck-weed from one aquarium to another. and consequently time and means for much migration. de Candolle. I think favourable means of dispersal explain this fact. also. But another agency is perhaps more effectual: I suspended the feet of a duck in an aquarium. they would not be likely to alight on the surface of the sea. which I have observed--and many others no doubt will be discovered--throw some light on this subject. so that a marine species belonging to a fresh-water group might travel far along the shores of the sea. Their distribution at first perplexed me much. are immediately killed by sea-water. which have very few aquatic members. it is probable. which frequent the muddy edges of ponds. that I have unintentionally stocked the one with fresh-water shells from the other. and in such cases there will have been ample time for great geographical changes. it has long been known what enormous ranges many fresh-water. Some species of fresh-water shells have very wide ranges. a Colymbetes. in damp air. When ducks suddenly emerge from a pond covered with duck-weed. in those large groups of terrestrial plants. Salt-water fish can with care be slowly accustomed to live in fresh water. which are continuous and consequently must. I could not even understand how some naturalised species have spread rapidly throughout the same country." when forty-five miles distant from the nearest land: how much farther it might have been blown by a favouring gale no one can tell. as if in consequence. and it has happened to me. would be sure to alight on a pool or rivulet. though at a somewhat more advanced age they would voluntarily drop off. both over continents and to the most remote oceanic islands. from an early period. Wading birds. This is strikingly illustrated. from twelve to twenty hours. Instances. Some fresh-water fish belong to very ancient forms. Birds of this order wander more than those of any other. I have before mentioned that earth occasionally adheres in some quantity to the feet and beaks of birds. or to any other distant point. and. I have twice seen these little plants adhering to their backs. and clung to them so firmly that when taken out of the water they could not be jarred off. Sir Charles Lyell informs me that a Dyticus has been caught with an Ancylus (a fresh-water shell like a limpet) firmly adhering to it. But two facts. and in this length of time a duck or heron might fly at least six or seven hundred miles. have completely prevented the inosculation of the river systems on the two sides. become adapted without much difficulty to the fresh waters of a distant land. have. if suddenly flushed. without any change of level. are descended from a common parent. would be the most likely to have muddy feet. and must have proceeded from a single source. a wide range. and could. there is hardly a single group of which all the members are confined to fresh water. and if blown across the sea to an oceanic island. and are occasionally found on the most remote and barren islands of the open ocean. Dr. and I found that numbers of the extremely minute and just-hatched shells crawled on the feet.

but will here give only the most striking case: I took in February three tablespoonfuls of mud from three different points. on the edge of a little pond. which have great powers of flight. or in some degree modified. beneath water. analogy makes me believe that it would have rejected the seeds in the pellet in a fit state for germination. even small fish swallow seeds of moderate size. yet as the number even in a well-stocked pond is small in comparison with the number of species inhabiting an equal area of land. de Candolle's remarks on the distribution of this plant. this mud when dry weighed only 6 and 3/4 ounces. it should be remembered that when a pond or stream is first formed. they then take flight and go to other waters. I have stated that fresh-water fish eat some kinds of seeds. and we have reason to believe that such beings become modified more slowly than the high. pulling up and counting each plant as it grew. though they reject many other kinds after having swallowed them. for instance on a rising islet. We should not forget the probability of many fresh-water forms having formerly ranged continuously over immense areas. when rejected many hours afterwards in pellets or in the excrement. Now this bird must often have flown with its stomach thus well stocked to distant ponds. and were altogether 537 in number. the plants were of many kinds. have gone on daily devouring fish. and. Although there will always be a struggle for life between the inhabitants of the same pond. We should also remember that many fresh-water productions are low in the scale of nature. situated at very distant points. and then having become extinct at intermediate points. apparently depends in main part on the wide dispersal of their seeds and eggs by animals. or are blown across the sea. and remembered Alph. Herons and other birds. the competition between them will probably be less severe than between terrestrial species. But the wide distribution of fresh-water plants. more especially by fresh-water birds. and naturally travel from one piece of water to another. and we have seen that seeds retain their power of germination.dirt on their feet would not be washed off. and when gaining the land. Other and unknown agencies probably have also played a part. The same agency may have come into play with the eggs of some of the smaller fresh-water animals. than in the case of terrestrial colonists. as of the yellow water-lily and Potamogeton. and a single seed or egg will have a good chance of succeeding. I kept it covered up in my study for six months. In considering these several means of distribution. I think it would be an inexplicable circumstance if water-birds did not transport the seeds of fresh-water plants to unstocked ponds and streams. ON THE INHABITANTS OF OCEANIC ISLANDS. then getting a hearty meal of fish. and of the lower animals. Hooker. they would be sure to fly to their natural fresh-water haunts. and yet the viscid mud was all contained in a breakfast cup! Considering these facts. When I saw the great size of the seeds of that fine water-lily. the Nelumbium. but Audubon states that he found the seeds of the great southern water-lily (probably according to Dr. consequently an intruder from the waters of a foreign country would have a better chance of seizing on a new place. I thought that the means of its dispersal must remain inexplicable. the Nelumbium luteum) in a heron's stomach. it will be unoccupied. and this will give time for the migration of aquatic species. I do not believe that botanists are aware how charged the mud of ponds is with seeds: I have tried several little experiments. century after century. whether retaining the same identical form. however few in kind. .

as already explained. New Zealand. and the comparison in some other respects is not quite fair. which I have selected as presenting the greatest amount of difficulty with respect to distribution. would be eminently liable to modification. If we compare. so that even an occasional cross would produce more effect than might have been . and partly on the frequent arrival of unmodified immigrants from the mother-country. are peculiar. extending over 780 miles of latitude. with the number found on any continent. for man has unintentionally stocked them far more fully and perfectly than did nature. de Candolle admits this for plants. for instance. and Wollaston for insects. I have already given my reasons for disbelieving in continental extensions within the period of existing species on so enormous a scale that all the many islands of the several oceans were thus stocked with their present terrestrial inhabitants. together with the outlying islands of Auckland. In the following remarks I shall not confine myself to the mere question of dispersal. He who admits the doctrine of the creation of each separate species. after long intervals of time in the new and isolated district. for. so that their mutual relations have not been much disturbed. species occasionally arriving. as they have in New Zealand and on every other oceanic island which can be named. but that allied species. It should be borne in mind that the offspring of such crosses would certainly gain in vigour. the number of endemic land-shells in Madeira. In St. and would often produce groups of modified descendants. but shall consider some other cases bearing on the truth of the two theories of independent creation and of descent with modification. with which the insular forms have intercrossed. with its lofty mountains and diversified stations. the birthplace of their early progenitors. although now inhabiting the most distant points. have proceeded from a single area. or of another section of the same class. those of another class. the proportion of endemic kinds (i. contain altogether only 960 kinds of flowering plants. Although in oceanic islands the species are few in number. or of endemic birds in the Galapagos Archipelago. for instance. and this difference seems to depend partly on the species which are not modified having immigrated in a body. has given rise to so great a difference in number. if we compare this moderate number with the species which swarm over equal areas in Southwestern Australia or at the Cape of Good Hope. we shall see that this is true. and then compare the area of the island with that of the continent. Campbell and Chatham. We have evidence that the barren island of Ascension aboriginally possessed less than half-a-dozen flowering plants.e. But it by no means follows that. and having to compete with new associates. but a few ferns and a few introduced plants are included in these numbers. independently of different physical conditions. Helena there is reason to believe that the naturalised plants and animals have nearly or quite exterminated many native productions. on the view that not only all the individuals of the same species have migrated from some one area. because in an island nearly all the species of one class are peculiar. yet many species have now become naturalised on it. This view removes many difficulties. we must admit that some cause. those found nowhere else in the world) is often extremely large. The species of all kinds which inhabit oceanic islands are few in number compared with those on equal continental areas: Alph. and the little island of Anglesea 764. but it does not accord with all the facts in regard to the productions of islands. will have to admit that a sufficient number of the best adapted plants and animals were not created for oceanic islands. Even the uniform county of Cambridge has 847 plants.We now come to the last of the three classes of facts. This fact might have been theoretically expected.

and three or four other species are confined to this island and to the Canaries. J. still retaining its hooks. the place of mammals. whereas of the eleven marine birds only two are peculiar. All such differences in number. Almost every year. or recently took. often arriving from the mother-country. and we know from Mr. some of the endemic plants have beautifully hooked seeds. Facility of immigration seems to have been fully as important as the nature of the conditions. it is in some degree doubtful whether it should be so ranked. Jones's admirable account of Bermuda. when settled in their new homes. in certain islands not tenanted by a single mammal. are generally accounted for by supposed differences in the physical conditions of the islands. yet few relations are more manifest than that hooks serve for the transportal of seeds in the wool or fur of quadrupeds. Again. islands often possess trees or bushes belonging to orders which elsewhere include only herbaceous species. or to the feet of wading birds. The different orders of insects inhabiting Madeira present nearly parallel cases. Hence. which would form a useless appendage. this island is inhabited by ninety-nine kinds. the Rev. Madeira again is inhabited by a wonderful number of peculiar land-shells. as Alph.V. as well as New Caledonia. from its geological character and the direction of its mountain ranges. which for long ages have there struggled together. Bermuda. though we do not know how sea-shells are dispersed. whereas not one species of sea-shell is peculiar to its shores: now. and the plant then becoming modified would form an endemic species. on the other hand. Harcourt. though very closely related to a European form. Turning to plants. perhaps attached to seaweed or floating timber. Dr. I will give a few illustrations of the foregoing remarks: in the Galapagos Islands there are twenty-six land birds. as I am informed by Mr. E. many European and African birds are blown to Madeira. Many remarkable little facts could be given with respect to the inhabitants of oceanic islands. For instance. Any tendency to modification will also have been checked by intercrossing with the unmodified immigrants. W. Oceanic islands are sometimes deficient in animals of certain whole classes. and which has a very peculiar soil. which lies at about the same distance from North America as the Galapagos Islands do from South America. might be transported across three or four hundred miles of open sea far more easily than land-shells. like the shrivelled wings under the soldered wing-covers of many insular beetles.B. that very many North American birds occasionally or even frequently visit this island. of these twenty-one (or perhaps twenty-three) are peculiar. does not possess a single endemic land bird. it is of large size.anticipated. should be considered as appurtenances of Australia. Hooker has shown that in the Galapagos Islands the proportional numbers of the different orders are very different from what they are elsewhere. and it is obvious that marine birds could arrive at these islands much more easily and frequently than land-birds. But a hooked seed might be carried to an island by other means. yet we can see that their eggs or larvae. Clarke has lately maintained that this island. and will consequently have been but little liable to modification. So that the islands of Bermuda and Madeira have been stocked from the neighbouring continents with birds. de .M. each kind will have been kept by the others to its proper place and habits. and have become mutually co-adapted. now trees. of which one alone is peculiar. and the absence of certain whole groups of animals and plants. but this explanation is not a little doubtful. Although New Zealand is here spoken of as an oceanic island. thus in the Galapagos Islands reptiles. take. and in New Zealand gigantic wingless birds. and their places are occupied by other classes. and is not separated from Australia by a profoundly deep sea.

Candolle has shown, generally have, whatever the cause may be, confined ranges. Hence trees would be little likely to reach distant oceanic islands; and an herbaceous plant, which had no chance of successfully competing with the many fully developed trees growing on a continent, might, when established on an island, gain an advantage over other herbaceous plants by growing taller and taller and overtopping them. In this case, natural selection would tend to add to the stature of the plant, to whatever order it belonged, and thus first convert it into a bush and then into a tree. ABSENCE OF BATRACHIANS AND TERRESTRIAL MAMMALS ON OCEANIC ISLANDS. With respect to the absence of whole orders of animals on oceanic islands, Bory St. Vincent long ago remarked that Batrachians (frogs, toads, newts) are never found on any of the many islands with which the great oceans are studded. I have taken pains to verify this assertion, and have found it true, with the exception of New Zealand, New Caledonia, the Andaman Islands, and perhaps the Solomon Islands and the Seychelles. But I have already remarked that it is doubtful whether New Zealand and New Caledonia ought to be classed as oceanic islands; and this is still more doubtful with respect to the Andaman and Solomon groups and the Seychelles. This general absence of frogs, toads and newts on so many true oceanic islands cannot be accounted for by their physical conditions; indeed it seems that islands are peculiarly fitted for these animals; for frogs have been introduced into Madeira, the Azores, and Mauritius, and have multiplied so as to become a nuisance. But as these animals and their spawn are immediately killed (with the exception, as far as known, of one Indian species) by sea-water, there would be great difficulty in their transportal across the sea, and therefore we can see why they do not exist on strictly oceanic islands. But why, on the theory of creation, they should not have been created there, it would be very difficult to explain. Mammals offer another and similar case. I have carefully searched the oldest voyages, and have not found a single instance, free from doubt, of a terrestrial mammal (excluding domesticated animals kept by the natives) inhabiting an island situated above 300 miles from a continent or great continental island; and many islands situated at a much less distance are equally barren. The Falkland Islands, which are inhabited by a wolf-like fox, come nearest to an exception; but this group cannot be considered as oceanic, as it lies on a bank in connection with the mainland at a distance of about 280 miles; moreover, icebergs formerly brought boulders to its western shores, and they may have formerly transported foxes, as now frequently happens in the arctic regions. Yet it cannot be said that small islands will not support at least small mammals, for they occur in many parts of the world on very small islands, when lying close to a continent; and hardly an island can be named on which our smaller quadrupeds have not become naturalised and greatly multiplied. It cannot be said, on the ordinary view of creation, that there has not been time for the creation of mammals; many volcanic islands are sufficiently ancient, as shown by the stupendous degradation which they have suffered, and by their tertiary strata: there has also been time for the production of endemic species belonging to other classes; and on continents it is known that new species of mammals appear and disappear at a quicker rate than other and lower animals. Although terrestrial mammals do not occur on oceanic islands, aerial mammals do occur on almost every island. New Zealand possesses two bats found nowhere else in the world: Norfolk Island, the Viti Archipelago, the Bonin Islands, the Caroline and Marianne Archipelagoes, and Mauritius, all possess their peculiar bats. Why, it may be asked, has

the supposed creative force produced bats and no other mammals on remote islands? On my view this question can easily be answered; for no terrestrial mammal can be transported across a wide space of sea, but bats can fly across. Bats have been seen wandering by day far over the Atlantic Ocean; and two North American species, either regularly or occasionally, visit Bermuda, at the distance of 600 miles from the mainland. I hear from Mr. Tomes, who has specially studied this family, that many species have enormous ranges, and are found on continents and on far distant islands. Hence, we have only to suppose that such wandering species have been modified in their new homes in relation to their new position, and we can understand the presence of endemic bats on oceanic islands, with the absence of all other terrestrial mammals. Another interesting relation exists, namely, between the depth of the sea separating islands from each other, or from the nearest continent, and the degree of affinity of their mammalian inhabitants. Mr. Windsor Earl has made some striking observations on this head, since greatly extended by Mr. Wallace's admirable researches, in regard to the great Malay Archipelago, which is traversed near Celebes by a space of deep ocean, and this separates two widely distinct mammalian faunas. On either side, the islands stand on a moderately shallow submarine bank, and these islands are inhabited by the same or by closely allied quadrupeds. I have not as yet had time to follow up this subject in all quarters of the world; but as far as I have gone, the relation holds good. For instance, Britain is separated by a shallow channel from Europe, and the mammals are the same on both sides; and so it is with all the islands near the shores of Australia. The West Indian Islands, on the other hand, stand on a deeply submerged bank, nearly one thousand fathoms in depth, and here we find American forms, but the species and even the genera are quite distinct. As the amount of modification which animals of all kinds undergo partly depends on the lapse of time, and as the islands which are separated from each other, or from the mainland, by shallow channels, are more likely to have been continuously united within a recent period than the islands separated by deeper channels, we can understand how it is that a relation exists between the depth of the sea separating two mammalian faunas, and the degree of their affinity, a relation which is quite inexplicable on the theory of independent acts of creation. The foregoing statements in regard to the inhabitants of oceanic islands, namely, the fewness of the species, with a large proportion consisting of endemic forms--the members of certain groups, but not those of other groups in the same class, having been modified--the absence of certain whole orders, as of batrachians and of terrestrial mammals, notwithstanding the presence of aerial bats, the singular proportions of certain orders of plants, herbaceous forms having been developed into trees, etc., seem to me to accord better with the belief in the efficiency of occasional means of transport, carried on during a long course of time, than with the belief in the former connection of all oceanic islands with the nearest continent; for on this latter view it is probable that the various classes would have immigrated more uniformly, and from the species having entered in a body, their mutual relations would not have been much disturbed, and consequently, they would either have not been modified, or all the species in a more equable manner. I do not deny that there are many and serious difficulties in understanding how many of the inhabitants of the more remote islands, whether still retaining the same specific form or subsequently modified, have reached their present homes. But the probability of other islands

having once existed as halting-places, of which not a wreck now remains, must not be overlooked. I will specify one difficult case. Almost all oceanic islands, even the most isolated and smallest, are inhabited by land-shells, generally by endemic species, but sometimes by species found elsewhere striking instances of which have been given by Dr. A.A. Gould in relation to the Pacific. Now it is notorious that land-shells are easily killed by sea-water; their eggs, at least such as I have tried, sink in it and are killed. Yet there must be some unknown, but occasionally efficient means for their transportal. Would the just-hatched young sometimes adhere to the feet of birds roosting on the ground and thus get transported? It occurred to me that land-shells, when hybernating and having a membranous diaphragm over the mouth of the shell, might be floated in chinks of drifted timber across moderately wide arms of the sea. And I find that several species in this state withstand uninjured an immersion in sea-water during seven days. One shell, the Helix pomatia, after having been thus treated, and again hybernating, was put into sea-water for twenty days and perfectly recovered. During this length of time the shell might have been carried by a marine country of average swiftness to a distance of 660 geographical miles. As this Helix has a thick calcareous operculum I removed it, and when it had formed a new membranous one, I again immersed it for fourteen days in sea-water, and again it recovered and crawled away. Baron Aucapitaine has since tried similar experiments. He placed 100 land-shells, belonging to ten species, in a box pierced with holes, and immersed it for a fortnight in the sea. Out of the hundred shells twenty-seven recovered. The presence of an operculum seems to have been of importance, as out of twelve specimens of Cyclostoma elegans, which is thus furnished, eleven revived. It is remarkable, seeing how well the Helix pomatia resisted with me the salt-water, that not one of fifty-four specimens belonging to four other species of Helix tried by Aucapitaine recovered. It is, however, not at all probable that land-shells have often been thus transported; the feet of birds offer a more probable method. ON THE RELATIONS OF THE INHABITANTS OF ISLANDS TO THOSE OF THE NEAREST MAINLAND. The most striking and important fact for us is the affinity of the species which inhabit islands to those of the nearest mainland, without being actually the same. Numerous instances could be given. The Galapagos Archipelago, situated under the equator, lies at a distance of between 500 and 600 miles from the shores of South America. Here almost every product of the land and of the water bears the unmistakable stamp of the American continent. There are twenty-six land birds. Of these twenty-one, or perhaps twenty-three, are ranked as distinct species, and would commonly be assumed to have been here created; yet the close affinity of most of these birds to American species is manifest in every character in their habits, gestures, and tones of voice. So it is with the other animals, and with a large proportion of the plants, as shown by Dr. Hooker in his admirable Flora of this archipelago. The naturalist, looking at the inhabitants of these volcanic islands in the Pacific, distant several hundred miles from the continent, feels that he is standing on American land. Why should this be so? Why should the species which are supposed to have been created in the Galapagos Archipelago, and nowhere else, bear so plainly the stamp of affinity to those created in America? There is nothing in the conditions of life, in the geological nature of the islands, in their height or climate, or in the proportions in which the several classes are associated together, which closely resembles the conditions of the South American coast. In fact, there is a considerable dissimilarity in all these respects. On

the other hand, there is a considerable degree of resemblance in the volcanic nature of the soil, in the climate, height, and size of the islands, between the Galapagos and Cape Verde Archipelagos: but what an entire and absolute difference in their inhabitants! The inhabitants of the Cape Verde Islands are related to those of Africa, like those of the Galapagos to America. Facts, such as these, admit of no sort of explanation on the ordinary view of independent creation; whereas, on the view here maintained, it is obvious that the Galapagos Islands would be likely to receive colonists from America, whether by occasional means of transport or (though I do not believe in this doctrine) by formerly continuous land, and the Cape Verde Islands from Africa; such colonists would be liable to modification--the principle of inheritance still betraying their original birthplace. Many analogous facts could be given: indeed it is an almost universal rule that the endemic productions of islands are related to those of the nearest continent, or of the nearest large island. The exceptions are few, and most of them can be explained. Thus, although Kerguelen Land stands nearer to Africa than to America, the plants are related, and that very closely, as we know from Dr. Hooker's account, to those of America: but on the view that this island has been mainly stocked by seeds brought with earth and stones on icebergs, drifted by the prevailing currents, this anomaly disappears. New Zealand in its endemic plants is much more closely related to Australia, the nearest mainland, than to any other region: and this is what might have been expected; but it is also plainly related to South America, which, although the next nearest continent, is so enormously remote, that the fact becomes an anomaly. But this difficulty partially disappears on the view that New Zealand, South America, and the other southern lands, have been stocked in part from a nearly intermediate though distant point, namely, from the antarctic islands, when they were clothed with vegetation, during a warmer tertiary period, before the commencement of the last Glacial period. The affinity, which, though feeble, I am assured by Dr. Hooker is real, between the flora of the south-western corner of Australia and of the Cape of Good Hope, is a far more remarkable case; but this affinity is confined to the plants, and will, no doubt, some day be explained. The same law which has determined the relationship between the inhabitants of islands and the nearest mainland, is sometimes displayed on a small scale, but in a most interesting manner, within the limits of the same archipelago. Thus each separate island of the Galapagos Archipelago is tenanted, and the fact is a marvellous one, by many distinct species; but these species are related to each other in a very much closer manner than to the inhabitants of the American continent, or of any other quarter of the world. This is what might have been expected, for islands situated so near to each other would almost necessarily receive immigrants from the same original source, and from each other. But how is it that many of the immigrants have been differently modified, though only in a small degree, in islands situated within sight of each other, having the same geological nature, the same height, climate, etc? This long appeared to me a great difficulty: but it arises in chief part from the deeply-seated error of considering the physical conditions of a country as the most important; whereas it cannot be disputed that the nature of the other species with which each has to compete, is at least as important, and generally a far more important element of success. Now if we look to the species which inhabit the Galapagos Archipelago, and are likewise found in other parts of the world, we find that they differ considerably in the several islands. This difference might indeed have been expected if the

islands have been stocked by occasional means of transport--a seed, for instance, of one plant having been brought to one island, and that of another plant to another island, though all proceeding from the same general source. Hence, when in former times an immigrant first settled on one of the islands, or when it subsequently spread from one to another, it would undoubtedly be exposed to different conditions in the different islands, for it would have to compete with a different set of organisms; a plant, for instance, would find the ground best-fitted for it occupied by somewhat different species in the different islands, and would be exposed to the attacks of somewhat different enemies. If, then, it varied, natural selection would probably favour different varieties in the different islands. Some species, however, might spread and yet retain the same character throughout the group, just as we see some species spreading widely throughout a continent and remaining the same. The really surprising fact in this case of the Galapagos Archipelago, and in a lesser degree in some analogous cases, is that each new species after being formed in any one island, did not spread quickly to the other islands. But the islands, though in sight of each other, are separated by deep arms of the sea, in most cases wider than the British Channel, and there is no reason to suppose that they have at any former period been continuously united. The currents of the sea are rapid and deep between the islands, and gales of wind are extraordinarily rare; so that the islands are far more effectually separated from each other than they appear on a map. Nevertheless, some of the species, both of those found in other parts of the world and of those confined to the archipelago, are common to the several islands; and we may infer from the present manner of distribution that they have spread from one island to the others. But we often take, I think, an erroneous view of the probability of closely allied species invading each other's territory, when put into free intercommunication. Undoubtedly, if one species has any advantage over another, it will in a very brief time wholly or in part supplant it; but if both are equally well fitted for their own places, both will probably hold their separate places for almost any length of time. Being familiar with the fact that many species, naturalised through man's agency, have spread with astonishing rapidity over wide areas, we are apt to infer that most species would thus spread; but we should remember that the species which become naturalised in new countries are not generally closely allied to the aboriginal inhabitants, but are very distinct forms, belonging in a large proportion of cases, as shown by Alph. de Candolle, to distinct genera. In the Galapagos Archipelago, many even of the birds, though so well adapted for flying from island to island, differ on the different islands; thus there are three closely allied species of mocking-thrush, each confined to its own island. Now let us suppose the mocking-thrush of Chatham Island to be blown to Charles Island, which has its own mocking-thrush; why should it succeed in establishing itself there? We may safely infer that Charles Island is well stocked with its own species, for annually more eggs are laid and young birds hatched than can possibly be reared; and we may infer that the mocking-thrush peculiar to Charles Island is at least as well fitted for its home as is the species peculiar to Chatham Island. Sir C. Lyell and Mr. Wollaston have communicated to me a remarkable fact bearing on this subject; namely, that Madeira and the adjoining islet of Porto Santo possess many distinct but representative species of land-shells, some of which live in crevices of stone; and although large quantities of stone are annually transported from Porto Santo to Madeira, yet this latter island has not become colonised by the Porto Santo species: nevertheless, both islands have been colonised by some European land-shells, which no doubt had some advantage over the indigenous species. From these

considerations I think we need not greatly marvel at the endemic species which inhabit the several islands of the Galapagos Archipelago not having all spread from island to island. On the same continent, also, pre-occupation has probably played an important part in checking the commingling of the species which inhabit different districts with nearly the same physical conditions. Thus, the south-east and south-west corners of Australia have nearly the same physical conditions, and are united by continuous land, yet they are inhabited by a vast number of distinct mammals, birds, and plants; so it is, according to Mr. Bates, with the butterflies and other animals inhabiting the great, open, and continuous valley of the Amazons. The same principle which governs the general character of the inhabitants of oceanic islands, namely, the relation to the source whence colonists could have been most easily derived, together with their subsequent modification, is of the widest application throughout nature. We see this on every mountain-summit, in every lake and marsh. For Alpine species, excepting in as far as the same species have become widely spread during the Glacial epoch, are related to those of the surrounding lowlands; thus we have in South America, Alpine humming-birds, Alpine rodents, Alpine plants, etc., all strictly belonging to American forms; and it is obvious that a mountain, as it became slowly upheaved, would be colonised from the surrounding lowlands. So it is with the inhabitants of lakes and marshes, excepting in so far as great facility of transport has allowed the same forms to prevail throughout large portions of the world. We see the same principle in the character of most of the blind animals inhabiting the caves of America and of Europe. Other analogous facts could be given. It will, I believe, be found universally true, that wherever in two regions, let them be ever so distant, many closely allied or representative species occur, there will likewise be found some identical species; and wherever many closely-allied species occur, there will be found many forms which some naturalists rank as distinct species, and others as mere varieties; these doubtful forms showing us the steps in the process of modification. The relation between the power and extent of migration in certain species, either at the present or at some former period, and the existence at remote points of the world of closely allied species, is shown in another and more general way. Mr. Gould remarked to me long ago, that in those genera of birds which range over the world, many of the species have very wide ranges. I can hardly doubt that this rule is generally true, though difficult of proof. Among mammals, we see it strikingly displayed in Bats, and in a lesser degree in the Felidae and Canidae. We see the same rule in the distribution of butterflies and beetles. So it is with most of the inhabitants of fresh water, for many of the genera in the most distinct classes range over the world, and many of the species have enormous ranges. It is not meant that all, but that some of the species have very wide ranges in the genera which range very widely. Nor is it meant that the species in such genera have, on an average, a very wide range; for this will largely depend on how far the process of modification has gone; for instance, two varieties of the same species inhabit America and Europe, and thus the species has an immense range; but, if variation were to be carried a little further, the two varieties would be ranked as distinct species, and their range would be greatly reduced. Still less is it meant, that species which have the capacity of crossing barriers and ranging widely, as in the case of certain powerfully-winged birds, will necessarily range widely; for we should never forget that to range widely implies not only the power of crossing barriers, but the more important power of being

victorious in distant lands in the struggle for life with foreign associates. But according to the view that all the species of a genus, though distributed to the most remote points of the world, are descended from a single progenitor, we ought to find, and I believe as a general rule we do find, that some at least of the species range very widely. We should bear in mind that many genera in all classes are of ancient origin, and the species in this case will have had ample time for dispersal and subsequent modification. There is also reason to believe, from geological evidence, that within each great class the lower organisms change at a slower rate than the higher; consequently they will have had a better chance of ranging widely and of still retaining the same specific character. This fact, together with that of the seeds and eggs of most lowly organised forms being very minute and better fitted for distant transportal, probably accounts for a law which has long been observed, and which has lately been discussed by Alph. de Candolle in regard to plants, namely, that the lower any group of organisms stands the more widely it ranges. The relations just discussed--namely, lower organisms ranging more widely than the higher--some of the species of widely-ranging genera themselves ranging widely--such facts, as alpine, lacustrine, and marsh productions being generally related to those which live on the surrounding low lands and dry lands--the striking relationship between the inhabitants of islands and those of the nearest mainland--the still closer relationship of the distinct inhabitants of the islands of the same archipelago--are inexplicable on the ordinary view of the independent creation of each species, but are explicable if we admit colonisation from the nearest or readiest source, together with the subsequent adaptation of the colonists to their new homes. SUMMARY OF THE LAST AND PRESENT CHAPTERS. In these chapters I have endeavoured to show that if we make due allowance for our ignorance of the full effects of changes of climate and of the level of the land, which have certainly occurred within the recent period, and of other changes which have probably occurred--if we remember how ignorant we are with respect to the many curious means of occasional transport--if we bear in mind, and this is a very important consideration, how often a species may have ranged continuously over a wide area, and then have become extinct in the intermediate tracts--the difficulty is not insuperable in believing that all the individuals of the same species, wherever found, are descended from common parents. And we are led to this conclusion, which has been arrived at by many naturalists under the designation of single centres of creation, by various general considerations, more especially from the importance of barriers of all kinds, and from the analogical distribution of subgenera, genera, and families. With respect to distinct species belonging to the same genus, which on our theory have spread from one parent-source; if we make the same allowances as before for our ignorance, and remember that some forms of life have changed very slowly, enormous periods of time having been thus granted for their migration, the difficulties are far from insuperable; though in this case, as in that of the individuals of the same species, they are often great. As exemplifying the effects of climatical changes on distribution, I have attempted to show how important a part the last Glacial period has played, which affected even the equatorial regions, and which, during

the alternations of the cold in the north and the south, allowed the productions of opposite hemispheres to mingle, and left some of them stranded on the mountain-summits in all parts of the world. As showing how diversified are the means of occasional transport, I have discussed at some little length the means of dispersal of fresh-water productions. If the difficulties be not insuperable in admitting that in the long course of time all the individuals of the same species, and likewise of the several species belonging to the same genus, have proceeded from some one source; then all the grand leading facts of geographical distribution are explicable on the theory of migration, together with subsequent modification and the multiplication of new forms. We can thus understand the high importance of barriers, whether of land or water, in not only separating but in apparently forming the several zoological and botanical provinces. We can thus understand the concentration of related species within the same areas; and how it is that under different latitudes, for instance, in South America, the inhabitants of the plains and mountains, of the forests, marshes, and deserts, are linked together in so mysterious a manner, and are likewise linked to the extinct beings which formerly inhabited the same continent. Bearing in mind that the mutual relation of organism to organism is of the highest importance, we can see why two areas, having nearly the same physical conditions, should often be inhabited by very different forms of life; for according to the length of time which has elapsed since the colonists entered one of the regions, or both; according to the nature of the communication which allowed certain forms and not others to enter, either in greater or lesser numbers; according or not as those which entered happened to come into more or less direct competition with each other and with the aborigines; and according as the immigrants were capable of varying more or less rapidly, there would ensue in the to or more regions, independently of their physical conditions, infinitely diversified conditions of life; there would be an almost endless amount of organic action and reaction, and we should find some groups of beings greatly, and some only slightly modified; some developed in great force, some existing in scanty numbers--and this we do find in the several great geographical provinces of the world. On these same principles we can understand, as I have endeavoured to show, why oceanic islands should have few inhabitants, but that of these, a large proportion should be endemic or peculiar; and why, in relation to the means of migration, one group of beings should have all its species peculiar, and another group, even within the same class, should have all its species the same with those in an adjoining quarter of the world. We can see why whole groups of organisms, as batrachians and terrestrial mammals, should be absent from oceanic islands, whilst the most isolated islands should possess their own peculiar species of aerial mammals or bats. We can see why, in islands, there should be some relation between the presence of mammals, in a more or less modified condition, and the depth of the sea between such islands and the mainland. We can clearly see why all the inhabitants of an archipelago, though specifically distinct on the several islets, should be closely related to each other, and should likewise be related, but less closely, to those of the nearest continent, or other source whence immigrants might have been derived. We can see why, if there exist very closely allied or representative species in two areas, however distant from each other, some identical species will almost always there be found. As the late Edward Forbes often insisted, there is a striking parallelism in the laws of life throughout time and space; the laws governing the succession of forms in past times being nearly the same

with those governing at the present time the differences in different areas. We see this in many facts. The endurance of each species and group of species is continuous in time; for the apparent exceptions to the rule are so few that they may fairly be attributed to our not having as yet discovered in an intermediate deposit certain forms which are absent in it, but which occur above and below: so in space, it certainly is the general rule that the area inhabited by a single species, or by a group of species, is continuous, and the exceptions, which are not rare, may, as I have attempted to show, be accounted for by former migrations under different circumstances, or through occasional means of transport, or by the species having become extinct in the intermediate tracts. Both in time and space species and groups of species have their points of maximum development. Groups of species, living d